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Home range behavior of Roosevelt elk in Strathcona Park Sovka, David G. 1993

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We accep this thesis as conforminge requ' ed standardHOME RANGE BEHAVIOR OF ROOSEVELT ELK IN STRATHCONA PARKbyDAVID GRANT SOVKAB.Sc., The University of Calgary, 1990A THESIS SUBMITTED IN PARTIAL FULFILLMENT OFTHE REQUIREMENTS FOR THE DEGREE OFMASTER OF SCIENCEinTHE FACULTY OF GRADUATE STUDIES(Department of Animal Science)THE UNIVERSITY OF BRITISH COLUMBIAApril 1993© David Grant Sovka, 1993In presenting this thesis in partial fulfilment of the requirements for an advanceddegree at the University of British Columbia, I agree that the Library shall make itfreely available for reference and study. I further agree that permission for extensivecopying of this thesis for scholarly purposes may be granted by the head of mydepartment or by his or her representatives. It is understood that copying orpublication of this thesis for financial gain shall not be allowed without my writtenpermission.Department of /40/144/-- SC:-/ '5771/ .The University of British ColumbiaVancouver, CanadaDate .4/E7/4- DE-6 (2/88)-II-AbstractThis thesis investigates the population, group composition, and home rangesizes of 5 groups of Roosevelt elk (Cervus elaphus roosev -It) in 4 regions ofStrathcona Provincial Park on Vancouver Island, British Columbia. Ten elk weresuccessfully radio-collared and re-located for 12-21 months, beginning in March1991, and ending in January 1993. Eight of the 10 study animals were non-migratory. Sizes of cumulative and seasonal ranges, and core use areas weredetermined for each study animal using the harmonic mean estimator of ProgramHome Range.Three seasonal models (summer, mild winter, and severe winter) of habitatsuitability were tested using a non-migratory elk group. A Geographic InformationSystem (GIS) was used to create a digital map of the study area. The models wereapplied to the map to generate habitat suitability values for each of the polygonscomprising the study area.This study provides additional support for a correlation between habitatsuitability as predicted by the model and elk habitat selection. Elk were notrandomly selecting habitats in the study area with respect to modeled habitatsuitability values. Habitat suitability of elk cumulative range was higher thanportions of the study area unused by elk. Core use areas had higher habitatsuitability than cumulative range. No one model best predicted the habitatsuitability of the home range selected by the non-migratory study animals.Use of the model as a predictor of suitable elk habitat in areas currentlyunused by elk is validated, but not warranted given the time and money required tocollect the information necessary to use the model.Table of ContentsAbstract^ iiTable of Contents^ iiiList of Tables vList of Figures^ viAcknowledgement viiGENERAL INTRODUCTION^ 1Classification and Distribution^ 1Current Elk Population Status on Vancouver Island^ 2Roosevelt Elk Management on Vancouver Island 3Roosevelt Elk Management in Strathcona Park 4CHAPTER 1. POPULATION AND HOME RANGE CHARACTERISTICS FORROOSEVELT ELK IN STRATHCONA PARK^ 7Introduction^ 7Population and Home Range Concepts^ 7Measuring Population and Home Range 9Study Area^ 10Study Area Boundaries^ 10Elk River Valley^ 13Thelwood Valley 14Heber River-Camel Creek Valleys^ 15Ucona River Basin^ 16Methods^ 17Trapping and Radio-Collaring Animals^ 17Population and Group Composition Data 18Animal Locations^ 18Home Range Delineation^ 22Animal/Season Selection Criteria^ 27Home Range Analysis 30Core Use Areas^ 31Statistical Analyses 33Results^ 35Mortalities^ 35Animal Locations^ 35Group Composition and Population^ 35Home Range Characteristics and Movements^ 38Elk River Valley^ 38Thelwood Valley 46-iv-Heber River-Camel Creek Valleys^ 48Ucona River Basin^ 49Discussion^ 50Mortalities^ 50Home Range Characteristics and Movements^ 51Elk River Valley^ 52Thelwood Valley 53Heber River-Camel Creek Valleys^ 54Ucona River Basin^ 55Migration and Non-migration as Movement Strategies^56Home Range Estimation 62Summary^ 65CHAPTER 2. TESTING SEASONAL MODELS OF ROOSEVELT ELK HABITATSUITABILITY^ 67Introduction 67Habitat Suitability Index (HSI) Model Development and Testing^68GIS Applications in Wildlife Habitat Modelling^ 70Study Background - Brunt's (1991) Thesis 72Study Area^ 75Study Area^ 75Historical Overview^ 76Methods^ 77Study Area Map Development^ 77Model Development and Application 78Forage^ 79Cover 82Interspersion 83Aspect/Elevation^ 85Habitat Suitability Calculations^ 87Home Range Suitability Calculations 88Analyses of Modeled Habitat Suitability^ 89A) Random Habitat Selection Test 89B) Habitat Suitability Within Study Area^ 90C) Comparisons Among Seasonal Model Prediction^92Results^ 93Habitat Suitability Model Validation Tests^ 93A) Random Habitat Selection Test 93B) Habitat Suitability Within Study Area 93C) Comparisons Among Seasonal Model Output^97Discussion^ 97Conclusions 105LITERATURE CITED^ 109-v-List of TablesTable 1.^Dates used to delineate seasonal home ranges for each study animal.28Table 2.^Home ranges analyzed using the harmonic mean contour method. 29Table 3.^Core use area delineation methods and core area and home rangesizes.^ 34Table 4.^Classification counts for summer and winter seasons.^36, 37Table 5.^Sizes (km2) of seasonal home range and core use areas for 2 non-migratory elk from the Heber River-Camel Creek valleys.^44Table 6.^Sizes (km 2) of cumulative (year-round home range using 100% ofutilization distribution) and core use areas for all study animals.^45Table 7.^Habitat types used by the Elk River valley group during summer 1992(May 21-August 30).^ 47Table 8.^Forage modifier values. 65Table 9.^Cover and potential forage suitability values by understory type.^66Table 10.^Interspersion modifier values.^ 69Table 11. Aspect/elevation modifier values used in the mild and severe wintermodels.^ 71Table 12.^Frequencies of expected (given random habitat selection) andobserved habitat suitability class values for the summer model.^80Table 13.Table 14.Habitat suitability values for total study area, unused portion of thetotal study area, cumulative range, and core use area for summer, mildwinter, and severe winter models.^ 81Mean habitat suitability values for animal locations in the core usearea, and random locations in the unused portion, of the Elk Rivervalley study area. 83Table 15.^Mean habitat suitability values of summer, winter, and cumulativeanimal locations estimated by each of the 3 seasonal models.^84Table 16.^Results of all possible two-tailed paired t-test comparisons amongsummer, winter, and cumulative animal location datasets using the 3habitat suitability models.^ 85-vi-List of FiguresFigure 1.^The 4 study areas in Strathcona Provincial Park on Vancouver Island,British Columbia.^ 11Figure 2.^Cumulative range and core use area for elk it's 120, 179, 218, and259, in the Elk River valley.^ 39Figure 3.^Cumulative range and core use areas for elk #580 in the Thelwoodvalley.^ 40Figure 4.^Summer and winter (1991-92) seasonal ranges and core use areas forelk #499 in the Heber River-Camel Creek valleys.^ 41Figure 5.^Summer and winter (1991-92) seasonal ranges and core use areas forelk #688 in the Heber River-Camel Creek valleys.^ 42Figure 6.^Cumulative range and core use areas of elk it's 159, 300, and 339 inthe Ucona River basin.^ 43Figure 7.^Total study area, unused portion of the study area, cumulative range,and core use area for elk it's 120, 179, 218, and 259 in the Elk Rivervalley, as delineated for seasonal habitat suitability model testing. 94-vii-AcknowledgementsThis research was supported financially by the Ministry of Environment,Lands and Parks; the Canadian Wildlife Service; the Vancouver Island Habitat IslandEnhancement Fund; and David Shackleton. I would like to thank Rik Simmons ofBC Parks for administrative support, and also Ron Quitter and Joanne McLeod ofthe Strathcona Zone Office. Kim Brunt provided assistance throughout my study,including darting and collaring the study animals, data analysis, and helpfulsuggestions in the field. I would like to thank David Shackleton, Fred Bunnell, andMike Pitt of the University of British Columbia, and Rik Simmons of BC Parks forparticipating on my graduate committee. Strathcona Park Ranger Rob Robertsonassisted in data collection several times. Mike Scott of VideoWave Productions,Inc. (Courtenay, BC) accompanied me on several backcountry trips to search forelk, and videoed a short documentary of the study. Thanks to Jerry Maedel andPeter Murtha of the University of British Columbia's FIRMS/GIS Laboratory forallowing me to use the equipment.I would like to thank my beloved for her support and encouragementthroughout this study. She tolerated my numerous absences over the past twoyears; missed me; loved me; and even accompanied me several times in the fieldup steep, dangerous logging roads in our 1972 Mercury, and all the while 6 monthspregnant with our good and perfect gift. I would like to thank him too, for thegreat joy that he is. Above all, though, thanks to Him who's creation I studiedwith wonder and fear; who protected and cared for me in spite of so much.GENERAL INTRODUCTIONClassification and DistributionCurrently, 2 subspecies of elk occur in British Columbia: Roosevelt elk(Cervus elaphus roosevelti) and Rocky Mountain elk (C. e. nelsoni). RockyMountain elk are the more abundant and widespread, occurring in greatest numbersin the Rocky Mountains and the Rocky Mountain trench of southeastern BritishColumbia. Roosevelt elk occur in Canada only on Vancouver Island, which is thenorthernmost limit of their present-day natural distribution; in valleys at the head ofseveral mainland inlets adjacent to northern Vancouver Island; and on the SecheltPeninsula as a result of several transplants in the late 1980's (Brunt 1990).Historically, elk were more widely distributed in coastal British Columbia thanthey are at present (Cowan and Guiguet 1965, Bryant and Maser 1982). Localextinctions were caused by human settlement and excessive hunting, particularlyon southern Vancouver Island. The present reduced numbers and limiteddistribution of elk on the Island result from a number of factors, including habitatquality, historical hunting and predation pressures, geographical barriers tocolonization, transplant efforts, and a certain amount of chance patterns ofdispersal (Brunt 1990). Roosevelt elk population numbers are considered to be farbelow their maximum sustainable level (Nyberg and Janz 1990).A discussion of the classification and historical distribution of elk in NorthAmerica can be found in Bryant and Maser (1982), together with a thorough-2-physical description of the Roosevelt elk subspecies. Geist (1982) has reviewedelk social behavior.Current Elk Population Status on Vancouver IslandAn accurate census of ungulate populations in the coastal forests found onVancouver Island is virtually impossible because cover is so dense. Recentestimates of elk populations prepared for the regional wildlife and habitat plans ofthe British Columbia Ministry of Environment, Lands, and Parks (MOELP) indicatethat Roosevelt elk number between 2170 and 2770 (Nyberg et al. 1990).The Wildlife Species Evaluation List describes the status and prognosis of thewildlife species in British Columbia. In 1991, the MOELP placed Roosevelt elk onthe blue list (sensitive/vulnerable species). The reasons for blue-listing speciesinclude the following: i) a significant current or predicted downward trends inpopulation numbers or density; and ii) significant current or predicted downwardtrends in habitat suitability that would further reduce the species' existingdistribution.Elk populations on Vancouver Island have declined over the past fifteenyears. Climate, habitat changes, and hunting have most affected elk on VancouverIsland. Effects of the severe 1968-69 winter were most pronounced on southernVancouver Island, where because of past logging history, few old-growth winterranges remained, while there were many large areas dominated by young,regenerating forests. Elk populations declined during this period because of deepsnow accumulations, resulting from the loss of canopy cover (Nyberg et al. 1990).-3-Over the next decade, elk numbers gradually recovered in response to a series ofmild winters, but increased illegal hunting and unregulated native hunting haveslowed the elk recovery (MOELP 1989a).Nyberg et al. (1990) describe the present status of Roosevelt elk on northernVancouver Island as "more encouraging," where most herds are stable or increasingbecause of the recent mild winters, a favorable habitat mosaic in many watersheds,and relatively low levels of illegal hunting. Wolf predation affects some herds inwatersheds where deer numbers are low and wolves are switching to elk as analternative prey (Janz and Becker 1986). In such areas, the effects of wolfpredation on elk populations are likely to increase during more severe winters as elkcongregate on small winter ranges and become less mobile and less healthy.Roosevelt Elk Management on Vancouver IslandManagement of the provincial wildlife resource is the responsibility of theWildlife Branch, Ministry of the Environment (MOELP). The mission of the wildlifeprogram is "...to manage the province's wildlife resources for the benefit andenjoyment of British Columbians, by maintaining an optimal balance betweenecological, cultural, economic, and recreational needs" (MOELP 1989b). A majorgoal is to maintain and enhance wildlife and their habitats to ensure an abundant,diverse, and self-sustaining wildlife resource throughout British Columbia.The present management objective for Roosevelt elk on Vancouver Island isto increase elk numbers to approximately 3000-3800 by 1996, which shouldgenerate 2350-3200 hunter-days of recreation and provide increased viewing-4-opportunities (Nyberg et al. 1990). Management activities required to meetpopulation objectives include the curtailment of illegal and unregulated hunting;habitat management focusing on protection of important old-growth winter rangesand special habitats such as wetlands and vegetated slides; provision of forage andcover in regenerating forests; and manipulation of young stands to provide snowinterception cover.At present, the status of Roosevelt elk in British Columbia is listed as "stableto increasing (2,500)" (BC Wildlife Branch 1991). However, significant losses totraditional elk habitat have occurred, even within supposedly protected areas suchas Strathcona Provincial Park on Vancouver Island (Strathcona Park ElkManagement Plan, 1990).Roosevelt Elk Management in Strathcona ParkWithin the MOELP, the Parks branch (BC Parks) is responsible for thedesignation, management and conservation of a land and water-based system ofparks, recreation areas and ecological reserves, containing the best representativeelements of British Columbia's natural and cultural heritage for the inspiration andrecreational use of British Columbians and their visitors (BC Parks' Mandate, 1990).As such, in 1990 BC Parks developed a management plan for Strathcona Parkwhich set out objectives outlining the government's stance on the future of elk inthe Park.The Strathcona Park Elk Management Plan states that "...maintaining ahealthy viable population of elk in the park is the primary goal of the park-5-management plan as well as satisfying park system objectives." One objective ofthe park system is to represent the biodiversity of the province. BC Parksdetermined that Roosevelt elk, as a keystone species of the province of BritishColumbia, "...must be maintained not only in their present but also their historicdistribution and abundance," (Strathcona Park Elk Management Plan, 1990). Atthe same time, however, it was realized that "...the present state of knowledge islimited and does [not) include a complete understanding of existing habitat,utilization, distribution and abundance by elk or allow an assessment of other areasof the park's significance to elk."In light of the Ministry's goal "...to provide an environment in which thecontinued presence of elk is assured in Strathcona Park in perpetuity," (StrathconaPark Elk Management Plan, 1990), a set of objectives was drafted, which includethe following goals: protecting and enhancing elk habitat, increasing elk abundanceand distribution, and increasing opportunities for viewing elk.Within the 3 broad goals identified in the Strathcona Park Elk ManagementPlan (1990), broad action steps were defined. For the goal of protecting andenhancing elk habitat, the defined action steps included determining the present elkdistribution and abundance in the Park, and developing "...a park specificunderstanding of elk ecology and capability." For the goal of increasing elkabundance and distribution, the defined action steps included enhancing the levelof knowledge of habitats and habitat utilization in the park, reintroducing extirpated-6-populations to the Park, identifying unused habitat, and designing and implementinga transplant of elk into unused habitat.In the interests of meeting the first and second goals, BC Parks, incooperation with the BC Wildlife Branch, trapped and radio-collared 10 elk indifferent areas within and adjacent to the Strathcona Park boundary. Applicableresults obtained in the present study form Chapters 1 and 2 of this thesis. Mystudy represents the first phase in a multi-level plan addressing the definedobjectives of the Strathcona Park Elk Management Plan (1990).The specific objectives of my study were to collect baseline ecological dataon the population and home ranges of Roosevelt elk of Strathcona Provincial Parkon Vancouver Island; and ii) to validate seasonal habitat suitability modelsdeveloped by Brunt (1991), and which may be a useful park management tool forpredicting high quality elk habitat throughout the Park. The main purpose of mystudy was to provide the baseline ecological and practical information that willallow Park managers to better understand the current elk-habitat relationship inStrathcona Park.-7-CHAPTER 1Population and Home Range Characteristics for Roosevelt Elk in Strathcona ParkINTRODUCTIONPopulation and Home Range ConceptsIndividual elk do not live naturally in isolation, but rather in a particular socialenvironment, and with other elk, as members of a group. "Group" can refer to aprovince-wide group, the population of a management unit, the population of awatershed, or a distinct group within a watershed. For the purpose of this thesis,an elk group is defined as > 1 Roosevelt elk which occupy a similar environment,and inhabiting a contiguous region, such as a watershed.As Taber et al. (1982) noted, such a population has 3 general characteristicswhich may be described mathematically, including those related to production, tomortality, and to movement. Analysis of these characteristics and theirinteractions can provide estimates of life history attributes which may not bemeasured readily, such as life expectancy. Gathering and analyzing population datais fundamental to deriving wildlife-habitat management guidelines and strategies.A concept closely related to the characteristics defining an animal group, andequally important to wildlife-habitat management decisions is that of home range.Burt's (1943) classic conceptual definition of home range is "...that area traversedby the individual in its normal activities of food gathering, mating and caring foryoung." Home range may be defined over daily, seasonal, or lifetime time frames(Harested 1979).-8-The size of an animal's home range is related to its energy requirements, andthe productivity of the habitat it is utilizing (Bunnell 1978). Larger animals, forinstance, are capable of covering a greater area in the course of a day, as well asrequiring more absolute energy than do smaller animals. Other factors affecting orinfluencing home range size include population density, season of year, interspecificinteractions, age, latitude, terrain, and precipitation (Janz et al. 1980b). Habitatquality and terrain are other key factors affecting home range size. Harper (1971)concluded that terrain was the most important factor regulating the home rangesize of Roosevelt elk in Oregon.Population characteristics have been described for Roosevelt elk onVancouver Island (Brunt et al. 1989), and for other North American elk sub-species(Taber et al. 1982, Morgantini 1988, Woods 1991). Seasonal and cumulativehome ranges have been estimated for several migratory and non-migratoryRoosevelt elk groups on Vancouver Island (Janz and Lloyd 1977, Janz et al.1980b, Brunt et al. 1989, Brunt 1991).Important to understanding the relationship between populationcharacteristics and home range selection in terms of size, stability, and seasonalityis the concept of migration. Migration is generally defined as the annual returnmovement of individuals between two geographically separate areas (Woods1991). It follows implicitly that migratory animals establish a degree of philopatryto seasonal ranges and make relatively little use of areas between these ranges.While many species of birds and mammals have been classified as being either-9-migratory or non-migratory, intraspecific variation in seasonal movement patterns iscommon (Baker 1978). Elk are often classified as migratory, despite observationsof co-existing non-migratory groups, and groups utilizing several concurrentmovement strategies (Woods 1991). Morgantini (1988) suggested that thevariation found in elk movement strategies is an example of the species' flexibilityin coping with a varied environment. Woods (1991) concluded that the occurrenceof different space-use patterns in elk populations was an evolutionary stablestrategy (ESS).Measuring Population and Home RangeThere are 2 levels of population structure analysis, including the structurethat can be detected through group composition counts (viz. recognition andenumeration of age and sex classes in a population), and that which is developedthrough the use of mathematical models of the population (Taber et al. 1982). Thefirst level is employed in the present study, where group composition countsconsisted of direct tallies of individual elk in the study area.Twenty years ago Craighead et al. (1973) noted that the techniques of radio-telemetry make possible the delineation of land areas vital to individual animals, todetermining seasonal use of those areas as the animal's seasonal needs are met,and to observing behavior and activity that has spatial and temporal significance inthe life of the animal. The use of radio-telemetry to study space and habitat usepatterns of animals is now considered standard practice in wildlife research andmanagement (Skovlin 1982, White and Garrot 1990, Samuel and Kenow 1992).-10-Radio-triangulation is the most common technique used to determine an animal'slocation (Springer 1979), and was used in the present study to identify animallocations, which were then imported into a home range estimating program.In a survey of recent studies presenting home range data, Boulanger andWhite (1990) commented that the application of different home range estimatorsproduces confusion in the interpretation of home range estimates because some ofthe differences observed between studies are due to the estimators themselves,and not to the behavior of the study animals. All home range estimations in thepresent study were made using the harmonic mean option in Program Home Range,second edition (Ackerman et al. 1990), a method of home range estimation rapidlygaining popularity in wildlife ecology (Boulanger and White 1990, Harris et al.1990, Brunt 1991).In this chapter of the thesis, I describe the composition and home rangecharacteristics of 5 distinct Roosevelt elk groups within or adjacent to the boundaryof Strathcona Provincial Park on Vancouver Island.STUDY AREAStudy Area BoundariesThe study area (Figure 1) includes regions of Strathcona Provincial Park, andareas immediately adjacent to the Park's boundaries. Strathcona Park, BritishColumbia's oldest provincial park, is centrally located on Vancouver Island, andcovers an area of approximately 210,000 ha. Most of the Park is mountainous,VANCOUVER ISLAND0 5.0 10.0 kmTHELWOOD VALLEYHighway 28HEBER RIVER - CAMELCREEK VALLEYSUpper Campbell LakeELK RIVER VALLEYUCONA RIVER BASINSTRATHCONAPROVINCIAL PARKStrathcona Provincial ParkVictoria•Study AreasButtle LakeFigure 1. The 4 study areas in Strathcona Park onVancouver Island, British Columbia.-12-forming a large part of the Vancouver Island Ranges, which are generally composedof a heterogeneous group of pre-Cretaceous sedimentary and volcanic rocks foldedby numerous granite batholiths (Holland 1964). Elevations in the Park range fromsea level to 2228 m at the peak of the Golden Hinde, the highest point onVancouver Island. Heavy snowfalls are common on the mountain slopes and alpineplateaus from November through March. Snow remains all year on the mountainpeaks, and may last into July at lower elevations. A detailed biophysicaldescription of the Park can be found in Kojima and Krajina (1975).In Strathcona Park's 82-year history, much resource exploitation hasoccurred within its confines, including logging and mining. Mining is presentlyoccurring in the south-east region of the Park, near the south end of Buttle Lake.Much of the area surrounding the Park has been, or is currently in the process ofbeing, clearcut logged. At present, 79%, (172,000 ha) of the Park has been zonedeither for preservation as wilderness conservation, or for wilderness recreation,thus controlling mechanized access and facility development (Strathcona Provincial .Park Master Plan Draft, 1992). All land within the Park is closed to hunting, whilethe surrounding areas have a limited-entry hunt for elk. The illegal hunting of elk isknown to occur in all areas surrounding and within the Park (BC Parks; pers.comm.).Janz and Lloyd (1977) reported that elk are primarily distributed in thenorthern and western regions of Strathcona Park and adjacent areas outside thepark. The southern and eastern regions of the park support very few elk. Four-13-regions within or near Strathcona Park's boundaries were chosen as study areas bythe Ministry of Lands, Environment, and Parks (Figure 1). Each of the 4 regionswas deemed important to the study of elk for various reasons, including supportinga past population of elk; proximity to major roadways and heavy public-use zones;easy access for trapping, radio-collaring, and subsequent monitoring of elk; andbeing politically and/or ecologically sensitive (BC Parks; pers. comm.). Thefollowing are brief descriptions of the 4 study areas, including suspected elkpopulation numbers.Elk River ValleyThe Elk River valley study area lies entirely within the northern region ofStrathcona Park. It covers an area of approximately 4450 ha, following along thecourse of the lower Elk River, from the Drum Lakes to the western shores of UpperCampbell Lake.Two roadways running parallel to one another, transect the study area.Highway 28 travels along the south side of the Elk River, and the ERT gravellogging road travels along the north side of the Elk River. Both roads are openyear-round, and are heavily used.Jones (1983) reported that the lower Elk River valley likely provided winterrange for approximately 30 elk, 10 of which he believed to summer in the elkwinter range area of the valley. He thought that in summer the other elk likelyspread out into the upper Elk River valley, Cervus Creek, the upper portions ofIdsardi and Tlools Creeks, and possibly other, unknown areas.-14-Janz and Lloyd (1977) observed 2 distinct groups of elk in the Elk Rivervalley; a small group of 7 adult females in the western portion of the valley, and alarger but unknown number of elk in the eastern portion of the valley up to thewestern shore of Upper Campbell Lake.The elk winter ranges in the lower Elk River valley were believed to be inlargely 30-40 year-old second-growth resulting from logging in the Park. Theproductivity of these habitats is relatively poor when compared to the natural old-growth forest (Jones 1983). In addition, the 1957-58 flooding of Upper CampbellLake permanently destroyed part of the original elk winter range.Thelwood ValleyThe Thelwood valley study area, at the southern end of Buttle Lake, iscompletely within Strathcona Park's boundaries, and consists of approximately 655ha of flood plain, alluvial fan, and valley slopes (Blood 1988). The area oncesupported a small herd of approximately 15 elk, which were last seen in 1973(Jones 1983). A series of natural disasters and human-caused impacts likelyresulted in the extirpation of elk in the Thelwood by the late 1970's.Hydroelectric development in 1957-58 raised the level of Buttle Lake 9 m,resulting in flooding and permanent loss of flood plain habitat in the lowerThelwood valley. The 1958 Thelwood fire drastically affected habitat on the entireelk winter range. Most herbaceous and browse forage produced in the summer of1958 was destroyed, resulting in an extremely poor range for elk in the winter of-15-1958-59. In addition, during the first several years following the fire, there was nocover in the lower Thelwood valley.The Westmin Mine Co. has been in operation since the late 1950's in thearea, and currently occupies much of the Myra Creek valley to the west of theThelwood valley. Westmin has constructed a hydroelectric powerhouse station inthe Thelwood, and a dam at Jim Mitchell Lake, above the valley. Noise anddisturbance associated with mine development, operation, and traffic in theThelwood and nearby Myra Creek valleys may also have had a negative impact inpast elk populations. It is believed that easy access resulting from mine activity ledto the illegal hunting of any remaining elk in the area (BC Parks; pers. comm.).Heber River-Camel Creek ValleysThe Heber River and Camel Creek watersheds are found on the periphery ofthe western boundary of Strathcona Park. The Heber River valley acts as summerrange and the Camel Creek valley as winter range for the population of migratoryelk in the area. Both watersheds are criss-crossed by logging roads, most of whichare open due to the ongoing active logging in some regions of the valley. The vastmajority of the watersheds have already been clearcut logged, as part of the treefarm license managed by Fletcher-Challenge Ltd.Jones (1983) stated that the Heber River area had approximately 100 ormore elk, the majority of which wintered and summered outside the Park. Becausemuch of the elk summer range in the upper Heber River valley is adjacent to thePark boundary, some of the elk may summer in tributaries of the Heber River,-16-within the Park. Observations by MOELP personnel indicated that during severewinter seasons, some elk from the Heber River area may winter in the Elk Rivervalley, within the Park (Jones 1983; Janz and Lloyd 1977).Ucona River BasinThe Ucona River region forms a large basin on the periphery of the central-western Park boundary, at the confluence of several valleys historically importantto elk. These valleys include Quatchka Creek, and the Ucona Basin itself, bothoutside of the Park, and Kunlin Lake, Donner Lake, Pamela Creek, and an unnamedhanging valley inside the Park boundary.All of the winter range for the elk in the Ucona study area, including thoseanimals which summer in the Park, lies outside the Park. The winter ranges havebeen adversely affected by logging (Jones 1983), and the majority of the UconaRiver basin has been clearcut as part of the tree farm license managed by CanadianPacific Forest Products Ltd.The Ucona River elk winter range is currently in an early vegetation serecreated by logging during the mid-1960s (Janz and Lloyd 1977). There are only afew small pockets of mature timber within the area to provide thermal cover duringextreme winters. Logging is currently occurring in the Quatchka Creek valley, andin the Pamela Creek valley on the Park boundary. Aside from the Donner Lakeregion which is old-growth timber and consequently a difficult area to access, theentire Ucona River basin is criss-crossed by an extensive network of well--17-maintained logging roads. Janz and Lloyd (1977) noted a plentiful supply of winterelk forage in the area.Jones (1983) noted that the Ucona River area supports a group ofapproximately 100 or more elk, most of which live outside Strathcona Park. Ofthese, approximately 20 spend at least a portion of the summer within the Park, inthe Pamela Creek valley.METHODSTrapping and Radio-Collaring AnimalsTo determine the seasonal movements and home ranges of elk in StrathconaProvincial Park, 10 adult (> 2 years) female elk were captured and fitted withLMRT-4 radio-collars (Lotek Engineering, Newmarket, Ontario) transmitting in the150-151 KHz range. In March of 1991, prior to the start of my study, a total of 4animals were successfully tranquilized from a helicopter a fitted with radio-collarsby MOELP personnel; 1 in the Thelwood valley, 1 in the Ucona River area, and 2 inthe Heber River valley. In January of 1992, a corral trap was constructed in theElk River valley, baited with alfalfa hay, and used to capture 13 elk, 4 of whichwere radio-collared. In the same year, on February 17 and on March 11, 2 moreelk in the Ucona River Basin were tranquilized from a helicopter and fitted withradio-collars. An upper canine tooth was extracted from each of the 6 tranquilizedanimals for later age determination by cementum analysis (Matson's Laboratory,Milltown, Montana).-18-Population and Group Composition DataFor group composition counts to be representative of the entire population,all sex and age classes should be equally observable (Taber et al. 1982). Thislikely is never true, but it is more likely to occur at some periods of the year thanothers; because of this, group composition counts were conducted throughout theyear. Whenever elk (radio-collared or not) were located visually, records weremade of the number of elk present in each of the following age-sex classes: adultmale ( > 2 years, based in antler development), juvenile mate ( < 2 years, but notyoung of the year), adult female (not young of the year), young of the year, andunidentified. Ground observations of elk groups were made through 8X30binoculars, and a 20-40X60 spotting scope. Aerial observations of elk were madewithout visual aid. In addition to my observations, Park personnel were asked tonote elk numbers and age-sex classes observed during the study period, and wereinterviewed monthly for this data.Animal LocationsField work during the study consisted of 2 intensive and 2 less-intensiveperiods. Intensive periods occurred during the summers of 1991 and 1992 (MayAugust). Less-intensive periods occurred during the winter seasons of 1991-92(September-April), and 1992-93 (September-January). During the second less-intensive study period, animal locations were obtained by BC Parks personnel.Locations of the 10 radio-collared elk were determined by triangulation usinga three-element directional Yagi antenna, a Lotek SRX-400 receiver (Lotek-19-Engineering, Newmarket, Ontario), and aviator headphones. The majority of animallocations were obtained from the ground or from the air during daylight hours, overthe course of a 2-year period commencing in the spring of 1991, and ending in thewinter of 1992-1993.Radio-locations of radio-collared elk obtained from the ground weredetermined in 1 of 2 ways. First, the area the radio-collared elk was initiallybelieved to be in (from the previous location, or from the "best guess") wasscanned with the telemetry equipment on a high gain setting from logging and/orprimary roads. If a signal was not detected, adjacent areas were scanned in thesame manner until a signal was detected. If still no signal could be heard, the areawas noted and left, until an aerial search for the radio-collared animal could beconducted. If a signal could initially be detected, the gain on the receiver wasdecreased incrementally until the signal was just barely audible in the headphones.A field compass was used to take a bearing in the direction from which the signalwas detected, and the bearing line was then drawn on a 1:50,000 topographicfield map of the area. At least 2 more bearings were also taken from differentpositions along the logging and/or primary roads. When the bearing lines did notintersect at a common point, the mid point of the polygon defined by the bearinglines was taken to be the location of the radio-collared animal. The triangulationprocess was conducted as rapidly as accuracy and travel between successivetelemetry "fixes" permitted.-20-For each bearing taken, the observer's exact location on the map had to befirst calculated; this involved taking a series of at least three bearings fromprominent geophysical features, and/or man-made structures such as roadintersections, and then plotting the location on the field map. Because error isassociated with each bearing taken, and with plotting each telemetry location onthe topographic map, permanent telemetry "stations" were set up in some areas.These stations were of known exact location, from which accurate bearings onradio signals could be quickly made.The second, and more commonly followed, process to determine animallocations from the ground was to complete the above procedure, and then to go inon foot to the location determined by triangulation to attempt to visually observethe animals. This added process allowed me to confirm the animal's location, testthe accuracy of the triangulation technique, determine sex and age classificationdata for each elk group observed, determine the habitat type the animals werefound in, and observe elk behaviour.A Bell Jet Ranger Ill or Bell 206 helicopter with a single 3-element Yagiantennae was used to determine animal locations aerially at the start of the fieldseason, and when an animal occasionally could not be located from the ground.On a high gain setting, radio signals from up to approximately 10 km away couldbe received and used to direct the helicopter to the correct watershed, and thegeneral location of the radio-collared animal. Repeated slow passes were madeover the radio-collared animal's general location with the receiver on a low gain-21-setting. When the radio-collared animal and/or any associated elk were visuallyobserved, the location was plotted on 1:50,000 topographic maps of the studyareas. When visual contact with the animal could not be made due to densevegetative cover, the point above where the loudest signal was detected withoutthe antenna, or with a wooden pencil used as an antenna, was taken as the radio-collared animal's location.A Cessna 172 fixed-wing aircraft was also used for aerially determininganimal-locations, primarily during the winter months when ground accessibility tosome areas was not feasible, and time and manpower were constrained. Duringfixed-wing flights, 2 three-element directional Yagi antennae were mounted on theaircraft, one on each wing, and each at an angle of approximately 45 degrees fromthe vertical axis. The procedure to locate radio-collared elk was similar to thatused in the helicopter assisted locations.The inherent errors associated with all two-dimensional representations ofthree dimensional space, coupled with those of the "approximate" nature of a smallscale map, means that each time a location was plotted on the 1:50,000topographic maps, whether from the ground or aerially, a certain amount of errorwas incurred. The 1:50,000 topographic maps were the most accurate mapsavailable for the entire study area. In addition, because these maps were used atall times, any errors associated with plotting location data were assumed to beuniformly distributed throughout the study. As such their use was felt to bejustified.-22-1 tested the accuracy of ground telemetry 4 times throughout the study byhaving someone else place a radio-collar in one of the study areas, and then usingthe above described triangulation procedures, searched for and located it on a map.The actual location of the radio-collar was generally found to be within 50-100 mof the triangulated location. Without visual confirmation, accuracy of radio-collarlocation from the helicopter was thought to be at least this accurate. From thefixed-wing aircraft, without visual confirmation, accuracy was found to be less thanfrom the helicopter; the actual location of the radio-collar was generally within 75-125 m of the aerially determined location.Due to the relatively large number of radio-locations and direct sightings ofthe Elk River valley study animals (elk #'s 120, 179, 218, and 259), theseobservations were recorded by habitat during the 1992 summer season, based onsimple features of structure and composition, and by physiographic features of theselected sites (habitat types modified from Janz and Lloyd 1977).Home Range DelineationProgram Home Range (second edition, Ackerman gt al. 1990) was used togenerate home ranges for all radio-collared animals. Program Home Range allowsestimation of home range size using 5 different methods: bivariate normal ellipse(Jennrich and Turner 1969), weighted bivariate normal ellipse (Samuel and Garton1985), Fourier transformation (Anderson 1982), convex polygon (Hayne 1949,Michener 1979, Bowen 1982), and harmonic mean (Dixon and Chapman 1980).The program is capable of testing underlying assumptions such as independence of-23-observations (Swihart and Slade 1985). Harmonic mean core areas can also becalculated (Samuel gt al 1985, Samuel and Green 1988). Individual location datapoints can be weighted to decrease or eliminate the effect of outliers (Samuel andGarton 1985), to reduce serial correlation (Swihart and Slade 1985), or to includeproportion of time in different activities (Samuel and Garton 1987).Both the convex polygon and harmonic mean methods were used todelineate radio-collared elk home ranges. The convex polygon method was initiallyused to determine home ranges for all study animals because of its historicalprominence as one of the first developed home range estimators, and one of themost widely used. The convex polygon estimator is a simple and intuitive methodfor calculating the area enclosed by a set of locations (Beckoff and Mech 1984). Init, the most peripheral locations in a set of location data are connected by a line insuch a way that the internal angles of the generated polygon do not exceed 180degrees. However, the convex polygon method is strongly biased at relativelysmall sample sizes; is severely affected by outliers, and may include large areas notused by the animal; provides information solely on the area used by the animalduring the exact periods of observation and not on the area potentially used by theanimal; and provides no information concerning how the area within the homerange is used (see Ackerman At a 1990).After initial home range estimation, the minimum convex polygon methodwas not used in this study primarily because I felt that it did not accurately reflectthe actual home ranges of the study animals. In particular, the home ranges-24-generated by this method often included large areas of mountain top not used bythe animals, especially when an animal made use of 2 or more parallel valleys, or alarge bowl and an adjoining small watershed, but not the area between them. Inaddition, the sample bias of the minimum convex polygon method, where thecalculated home range tends to increase as the number of locations increases(Jennrich and Turner 1969), would have made comparisons between animals andseasons invalid. The inaccessibility of certain study areas resulted in largedifferences in sample sizes of animal locations among both seasons and studyanimals.In comparing currently popular home range estimation techniques, includingthe Fourier series, minimum convex polygon, harmonic mean, and 2 - 95% ellipsehome range estimators, Boulanger and White (1990:314) concluded that overall,the harmonic mean estimator shows the "best performance." The harmonic meanmethod of home range estimation is a non-parametric procedure based on thevolume under a fitted three-dimensional utilization distribution (Ackerman el al.1990). The distribution is based on harmonic mean values calculated at grid pointssystematically located throughout the animal's home range (Dixon and Chapman1980). The probability of use at any location in the home range is estimated, andthe utilization distribution is then determined by calculating the harmonic values ateach grid point. When their harmonic values exceed the highest harmonic value forany animal location, grid points are considered outside the animal's home range,and thus excluded from the utilization distribution. Thus harmonic mean estimation-25-is less affected by outliers (Ackerman et al. 1990), and can in fact be used toidentify outliers.The utilization distribution is based on a rectangular grid of square cells. Thisgrid is used to calculate the non-parametric harmonic mean distribution from theanimal's location data. The size of each grid cell in the rectangle is measured inreal world units (e.g. metres), and is dependent on the number of cells and thescale parameter. Each of the animal locations, and all of the generated harmoniccontours, must fit on the harmonic grid, or the estimates will be invalid. Thechoice of the scale and number of grid cells in the rectangle are crucial to ensuringthat the size of plot area is large enough to accommodate all of the data.Ackerman et at. (1990) suggest choosing the scale parameter so that it isapproximately the larger of 1 /8 th of the range of the Y coordinates, or 1 120t h of therange of the X coordinates. However, these suggested levels were found to mostoften result in a "plot scale too small to determine total utilization volume" errormessage, and so were only used as a starting point for determining the optimumscale value. The scale value was then adjusted by increments of 1.0 metres untilthe error message was no longer generated. The grid density parameter shouldachieve an average of 1 observation per grid cell (Ackerman et al. 1990), and thiswas accomplished automatically using the "optimization of grid density" option inthe program.The harmonic mean method of home range estimation was criticized bySpencer and Barrett (1984) and by Worton (1987, 1989), as being overly sensitive-26-to the choice of scale parameters. I also noticed, as did Brunt (1991), that eachanimal's home range size and shape could be severely influenced by changing thescale and grid density parameter values in the home range calculations.The harmonic mean estimator method allows for the use of weighting factorsto be applied to each animal location, to account for the relative amount of timethe animal spent at each location (Braun 1985, Samuel and Garton 1987). Withoutweighting, the program assumes that each location represents an equal amount oftime spent there by the animal, and so tends to overemphasize the relativeimportance of areas sampled more often. I chose to use a weighting factor whenmore than one location per animal was obtained within a 24-hour period. In thosecases, the weighting factors summed to 1.0, and so equalled the weighting factorgiven to all independent location observations.Program Home Range requires selection of a minimum distance betweenobservations, which is essentially the typical accuracy of the animal locations(Ackerman et al. 1990). I set the minimum distance between observations at 50 mfor all home range calculations, because this was considered to be the minimumerror value of triangulated animal locations.Dates for delineating seasons were based on study animal movementsbetween seasonal ranges. Not all study animals exhibited obvious migratorybehavior or seasonal shifts in areas of use. For those elk demonstrating migratorybehavior, the timing of migratory movements differed between years, and between-27-study animals, so seasons delineated for the purpose of home range calculationsare presented in Table 1.Animal/Season Selection CriteriaOver the course of the 2-year field study, some radio-collared animalsassociated with others for varying time periods. For the purpose of home rangeestimation, elk were considered to be associated with the same group wheneverthey were located within approximately 200 m of each other (Brunt 1991). When2 or more elk were considered to be associated together during part or all of aparticular season, a degree of association was assessed to determine whichanimals had sufficiently different seasonal range selection patterns to warrantcalculation of individual animal/season home ranges. If the proportion of a givenseason spent with other study animals was > 90%, one member of the associatedanimals was randomly selected for seasonal home range calculations (Brunt 1991).By contrast, individual home ranges were calculated only for animals whoseassociation with other study animals was < 90%, and for groups consisting of 2 ormore study animals whose association with each other was > 90%. Table 2outlines the 13 animal/seasons for which home ranges were calculated.The 4 radio-collared adult females in the Elk River valley (#'s 120, 179 ,218, and 259) were found to be closely associated with each other > 90% of thestudy period commencing with their collective capture in January of 1992. Forbrief periods during the calving season in the spring of 1992, 1 or more of the 4Table 1. Dates used to delineate seasonal home ranges for each study animal.ERV Groupe Cumulativeb Jan 24/92 - Jan 6/93159 Cumulativeb Mar 11/92 - Jan 6/93300 Cumulativeb Mar 3/92 - Jan 6/93339 Cumulativeb Jun 15/91 - Jan 6/93499Cumulative Jun 14/91 - Dec 10/92Summer 1991 Jun 14/91 - Nov 9/91Winter 1991-92 Nov 10/91 - Jun 4/92Summer 1992 Jun 5/92 - Sep 9/92Winter 1992-93 Sep 10/92 - Dec 10/92580 Cumulativeb Jun 13/91 - Jan 6/93688Cumulative Jun 17/91 - Jan 6/93Summer 1991 Jun 17/91 - Dec 21/91Winter 1991-92 Dec 22/91 - May 29/92Summer 1992 May 30/92 - Nov 7/92Winter 1992-93 Nov 8/92 - Jan 6/93"ERV = Elk River valley, and includes elk #'s 120, 179, 218, and 259 (Note: #218mortality on December 20, 1992).bBecause the elk did not demonstrate migratory behavior, seasonal ranges could notbe delineated.-28-Table 2. Home ranges analyzed using the harmonic mean contour method.Home Range Estimated Study Animals Total Ranges AnalyzedCumulative' ERV Groupb , 159, 300,339, 499, 580, and 6887Summer 1991 499, 688 2Winter 1991-92 499, 688 2Summer 1992 499, 688 2aYear-round home range using all animal locationsbERV= Elk River valley, and includes elk #'s 120, 179, 218, and 259-30-study elk were located in areas distinctly separate from the main group. Elk #259left the main group for a short period in the fall of 1992.The 2 adult females radio-collared in the Heber River watershed wereassociated with each other for portions of both winter and summer seasons, butnot for periods > 90% of any 1 season. Therefore, the 2 Heber study animalswere treated as non-associated elk, and separate home ranges were calculated foreach. Two of the 3 study animals in the Ucona River basin (#159 and #339) wereclosely associated with the same group during the summer of 1992. However,whereas elk #339 was radio-collared in the spring of 1991, elk #159 was radio-collared in the spring of 1992, so it is not known whether the 2 were associatedprior to the summer of 1992. No other study animals (elk #580 in the Thelwoodvalley, and elk #300 in the Ucona River basin) was observed In close association"with other radio-collared animals.Home Range AnalysisThree types of home range were calculated and compared for mostanimal/seasons: i) cumulative home ranges were determined from all animallocations obtained during the study period; ii) seasonal home ranges wereestimated from all non-outlier locations obtained during the seasons listed in Table1; and iii) core use areas were delineated as areas of concentrated use (see below)within each seasonal range.Cumulative home ranges were delineated for each study animal as the areaincluded by 100% of the locations obtained during the study. The cumulative-31-range provides a means of defining all areas available to the animal for seasonalrange selection (Brunt 1991). However, animals may occasionally and temporarilyleave their normal activity areas, producing extreme locations (outliers) that canseriously affect home range estimates. Outliers may be the result of mapping andcoding errors in the location data (Ackerman et al. 1990), and can also representtransitional locations between seasonal ranges. In accordance with Burt's (1943)classic definition of home range, these occasional excursions outside the normalarea of use were not considered part of the home range. Program Home Rangeallows for the calculation of contours which contain a specified percentage of theanimal's utilization distribution, and thus identifies outliers.To calculate seasonal home ranges, I used the 95% contour to reduce theeffects of potential outlier values, in accordance with Burt's (1943) home rangedefinition and the convention followed by others (Jennrich and Turner 1969,Schoener 1981).Core Use AreasThe concept of core use areas has received considerable use in theecological literature, and has generally been used to define central areas of intenseuse, in an effort to describe the internal anatomy of the home range (Ackerman etal. 1990). Core use areas are important because they are areas of particularlyhigh home range usage, and thus they often may provide a more clear measure ofthe changing pattern of range use than does the total home range area (Harris et al.1990). Core use areas are often more useful for understanding both intraspecific-32-and interspecific patterns of home range use, than are the more peripheral contours(e.g. 100% minimum convex polygon contour, 99% harmonic mean contour, etc.).Harris et al. (1990) cited an example of this phenomenon in a study comparing roedeer (Capreolus capreolus) and muntjac (Muntiacus reevesi) in-the same habitat.The number and relative position of core use areas differs between the species, butsuch differences are masked when comparing total home ranges. In addition, whilemale and female core use areas do overlap, they do not occupy the same space. Inthe case of male muntjac ranges, which may overlap to some extent, their core useareas are mutually exclusive.An option in Program Home Range allows core use areas to be automaticallyidentified by comparing the utilization distribution from harmonic mean calculationswith a uniform use model (Samuel ex t. 1985). Core areas are defined as themaximum area where the observed utilization distribution (based on harmonicvalues) exceeds a uniform distribution (Ackerman et al. 1990). Program HomeRange then uses a chi-square test on the ordered cumulative distribution ofobserved data, compared to the uniform model (Samuel and Green 1988). Anillustration of the statistical test and further description of the above describedmethod are presented in Samuel et al. (1985) and Samuel and Green (1988). Theaverage utilization distribution contour of the 10 home ranges calculated using thismethod was 60.0% (SD= 5.8%).Dixon and Chapman (1980) also described the core use area as the 50%harmonic mean contour. If Program Home Range's automatic core use area-33-estimator was unable to detect a significant core use area present in a studyanimal's home range, the 50% harmonic mean contour was used instead. Theharmonic mean contour actually plotted was the closest one to 50% that could becalculated given the study animal's location data. For the 3 core use areas forwhich Program Home Range's automatic method could not determine the presenceof a significant core use area, the average of the contours plotted was 69.7%(SD =10.209%). This was not significantly different from the average utilizationdistribution of 60.0% for core use areas determined by the automatic method(t = 2.098, tcritical = 2.201, df =11, a = 0.05).Core use areas are not considered reliable when S 12 animal locations areused to delineate a home range (Ackerman et al. 1990). Therefore, core use areasdetermined for the following 4 seasonal ranges are not considered reliable: summer1992 and winter 1992-93 for elk #'s 499 and 688. The 13 core areas that wereestimated and plotted are presented in Table 3, along with the method used tocalculate them (Program Home Range's automatic core use area calculation, or the50% harmonic mean contour).Statistical AnalysesA probability level of 0.05 (95% confidence level) was used in all statisticalanalyses. Walpole (1982), Zar (1984) and A. Kozak (pers. comm.) were consultedto determine appropriate statistical procedures. Statistical tests were conducted ona hand calculator.-34-Table 3. Core use area delineation methods and core area and home range sizes.HomeRangeEstimatedStudyAnimalCore AreaCalculationMethodProportionUtilizationDistributionSeasonalRange Size(km2)Core UseArea Size(km2)Summer1991499 50% Contour 0.64 31.82 6.53688 Automatic 0.68 36.97 13.12Winter1991-92499 Automatic 0.57 61.16 17.56688 50% Contour 0.61 18.59 1.37Summer1992499 50% Contour 0.84 7.24 0.43688 Automatic 0.64 63.25 11.86CumulativeERVa Automatic 0.65 32.38 10.14159 Automatic 0.67 41.53 13.71300 Automatic 0.54 28.15 6.21339 Automatic 0.63 26.14 10.90499 Automatic 0.57 95.98 32.28580 Automatic 0.51 15.93 3.51688 Automatic 0.54 155.67 29.11aERV= Elk River Valley Group, and includes elk #'s 120, 179, 218, and 259.-35-RESULTSMortalitiesOne of the 2 adult female elk first radio-collared in the Ucona River Basindied within a month after trapping and radio-collaring activities. Cementum annulianalysis indicated that elk #199 was 4 years old (accuracy + I- 1 year). It wasclear from the mortality site that elk #199 had been preyed upon by anundetermined number of wolves, whose tracks and bite marks on the radio-collarwere obvious. A second study animal mortality occurred on December 20, 1992,when one of the Elk River valley study animals, elk #218, was shot by a limitedentry hunt ticket holder within 50 m of the Strathcona Park boundary (BC Parks,pers. comm.).Animal LocationsA total of 535 locations was obtained during 5 seasons from June 13, 1991through January 6, 1993. Ground radio-telemetry was used to obtain279 (52.15%) of the locations, and aerial radio-telemetry was used to obtain theremainder (256 or 47.85%).Group Composition and PopulationGroup composition counts were conducted throughout the year in theThelwood valley, Heber River-Camel Creek, and Ucona River study areas; the ratiosand percentages of group sex-age classes were determined for both summer(1991-92 combined) and winter (1991-92 combined) seasons (Table 4). Groupcomposition counts for the Elk River valley were also conducted throughout theTable 4. Classification counts for summer and winter seasons in study area.I Elk River Valley - Summer° 1992Juvenile Male Adult Male Adult Female YOYb Total# 2 3 9 4 18Ratio 22.2 33.3 100 44.4% 11.1 16.7 50 22.2 100(Elk River Valley - Winter' 1992Juvenile Male Adult Male Adult Female YOY Total# 1 2 9 1 13Ratio 11.1 22.2 100 11.1% 7.7 15.4 69.2 7.7 100The!wood Valley - Summers 1991-92Juvenile Male Adult Male Adult Female YOY Total# 0 2 1 0 3Ratio 0 200 100 0% 0 66.7 33.3 0 100^IThelwood Valley - Winters 1991-92Juvenile Male Adult Male Adult Female YOY Total# 0 2 1 1 4Ratio 0 200 100 100% 0 50.0 25.0 25.0 100aSummer season from May through OctoberbY0Y = young of the year (calves)°Winter season from November through April-36-Table 4 (continued). Classification counts for summer and winter seasons.Heber River Watershed - Summers' 1991-92Juvenile Male Adult Male Adult Female YOYb Total# 14 27 216 51 308Ratio 6.5 12.5 100 23.64.5 8.8 70.1 16.6 100Heber River Watershed - Winters` 1991-92Juvenile Male Adult Male Adult Female YOY Total# 0 0 49 7 56Ratio 0 0 100 14.3% 0 0 87.5 12.5 100Ucona River Basin - Summers 1991-92Juvenile Male Adult Male Adult Female YOY Total# 1 19 40 8 68Ratio 2.5 47.5 100 20% 1.5 27.9 58.8 11.8 100Ucona River Basin - Winters 1991-92Juvenile Male Adult Male Adult Female YOY Total# 0 2 9 1 12Ratio 0 22.2 100 11.1% 0 16.7 75.0 8.3 100aSummer season from May through OctoberbY0Y =young of the year (calves)`Winter season from November through April-37--38-year (1992); however, I believe the total sex-age class numbers presented areaccurate, and not just indices of group composition, as in the other study areas.Home Range Characteristics and MovementsHome ranges and core use areas were estimated for all 10 radio-collaredstudy animals (Figures 2-6). Location data indicated that the 2 Heber River elk(#499 and #688) were migratory, and the remaining 8 radio-collared elk were non-migratory. Seasonal home ranges and core use area sizes were determined for the2 migratory elk for summer 1991, winter 1991-92, and summer 1992 (Table 5).Although data continue to be collected at the time of this writing, seasonal rangeand core use area size could not be estimated for the winter 1992-93 season, dueto insufficient locations having been obtained by the end of my study period.Cumulative home range and core use area sizes using all location data obtainedover the study period were calculated for all study animals and are presented inTable 6.Elk River ValleyThe core group of 16 elk in the Elk River study area consisted of 9 adultfemales, 4 of which were radio-collared, 4 calves, 3 adult males, and a juvenilemale (Table 4). This group was directly sighted over the 1992 summer period onaverage once every 4 days, and numerous times throughout the winter season;consequently the group size and composition is believed to be accurate.The largest number of elk in the Elk River valley during the summer seasonwas found to be 18 animals. From the distribution of sightings in the valley, it isglIM OM OM 4PBHighway 28ERT Logging Road0 1.5^3.0 kmNCORE USE AREADrum Lakes...,-T1EC-CDco0CDCDXCUMULATIVE RANGEFigure 2. Cumulative range and core use area for elk #'s 120, 179, 218, and 259, in the Elk River valley.Figure 3. Cumulative range and core use areas for elk #580 in the Thelwood valley.2 4kmWINTER RANGEWINTER CORE USE AREASUMMER RANGESUMMER CORE USE AREASTRATHCONA PARK BOUNDARY- N^ Crest Lake— — -- Logging RoadHighway 28Figure 4. Summer and winter (1991-92) seasonal ranges and core use areas for elk #499 in the Heber River-Camel Creek Valleys.este k•^ ■SMA Ns.^- • - ' --BOUNDARYCQA PARKFigure 5. Summer and winter (1991-92) seasonal ranges and core use areas for elk #688 in the Heber River-Camel Creek Valleys.Figure 6. Cumulative range and core use areas of elk #'s 159, 300, and 339in the Ucona River Basin.Table 5. Sizes (km 2) of seasonal home rang& and core use areas for 2 non-migratory elk from the Heber River-Camel Creek valleys.Season Dates StudyAnimalnLocationsSeasonalRange SizeCore UseArea SizeSummer1991Jun 14-Nov 9 499 5\11b 31.82 6.53Jun 17-Dec 21 688 9\6 36.97 13.12Winter1991-92Nov 10-Jun 4 499 13\4 61.16 17.56Dec 22-May 29 688 9\4 18.59 1.37Summer1992Jun 5-Sep 9 499 5\7 7.24 0.43May 30-Nov 7 688 7\4 63.25 11.86aHome range estimated using 95% of utilization distributionbAerial locations/ground locationsTable 6. Sizes (km 2) of cumulative (year-round home range using 100% ofutilization distribution) and core use areas for all study animals.Study Area Study Animal n Locations CumulativeRange SizeCore UseArea SizeElk River120 24/478 32.38 10.14179 24/47 32.96 9.77218 22/47 29.70 9.54259 23/47 32.26 9.58Ucona RiverBasin159 21/6 26.14 10.90300 21/7 28.15 6.21339 32/14 41.53 13.71Thelwood 580 31/28 15.93 3.51Heber River-Camel Creek499 29/22 95.98 32.28688 28/14 155.67 29.11-45-aAerial locations/ground locations-46-possible that several more elk, likely in small "bachelor" groups of 2-3 adult malesmake use of the valley in the early spring and late fall. If this is indeed the case,the maximum population of elk in the Elk River valley approaches 20 individuals.In the winter season, the maximum number of elk observed was 13 animals.The Elk River valley group was found to use a total of 7 habitat types duringthe summer period of May 21 - August 30, 1992 (Table 7). Riparian willow (Salixspp.) flats, wet meadows or bogs, the grassy right-of-way below the powertransmission fines, and second growth coniferous forest (predominantly Douglas firPseudostugas menziesii)  and western hemlock (Tsuga heterophylla), with somelodgepole pine (Pinus contorta latifolia)) were the habitat types most frequentlyused by this group. While the Elk River valley group made repeated use of theirentire home range throughout the summer, I observed that as the water levels inthe Elk River dramatically decreased over the course of the summer season, the elkmade greater use of wet meadows or bogs and the river estuary, and less use ofriparian willow flats.Thelwood ValleyThe Thelwood valley presently supports a known population of 4 elk,consisting of 1 adult female, 1 calf of undetermined sex, and 2 adult male elk. Thecalf was born in the spring of 1992.The majority of locations in the Thelwood valley of elk #580 were nearlyevenly spread between 2 areas; low elevation (223 m), 30-year old coniferousTable 7. Habitat types' used by the Elk River valley group b during summer 1992(May 21-August 30).Habitat Type # Observations` %UseSecond growth conifer forest 4 8Mature conifer forest 1 2Power transmission lines 12 24River estuary - lakeside 2 4Bog - wetland 8 16Riparian willow flats 5 10Riparian second-growth deciduous forest 18 36Totals 50 100*modified from Janz and Lloyd (1977)bElk #'s 120, 179, 218, and 259`Direct sightings or locations later confirmed by presence of elk sign-48-second growth and adjacent regenerating riparian alder (Alnus rubra) flats, andhigher elevation (450-700 m) riparian areas and vegetated slides. Young alderbushes, salmonberry (Rubus spectabilis), thimbleberry (Rubus parviflorus), andbracken fern (Pteridium aquilinum oubescens)  were the among most commonunderstory vegetation associated with animal locations.Heber River-Camel Creek ValleysThe Heber River watershed likely supports a population of approximately 100elk during the summer season. These 100 elk are found in several groups of 10-30individuals which repeatedly congregate over the summer in groups of up to 70 elk.In the winter season, the lower Heber River valley supports few elk, probably nomore than 30 individuals; the upper valley, with a higher elevation and almost nosnow-interception or thermal cover, supports even fewer. It is not known howmany elk use the Camel Creek valley during the summer season. During thewinter, I believe that between 20 and 40 use the area.In the summer season, elk in the Heber River watershed (#499 and #688)made use of the 20-30 year-old dense Douglas fir and western hemlockregeneration along the valley bottom, and the clearcut sideslopes above.Vaccinium spp. dominated the understory layer. Summer range elevation variedfrom 750-1400 m. Elk #688 left the Heber River watershed and spentapproximately half of the summer seasons outside of the study area.In the winter season, both elk #499 and #688 were associated together witha small group which migrated to the Camel Creek valley, a distance of-49-approximately 43 km (measured from the northernmost point in elk #499's summerrange to the southernmost point in #499's winter range). Winter range elevationvaried from 350 m in the lowermost Heber River watershed to 750 m in the CamelCreek drainage system. Most of the Camel Creek valley has been clearcut logged,the lowermost portions of which consist of Douglas fir regeneration forest.Ucona River BasinThe Ucona River Basin elk population likely numbers approximately 100individuals, which spend both winters and summers in the basin and its associatedtributaries. During the winter season this number may increase to approximately125-140 elk, which migrate to their winter range in the Ucona River Basin fromsurrounding watersheds.The Ucona River Basin study animals made similar use of habitat over thecourse of the study period, although in different areas of the river basin. In thefirst year of the study, elk #339 made extensive use of the moist riparian areas inthe Quatchka Creek drainage, and the thick Douglas fir and western hemlockregeneration forests on the sideslope. Almost all locations were found in areas ofvery thick cover, where the predominant understory vegetation was thimbleberry,salmonberry, and sword fern (Polystichum munitum).  In the second year of thestudy, elk #339 was closely associated with elk #300 during the winter, and elk#159 in the summer. In the summer season, elk it's 339 and 159 were found in aclearcut "hanging valley" recently added to Strathcona Park (1987), grazing on the-50-sideslopes. Elk #300 made use of moist riparian habitat in the Pamela Creekdrainage, and of the vegetated slides adjacent to the riparian areas.DISCUSSIONMortalitiesWolves are the most important natural predators of elk on Vancouver Island(Janz and Becker 1986), and are known to live in the Ucona River Basin (BC Parks;pers. comm). The death of elk #199 was likely a natural occurrence of predation,and unrelated to earlier trapping and radio-collaring activities. Janz at al. (1980b)noted that predation, especially by wolves, on ungulate populations has in somecases been shown to be a major limiting factor on population growth, althoughrelatively little is known of their impact on elk populations, particularly on Rooseveltelk. However, in a study of wolf food habits on Vancouver Island, Scott (1979)found that Roosevelt elk made up 28% by relative weight of wolf scat contentduring her May - October study period. In Banff National Park, in addition tocollisions with trains and automobiles, predation by timber wolves is the majorsource of mortality for Rocky Mountain elk (Woods 1991).The Elk River valley group, including elk #218 which was legally hunted, waslocated within the Park on December 10, 1992, and later on December 23approximately 2 km east of the Park boundary, where they were using the rockyoutcrop habitat on the south-facing slopes above Upper Campbell Lake. I believethat the elk had moved out of the Park because of deep snowfall accumulation-51-which had occurred in the latter half of December. Although the Elk River grouphad not previously been observed outside the valley, Janz and Lloyd (1977)indicated that this area outside the Park was part of the group's winter range.Home Range Characteristics and MovementsBrunt et al. (1989) found that on Vancouver Island the home range sizes ofmigratory elk were approximately 6 times larger than those of non-migratory elk.This difference is consistent with my findings, in which the cumulative home rangesizes of migratory elk were 2.3 - 9.8 (mean = 6.1) times larger than those of non-migratory elk. Brunt gt al. (1989) estimated the winter home range size ofmigratory Roosevelt elk to vary from 2.2 - 43.8 km 2 , and the summer home rangesize to range from 3.5 - 29.2 km 2 . The present study estimated the winter homerange size to vary from 18.6 - 61.2 km 2 , and the summer home range size to varyfrom 7.2 - 63.3 km2 . Graf (1955) reported that non-migratory Roosevelt elk in theCoast Range of Oregon occupied home ranges of approximately 2.6-5.2 km 2, andin California, Franklin Di al. (1975) estimated the home ranges of non-migratoryRoosevelt elk to be 2.9 km2 . These values are smaller than those I estimated forall study animals in the present study except for the Thelwood elk. However,operational differences concerning the nature of "home range" described in eachstudy precludes realistic comparisons among home range sizes listed in these 2published studies; neither study reported how the home ranges were calculated,nor what kind of home range was estimated. And as Arcese (1989) observed,-52-home range sizes may vary with population size, and therefore do not provide auniversal scale for inter-specific comparisons.Elk River ValleyThe elk population estimate of the Elk River valley in the present study isreasonably close to Jones' (1983) estimate, which was "...winter range for about30 elk... probably about 10 of these summer in the winter range area." Thediscrepancy may be a result of Jones' (1983) underestimation of the elk carryingcapacity of the area. Given the amount of highly suitable elk habitat in the area(see Chapter 2), I believe that the Elk River valley could sustain at least 50% moreelk that it presently does.Janz and Lloyd (1977) determined that movements of the elk in the lowerElk River valley were confined to the valley bottom and low elevation, windwardsidehills. This is consistent with the findings of my study in which all 4 radio-collared elk showed a high degree of philopatry to the lower Elk River valley overthe course of the study period. The tenacity with which an animal returns tospecific areas or groups of like animals is generally referred to as philopatry(Greenwood 1980). Philopatry, and its opposite - dispersal, are relative rather thanabsolute terms. Several studies have shown adult elk to have high degrees ofphilopatry (Knight 1970, Adams 1982, Morgantini 1988, Woods 1991), however,as Woods (1991) noted, theoretical criteria for defining philopatry (or dispersal) arelacking.-53-Population trend is one of the major indicators of elk response to habitatchanges (Janz et al. 1980a), and as such is important to understanding the presentwildlife-habitat situation, and for forming the context in which future managementdecisions can be made. During Janz and Lloyd's (1977) 1976 winter study, the"A-frame" group and the "Campbell River" group (comparable to the "Elk Rivervalley" in the present study) were the only elk found to inhabit the Elk River valleystudy area. The "A-frame" group consisted of 7 adult females, and confined itsactivity to an area composed of moss and lichen-covered rock bluffs and theadjacent second growth (predominantly hemlock and lodgepole pine) in thewestern-most region of the Elk River valley outside of the "Campbell River" group'shome range. This area was repeatedly searched for signs of elk presence in thesummer, and although some fresh and old sign of deer, and old bear scat wasfound, no sign or observations of elk were made either in the summer or winter. Ibelieve that this area is not currently used by elk, and may only be used in severewinters where the requirements of a small group of elk can be met in acomparatively small area with high structural and compositional diversity ofvegetation.The'wood ValleyThe very small number of elk (4) in the Thelwood valley is of concern giventhe apparent ease with which the previous population of at least 15 animals wereextirpated (BC Parks; pers. comm.). Paternity of the calf is presently unknown, asis the possible maternal or sibling relationship between the adult female and each-54-adult male. It is unlikely, however, that the female is the mother of either malebecause she is 4 years old ( + /- 1 year accuracy) based on cementum annulianalysis, and the males are probably at least as old, based on the size anddevelopment of their antlers. The ultimate survival of the current population hasbeen questioned (BC Parks; pers. comm.), and an immediate transplant of non-migratory elk into the valley recommended (Blood 1988). I believe that such atransplant is warranted.The 3 distinct core use areas in the Thelwood valley used by the elk werenot seasonally divided; movement between each area occurred throughout theyear. Some of the movement between the areas may be related to the noise andother disturbances associated with the Westmin Mine Co.'s test-drilling operationsin the 1991-92 and 1992-93 winters. It was observed that when the drillingprocess began, in which heavy machinery and equipment generating loud noiseswas moved into the lower area used by the elk, the animals began using the higherarea. However, animal locations during the winter seasons were not obtainedfrequently enough to confirm the hypothesis that the elk moved in reaction todisturbance by humans.Heber River-Camel Creek ValleysDuring 2 years of study, primarily in the summer season, elk #688 spentapproximately 1/2 of the season outside of the study area. It is believed that shemade use of the ridge tops north and west of the Heber River valley, on the sideslopes and moist valley bottoms approximately 8 km to the north, near Gold Lake,-55-and areas >10 km north of Gold Lake (BC Parks; pers. comm.). Due to theinaccessibility of the regions this animal used, and because she could not beaerially located for much of the time, location data for #688 are limited andseverely underestimate the animal's cumulative and summer season home range,and her core use areas, and seasonal movements. This must be considered whenany comparisons are made between the home ranges occupied by the 2 migratorystudy animals.Ucona River BasinWhile elk #339 repeatedly used different areas of the Ucona River Basinregardless of season, I believe that it is likely the animal shifted its primary use ofthe Quatchka Creek valley to the hanging valley as a result of clearcut logging ofthe Quatchka area. However, the relatively few locations obtained between thefirst and second summer seasons when the shift occurred cannot substantiate thishypothesis.Additional years of animal location monitoring will determine the degree ofassociation between the 3 study animals, and may determine the number ofdistinct elk groups found within the Ucona River Basin study area. Franklin et(1975) observed that precise boundaries between distinct Roosevelt elk groups didnot exist, and concluded that the absence of outside animals in core use areas andtheir infrequent use of other parts of the home range showed that spacing betweenpopulations does occur. This confirms Graf's (1955) observation that closely-56-adjoining home ranges seldom overlap, and that there is little trespassing by onegroup on the range of another.Dasmann and Taber (1956) noted that in areas of dense cover, cervids oftenform small groups or are solitary, and form larger groups when inhabiting openareas. Franklin at al. (1975) concluded that Roosevelt elk population size appearsto be affected by habitat in a similar way. Observations made in this study areconsistent with these findings. During the summer season in open, cut over areassuch as those found on the clearcut logged slopes in the Heber River and UconaRiver watersheds, elk were more often found in large groups. In areas of moredense cover within the above watersheds, smaller elk groups and sign of smallerelk numbers were observed.Migration and Non-migration as Movement StrategiesCervus elaphus have the widest natural distribution of any wild ungulate inthe world (Clutton-Brock et al. 1982), and have a range of movement behaviorsincluding migration, non-migration, and partial migration (Adams 1982). Sinclair(1983) defined migration in vertebrates as the repetitive seasonal movements ofindividuals between distinct areas. Non-migration refers to the movement behaviorof individuals which use the same area throughout the seasons.Seasonal movements of elk are population-specific, thus it is important torecognize inter-population differences. Significant variation was seen in themovement strategies of the elk in the present study. In the case of the 2 HeberRiver-Camel Creek study animals, each elk spent a portion of both summer and-57-winter seasons together on the same ranges, but migrated at different times andvia potentially different routes. In the Elk River valley, all 4 study animals wereobserved to be closely associated with the rest of their group throughout the year,save for very brief periods during the calving and rutting seasons. The Thelwoodvalley study animal made use of 3 separate areas in its home range, but notseasonally, and the 3 Ucona River Basin elk shifted their core use areas somewhatover the course of a year, but not distinctly in that all regions of the study area stillreceived use. The literature is full of similar variations in migratory, non-migratory,and a mix of the 2 types of movement strategies.While Rocky Mountain elk at Jackson Hole, Wyoming, migrated in largegroups (Boyce 1991), elk in a Montana population moved individually or inmatriarchal groups (Knight 1970), and elk in Banff National Park, Alberta, migratedsingly or in small groups (Woods 1991). Although most migratory elk winter at lowelevations, and summer at high elevations, individual behaviors such as multiplemigration cycles per year and specific migrations to rutting areas illustrate thatmigration patterns vary widely among individuals (Woods 1991).Harper et al. (1967) suggested that historically, 2 kinds of Roosevelt elkwere found in California, migratory and non-migratory. Most groups of Rooseveltelk in Washington, Oregon, and California did not demonstrate definite migratorymovements, although they do make seasonal movements in response to forageconditions (Graf 1943, Franklin et al 1975). Adams (1982) noted that suchmovements resemble migration in that they generally are to higher elevations in-58-summer and to valleys and lower elevations in winter. These movements differfrom migration in that they are not periodical. If the right combination ofenvironmental variables are found in an area, the elk will remain there year-round;for example, if forage becomes less available, they may move to or in search of amore suitable supply area. However, some groups of elk, such as those of theOlympic Peninsula, Washington (Schwartz and Mitchell 1945), and on AfognakIsland, Alaska (Troyer 1960), are migratory in response to severe winter weatherconditions.Madison (1966) observed that the Tule elk (C. e. nannodes) in California isnon-migratory, and that there is no evidence that it ever was. The Tule elk utilizevarious portions of their range in response to seasonal variations in foodavailability. McCullough (1969:47) stated that:...such seasonal movements are not regarded as migrations forthe following reasons: i) summering areas are not inaccessiblebecause of weather during the winter period; ii) movements arenot consistent among herds; and iii) the timing of movementsdiffers from herd to herd.McCullough (1969) considered these movements to be solely local shifts inresponse to local conditions.Considering the wide range of movement strategies utilized by elkthroughout western North America, why then are some of the elk in the presentstudy utilizing a non-migratory strategy, while others a migratory strategy?Migration should be favored when the benefits of moving outweigh the benefits ofnot moving (Baker 1978). Woods (1991) listed the following potential benefits of-59-seasonal migration: i) access to better quality and quantity of food; ii) reducedcompetition for food; iii) escape from predation and other forms of harassment; iv)access to increased mating opportunities; and v) avoidance of extreme seasonalweather. Potential costs to migration include: i) increased energy requirements fortravel; ii) time lost from other activities; and iii) increased risk of mortality due totravel hazards and predation. Within a species, or within a population, allindividuals may not face the same costs and benefits of migration. Woods (1991)cited the example that the cost/ benefit ratio evaluation may vary with the size ofthe individual, and that individuals may vary in their genetic tendencies to migrate.By far the most favored proposal for the benefit of altitudinal migration inNorth American ungulates living in mountainous areas is that of a nutritionaladvantage over non-migration (Hebert 1973, Shackleton 1973, Morgantini andHudson 1983). Because forage quality generally peaks in young, rapidly growingplants and then declines as the plants mature (Nelson and Leege 1982), animalscould benefit from tracking early growth stages as spring advances from lower tohigher elevations. If this is the case, then at any time during the growing season,altitudinal migratory animals should have access to more nutritious forage than ifthey had remained at low elevations. However, Woods (1991) concluded thatempirical data for various North American ungulates regarding forage quality,animal condition, and animal movements, do not consistently support thehypothesis that altitudinal migratory animals have access to better quality food.Using a model, Boyce (1991) showed that seasonal fluctuations in food availability-60-on high, summer Rocky Mountain elk ranges could determine the fitness ofmigratory and non-migratory elk. In areas with low seasonality, non-migratoryindividuals had higher fitness over migratory animals because they had no migrationcosts. At moderate levels of seasonality, fitness of both migratory and non-migratory animals decreased, although the fitness of non-migratory individualsdecreased more sharply. Migrants do not universally gain access to better qualityfood, even by moving to higher elevational ranges.When one considers the large geographical range of Cervus elaphus (Clutton-Brock in al 1982) and the diverse habitats occupied by the species (Geist 1982), itis likely that the "nutritional payoff" for migration will vary widely amongpopulations. In areas like the upper Heber River valley, where summer conditionson low elevation winter ranges are relatively hot and dry, large differences in foragequality may exist between low and high elevation sites. In more mesic and coolerareas like the Elk River valley, the nutritional benefits of migration could be reducedor non-existent.Murie (1951), Baker (1978), and McCullough (1985) have presented analternative explanation for choice of movement, based on learning. The suggestionis that young animals develop patterns of home range use through individualexperience as well as association with others (particularly maternal association).Woods (1991) observed that in Banff National Park, where a mother-young pairwas continuously tracked, the daughter adopted the movement strategy of themother. However, although Clutton-Brock g al. (1982) demonstrated a close-61-association between mother-young pairs in red deer during the first years of thedaughter's life, Woods (1991) notes that this trait has not been well documented inelk, or in other ungulates.Life history traits are often phenotypically plastic; that is, a single genotypeproduces a range of phenotypes depending on the environment (Lessells 1991).Morgantini (1988) commented that flexibility in movement strategies and patterns,allows elk to adjust to a variable environment, and Boyce (1991) noted that suchphenotypic plasticity could be shaped by natural selection. Bergerud (1974)suggested that such adaptability explained variation in caribou (Rangifer tarandus)movements, and was a major adaptation in that species. Woods' (1991)observations of the movement patterns and variations in a Rocky Mountain elkpopulation supports these views. Woods (1991) noted that such flexibility allowselk to make a conditional assessment of density and environmental variables, andto either migrate or stay accordingly.As Morgantini (1988) and Woods (1991) concluded, migration may not be aspecies characteristic of elk, as has been implied by many authors, but rather anadaptive and versatile response to a specific environmental situation. This isconsistent with Geist's (1982) "opportunism" theory of elk behavior, and Woods'(1991) observation that no one variable (e.g. diet quality, previous experience,harassment by predators) is likely to explain the range of foraging and movementbehaviors seen in the species.-62-Home Range EstimationThe basic purpose of a home range estimator is to provide a quantifiable areadescribing the area traversed by an individual animal in the course of its normalactivities over a given time period. Because each home range estimator definesthis quantity differently, home range estimates should be viewed only as a generalmeasure of animal activity areas, and comparisons between studies of home rangesshould be taken only as general guidelines given the limitations of estimators(Boulanger and White 1990). Even within the harmonic method of home rangeestimation in a program such as Program Home Range, an animal's home range sizeand shape can be severely influenced by changing the scale and grid densityparameter values in the home range calculations. Ackerman et al. (1990)suggested that in order for harmonic mean home range estimates to be mostcomparable between animals, the scale and grid cell parameters should be thesame. Their suggestions for accomplishing this were followed in the present study,where each animal's location data was run separately with a "reasonable" choiceof scale parameter value, and the automatic grid density calculation option. An"average or typical" value for the scale and grid density values, based on the initialruns for each animal, was chosen and used to re-run all of data for each animal.Scale and grid density values are only comparable between study animals of similarmovement patterns; that is, the home ranges of non-migratory study animals areonly strictly comparable with those of other non-migratory study animals using thesame average scale and grid density values. Changing the home range size and-63-shape in this manner reinforces the notion that quantified home ranges should beaccompanied by detailed descriptions and maps of range areas to supplementcalculated areas (Harris et al. (1990).Harris et al.'s (1990:97) review of 93 papers on home range analysis usingradio-tracking data, and published in the 5-year period to the end of 1988, showedthat even 25 years after the first radio-tracking studies, in the majority of papersthere was still "insufficient attention given to accurate and sufficient datacollection, and to using appropriate analytical techniques to assess home range sizeand configuration." Less than 33% of the papers Harris et al. (1990) reviewedincluded an assessment of the accuracy of the radio-locations obtained (fixes),<10% considered whether or not locational fixes were autocorrelated, < 33%stated how many fixes were used to calculate home range size, and < 25%considered whether sufficient locations had been obtained for the home-rangeestimation to reach an asymptote.The number of radio-fixes required to estimate a home range size must beknown before the majority of field work is completed (Harris et al. 1990). This canbe accomplished by initiating a trial radio-tracking study period, where a largenumber of fixes can be collected from a range of animals in order to encompassvariations due to sex and age, and to determine at what point home range sizereaches an asymptote. This is determined by plotting home range size vs. numberof locations, and defined as the point after which additional locations result in aminimal increase in range size. Because asymptotes for individual home range-64-estimates are reached at different values and with a variety of curves depending onthe pattern of home range utilization and size, it is important that the selected timeinterval for the home range estimation is based on a sound assessment of theanimal's biology (Harris et al. 1990). In the present study I did not initiallydetermine the number of radio-fixes required; however, the data here presentedmay act as such information for any subsequent studies of the Strathcona ParkRoosevelt elk population.Swihart and Slade (1985) showed a bivariate test of independence usingempirically derived critical values for the ratio of mean squared distance betweensuccessive observations (t2) to mean squared distance from the centre of activity(r2). Using this, and 2 other tests of independence, Program Home Rangeautomatically determined whether or not each animal location dataset wasautocorrelated. The bivariate test of independence showed that over half of thedatasets were indeed autocorrelated in the present study. Harris at al. (1990)noted that strict adherence to the collection of non-autocorrelated data isworthwhile when the estimation of home range size is the primary aim of the studyand/or a large time interval between successive locations is possible. However,they concluded that most radio-tracking studies require the collection of data whichare dependent to some degree. So, even with a very large database, it may not bepossible to translate autocorrelated data into an independent form and still retain asample size that is adequate for home range estimation. This was the case in thepresent study, where it was not possible to translate autocorrelated data into-65-independent observations (see Swihart and Slade 1985, Samuel and Garton 1987,Ackerman ei al. 1990) and still retain a large enough sample size for home rangeestimation to occur.SUMMARYThe main purpose of my study was to provide the baseline ecological andpractical information that will allow Park managers to better understand the currentelk-habitat relationship in Strathcona Park before being able to manage for anothermore "desirable" one.Home ranges and core use areas were estimated using Program Home Rangefor all 10 radio-collared study animals. Location data for all study animals indicatedthat the two Heber River elk (#499 and #688) were migratory, while the other 8radio-collared elk were non-migratory.Population trend is one of the major indicators of elk response to habitatchanges (Janz g al. 1980a), and as such is important to understanding the presentwildlife-habitat situation, and for forming the context in which future managementdecisions can be made. Population and age-class composition estimates werepresented for each of the 4 study areas. Of concern is the low number of elk (4) inthe Thelwood valley.Brunt At al. (1989) estimated the home range sizes of migratory and non-migratory elk on Vancouver Island. Other studies (Graf 1955, Franklin etal. 1975)and reviews (e.g. Adams 1982) have reported a wide variation in home range size-66-within and among elk supspecies in western North America. Home range sizesestimated in the present study are within the variation reported in the literature;however, as Arcese (1989) observed, home range sizes may vary with populationsize, and therefore do not provide a universal scale for inter-specific comparisons.The different movement strategies used by the elk in the present study isconsistent with notion that migration is likely not a species characteristic of elk,but rather a versatile response to a given environmental situation.-67-CHAPTER 2Testing Seasonal Models of Roosevelt Elk Habitat SuitabilityINTRODUCTIONCentral to the study of wildlife ecology is how an animal uses itsenvironment; in particular the kinds of foods it consumes and the types of habitatsit occupies. Harris et al. (1990) noted that measuring an animal's home range size,shape, and pattern of utilization, are important for most ecological and/orbehavioral studies, and particularly for those concerned with habitat selection. Theability to accurately evaluate habitat and to predict wildlife habitat values frompreviously or readily gathered mapped data, would be of significant value to park,wildlife, and resource extraction managers. One of the first major steps takentoward such evaluative and predictive capabilities was the development of habitatsuitability models (U.S. Fish and Wildlife Service 1981). The ability of most modelsto provide accurate quantitative predictions is limited, but they are useful forunderstanding and evaluating how systems operate, and can facilitate thedevelopment of useful resource management plans (Bunnell 1989).Models of seasonal habitat suitability for Roosevelt elk in forestedwatersheds were first developed during phase I of the Integrated Wildlife-IntensiveForestry Research (IWIFR) Program (Brunt et al. 1989). These habitat suitabilitymodels were designed to provide a reliable means of assessing impacts of forestrydevelopment on potential Roosevelt elk habitat (Brunt 1991). The purpose oftesting the validity these models in my study, was to provide park, wildlife, and-68-forest managers with a reliable tool for assessing Strathcona Park for areas forgeneral elk habitat suitability, and for determining the impacts of recreational andindustrial development on elk habitat. In particular, this should assist parkmanagers in evaluating the land within Strathcona Provincial Park for the presenceof elk, for identifying areas suitable for the future placement of elk, and indetermining areas important to elk which require special protection or habitatenhancement.Johnson (1980) identified a 4-level ordering of the habitat selection process.First-order selection involves the physical or geographical range of a species. Thehome range of an individual animal or a social group within that range is determinedby second-order selection. Third-order selection refers to the usage made ofhabitat components within the home range, and fourth-order selection determinesthe actual procurement of food items from those available at that site.In this chapter, I examine seasonal range selection by a non-migratoryRoosevelt elk population in the Elk River valley of Strathcona Provincial Park(second-order selection) in relation to the relative suitability of the range's habitatcomponents (third-order selection).Habitat Suitability Index (HSI) Model Development and TestingThe habitat suitability index (HSI) models developed by the U.S. Fish andWildlife Service (U.S. Fish and Wildlife Service 1981) are based on assessment ofthe physical and biological attributes of habitat, under the assumption that habitatsuitability is proportional to carrying capacity (K. Berry 1986). Habitat use-69-documents a species' use of, or preference for, particular areas within its homerange. Two assumptions underlie this notion: i) an individual selects and usesareas that are best able to satisfy its life history requirements; and because of this,ii) greater use should occur in higher quality habitats (Schamberger and O'Neil1986). These 2 assumptions are not always valid, either because factors otherthan habitat characteristics may affect an animal's use of a site, or because of ourperception of that use.While there has been a recent increase in the number of wildlife habitatmodels being published in refereed journals, few models have been tested (K.Berry1986, Bunnell 1989). Most wildlife HSI models have been constructed using theliterature and opinions of professionals (K. Berry 1986). Because models aresimplifications of the systems, depicting and requiring numerous assumptions, theycan never completely mimic the real world (Maynard Smith 1974), and so areincomplete pictures of reality (Bunnell 1973).Bunnell (1989:6) made the important distinction between model validationand model verification, where "validate" means "sufficiently supported by actualfact," and "verify" means "to establish the truth, accuracy, or reality of." Thepresent study proposes to validate Brunt's summer season habitat suitabilitymodel. Lancia ?I al. (1982) considered habitat use by individual animals to be themost reliable method of model validation.According to Schamberger and O'Neil (1986), model testing serves 2important purposes: i) to provide information concerning model performance and-70-reliability; and ii) to provide data that may lead to model improvement. Beyondthese, model tests are intended to determine how well a model meets its statedobjectives. Models should be developed and tested using 2 or more different setsof the same type of data (Schamberger and O'Neil 1986). The present studyrepresents the second set of data used to validate Brunt's (1991) seasonal habitatsuitability models. The first testing of the validity of the models was presented inBrunt (1991). A third testing is currently underway on northern Vancouver Island(K. Campbell, unpubl. data).In testing a model, hypotheses can be formulated at different levels within amodel, including tests of assumptions, variables, components, or overall output.Most wildlife habitat model tests have been at the overall output level, whichprovides little information for improving model performance; testing individualmodel variables provides information for determining and improving model reliability(Schamberger and O'Neil 1986). This study was designed to test Brunt's (1991)summer season habitat suitability model at the overall output level.The primary means of testing the model was accomplished through linking aGeographic Information System (GIS) with Brunt's (1991) seasonal models.Throughout the process, Brunt's (1991) warning, which cautioned researchers toconstantly beware of the "black box" syndrome of computer modeling of ecologicalrelationships was heeded (see also Bunnell 1973, 1989).GIS Applications in Wildlife Habitat Modeling-71-Although the term GIS is relatively new, the use of spatial information formanagement decisions has existed since the first map was drawn. A GIS allowsthe efficient and useful organization, storage, retrieval, and display of informationfor management decisions (Devine and Field 1986). GIS technology is similar toconventional map processing, yet provides advanced analytic capabilities which canenable managers to address complex issues in entirely new ways (J. Berry 1986).GIS technology was introduced to park, wildlife, and forest managers as atool for storing, retrieving, and analyzing map and tabular information (Schwallerand Dealy 1986). The main purpose of a GIS is to process spatial information; itsmain power is that the relationships of the map data can be summarized (databaseinquiries), or manipulated (analytic processing). These procedures rely on thestorage abilities of GIS, which can efficiently organize and search large sets of datafor frequency statistics and coincidence among variables (J. Berry 1986). Existingmaps can be quickly and efficiently edited with GIS techniques. Predeterminedattributes can be entered into the GIS for each polygon (a defined area of particularimportance), independent of map creation, so that when complete, an attribute fileexists for every polygon (Consoletti 1986). Spatial information is representednumerically, rather than by analog means, such as in the inked lines of a map. Thisdigital representation has the potential for quantitative as well as qualitativeprocessing (J. Berry 1986).A key feature of this study is the importance of assessing habitatinterspersion. The use of a GIS allows the determination of spatial interspersion-72-values while retaining the spatial integrity of the habitat data. Eng et al. (1990)noted that previous attempts to incorporate habitat interspersion into wildlifeplanning have failed to adequately represent wildlife habitat because interspersionindices were added up over large areas. This has resulted in a failure to representthe relationships among individual habitat polygons.Study Background - Brunt's (1991) ThesisBrunt's (1991) study was based on the general movements and seasonalhabitat use of a transplanted group of migratory elk. A transplanted group usedprimarily for 3 reasons: i) habitat use by transplanted animals is more likely to bein response to existing habitat conditions than it is to long-established patterns; ii)for the first several years after the transplant, elk densities in the study area wouldbe lower than in most other Vancouver Island watersheds, thus minimizing thepossible confounding effects of density-dependent habitat selection patterns; andiii) a more precise test of model performance may be evaluated because exploratorymovements of the transplanted elk can be closely followed to assess which areaswithin the watershed the animals were or were not familiar with.The seasonal models of elk habitat suitability that Brunt (1991) developedand refined, were thus based on a low-density population of elk in an areacompletely new to the animals. Brunt's models may in fact primarily reflect (or beconfounded by) these 2 factors. In order to further validate the models, it isnecessary to test model performance and reliability with a group of elk-73-unconstrained by these factors; namely, a group of elk in a more densely populatedwatershed with which they are completely familiar.The specific objectives of Brunt's (1991) study were to develop modelsquantifying Roosevelt elk habitat suitability, and to test these models against elkseasonal range selection patterns in a different area from where they weredeveloped. The ultimate function of these models was to provide wildlife andforest managers with a reliable means of assessing the impacts of forestrydevelopment on elk habitat suitability (Brunt 1991).While habitats impose selection pressures which cause particular life historytraits to evolve, a complete description of the components of a habitat relevant tothe evolution of life histories would probably involve huge numbers of variables,and would become unwieldy for comparing different species (Lessells 1991).Habitat classification is an attempt to summarize the kinds of selection pressuresimposed by different habitats in just a few variables.The main factors affecting habitat suitability in Brunt's (1991) models areforage, cover, and the interspersion of forage and cover. Seasonal forage andcover quality for a polygon were calculated from information provided by theunderstory types, which are similar to ecosystem associations as described byKlinka gt al. (1984). A polygon's forage value was calculated from potential foragevalues modified according to overstory characteristics of the polygon.In testing Brunt's (1991) habitat suitability models, all potential cover andforage suitability values by understory type, and the modifier values of cover,-74-forage, and interspersion of cover and forage used by Brunt, were used here. I feltthat these values are based on sound reasoning and professional experience, andso assumed that they reflect the true (sensu Popper 1959) relative values of modelcomponents in each identified habitat polygon.Brunt (1991) developed and tested 3 models of seasonal habitat suitability(summer, mild winter, severe winter seasons) in the context of migratory Rooseveltelk habitat selection patterns on Vancouver Island. The question of seasonal rangeselection patterns of elk that do not migrate to higher elevations during the summerseason as a matter of habit was not addressed in the IWIFR program research inwhich the habitat relationships used in the models were initially developed. InBrunt's (1991) study, this question was only addressed in a recommendation forfurther study. Contrary to the importance placed on migratory elk in the IWIFRprogram and in Brunt's (1991) study, it is known that the majority of Roosevelt elkare non-migratory, as has been previously discussed in Chapter 1. The primary aimof my study was to validate the seasonal habitat suitability models, as applied tothe non-migratory Elk River valley elk population, and to determine which modelprovides the most accurate prediction of seasonal range selection patterns by non-migratory elk.The Elk River valley elk population was chosen as the study group for habitatmodel testing because: i) the area is within Strathcona Provincial Park and thus ofinterest to BC Parks; ii) funding was available for the required habitat mapping ofthe area; iii) a stable elk population was believed to be present; iv) the area was-75-relatively small and easily accessed; and v) the Elk River valley has somewhat lessdense vegetation than other areas inhabited by radio-collared elk, which facilitatedfrequent direct sightings of the study animals.STUDY AREAStudy AreaThe Elk River valley study area (Figure 1) is an approximately 4450 ha area,fallowing the course of the lower Elk River. The elevation gradient along the entireElk River changes dramatically from its upper valley headwaters (731 m atLandslide Lake) to its terminus (221 m at Upper Campbell Lake).The Elk River valley may be divided into 2 sections, the upper and lower.The lower valley currently begins at the confluence of the Elk River and the DrumLakes effluent (diverted water from the Heber River and Crest Creek; Kellerhals1992), and flows east into its terminus, Upper Campbell Lake. The abovedescribed lower Elk River valley is considered as the "study area" in the presentstudy. The upper valley, which flows into the lower, begins approximately 8.5 kmsouth of Drum Lakes, at Landslide Lake.Vegetation in the Elk River valley is characterized by the Coastal WesternHemlock zone (CWH), and above approximately 700-900 m elevation, theMountain Hemlock zone (MH), both as described by Krajina (1965). A detailedbiophysical description of the area can be found in Kojima and Krajina (1975). Themost recent description of habitat types along the Elk River valley can be found inRussell (1979).-76-Historical OverviewJanz and Lloyd (1977) noted that the major historical land use practicewithin Strathcona Park influencing present elk distribution is the logging of oldCrown-granted lands adjacent to the Elk River. They reported that the majority ofthe low elevation mature forest was logged during the early 1940's, from valleybottom (221 m elevation) to approximately 230-365 m elevation on the side hills.Although the entire Elk River drainage has been part of Strathcona ProvincialPark since its creation in 1911, the lower Elk River, in particular, has beenextensively modified by direct and indirect, primarily man-made interferences. Earlyphotographic records and written accounts dating back 50 years indicate a muchmore stable, productive, and aesthetically pleasing environment previously existed(Tredger et al. 1980). Kellerhals (1992) notes that the main impacts on the studyarea are thought to be:i) clear-cut logging of the flood plain (mid 1940's);ii) flooding due to a major landslide into a headwaters lake (1946);iii) road construction on the flood plains (mid 1940's - present);iv) flooding of the lowermost 10 km of river by BC Hydro's Strathcona Dam;v) complete diversion of Crest Creek into the Elk River; andvi) diversion of parts of the Heber River into the Elk River, which, togetherwith item 5 above, increases the low to intermediate water flows by 60-80%.The overall effects of these human-caused interferences has been a dramaticchange in river morphology along the lowermost alluvial reaches of the Elk River.-77-The active un-vegetated channel zone is now 2-3 times as wide as it was before1940, and this process of flood plain erosion is continuing (Kellerhals 1992).During this process, much fish and wildlife habitat has been irrevocably lost(Tredger et al. 1980).METHODSStudy Area Map DevelopmentUsing the Terrasoft 9c GIS (Digital Resources Inc., Nanaimo, B.C.), a digitalbase map of the Elk River valley was created from 1:20,000 understory associationmaps, 1:15,000 (approximate) air photographs, and the existing 1:50,000 series92F/13 - Upper Campbell Lake topographic map. Features digitized into the GISincluded biogeoclimatic subzones, understory association, canopy cover, aspectand elevation class, watercourses, roads, and the powerline transecting the studyarea.The 1:20,000 biogeoclimatic subzone and understory association map wascreated by Gartner Lee Ltd. (Burnaby, B.C.), under contract to the MOELP, and wasbased on the biogeoclimatic codes from Green et al. (1984) and understoryassociations described in Nyberg and Janz (1990). Mapped understoryassociations were delineated by the contractor from existing topographic and forestcover maps, color air photographs, and 40 field plots and ground truthing lines inthe study area. I then digitized the polygons delineating unique habitat types,based on biogeoclimatic subzone and understory type from the 1:20,000 map, intothe GIS, in addition to the above listed features.-78-Eight biogeoclimatic subzone units were delineated in the area, and 21unique habitats based on understory association types. In addition to the 40 fieldplots and ground truthing lines established by Gartner Lee, Ltd., I placed 10 - 10 mX 10 m field plots in different habitat types within the study area to further test theaccuracy of the understory association map. However, the limited number of fieldplots and the their highly biased placement relative to accessibility, cannot confirmthe accuracy of understory association map. Without more extensive verification,the map should be considered unconfirmed.Model Development and ApplicationThe main factors affecting habitat suitability in each of the 3 modelsdeveloped by Brunt (1991) are forage, cover, and the interspersion of forage andcover. The latter is important because elk is an ectone species whose habitat useis concentrated along the edge of more open areas which provide forage, and moredense areas which provide cover (Skovlin 1982). All 3 factors have long beenrecognized as the basic, seasonally fluctuating, habitat requirements whichdetermine elk abundance (Nyberg and Janz 1990). The model also considersaspect and elevation to be important determinants to winter habitat selectionbecause of their influence on snow accumulation on elk winter range (Brunt 1991),and are combined to form an additional factor used to calculate winter habitatsuitability in the mild and severe winter models.Brunt (1991) did not consider the location and density of roads to beimportant factors affecting habitat suitability in the 3 models. He believed that the-79-potential negative influences of the presence of roads, such as increased huntingpressures, may likely be balanced by the potential positive benefits, which includean abundance of preferred forage along roadsides, and the provision of relativelyeasy travel routes. The location and density of roads were not considered in mystudy in accordance with the above argument, and also because the 2 roads andpowerline which occur in the study area completely transact the valley, likelyaffecting the habitats within the entire area in a consistent manner.ForageForage values used in the model were based on understory types (Nybergand Janz 1990) modified by overstory conditions. Potential forage values byunderstory type for summer and winter seasons were developed based onVancouver Island Roosevelt elk diet information (Janz 1983, Brunt et al. 1989).These potential forage values were then modified depending on overstoryconditions; the underlying notion being that an increase in canopy closure results ina decrease in understory forage production (Nyberg and Janz 1990).Potential forage values ranged from 0 to 0.99, with 0.99 being optimal.Forage and cover values of 0.99 were used rather than 1.0 as optimal to avoiddivision by 0 in the calculation of overall habitat suitability, as discussed in detaillater under "Habitat Suitability Calculations." Forage modifier values are presentedin Table 8; potential forage values for the study area are presented in Table 9. ATable 8. Forage modifier valuesa.—^-Habitat ForageModifier Value4^/Logged 5 2 years previously 0.40Logged 3- 5 years previously 0.75Logged 5- 15 years previously 1.00Unlogged - deciduous dominated overstory b 0.75Unlogged - conifer dominated overstoryc 0.50Bog/Wetland; Rock outcrop; Slide complex 1.00From Brunt (1991)'Deer fern, Salmonberry, and Swordfern understory types°All other understory types with the exception of Bog/Wetland, Rock outcrop, andSlide complex-81-Table 9. Cover and potential forage suitability values by understory type'.Primary1^UnderstorySecondaryUnderstoryAb B C D- 0 0 0.99 0.90- 0.99 0.99 0.20 0.30Moss 0.99 0.99 0.18 0.26• Rock outcrop 0.99 0.99 0.22 0.28Salmonberry 0.99 0.99 0.32 0.40Slide complex 0.99 0.99 0.26 0.44Sword fern 0.99 0.99 0.32 0.38- 0.99 0.99 0.10 0.10• Rock outcrop 0.99 0.99 0.14 0.12• Dull Oregon grape 0.99 0.99 0.2 0.1- 0 0 0.30 0.20- i 0.99 0 0.30 0.20- 0.99 0 0.40 0.20Huckleberry-Moss 0.99 0 0.36 0.22• Lichen-salal 0.99 0 0.36 0.18Rock outcrop 0.99 0 0.38 0.20- 0.99 0 0.80 0.80- 0 0 0.50 0.99• - l 0.99 0.99 0.80 0.70• Salmonberry 0.99 0.99 0.80 0.72- 0 0 0 0'Modified from Brunt (1991)bA = Security Cover; B = Thermal/Snow Interception Cover; C= Potential WinterForage; D= Potential Summer Forage-82-more detailed explanation of the rationale used in potential forage and modifiercover value selection may be found in Brunt (1991).CoverNyberg and Janz (1990) discussed 3 types of cover as being important forelk: security, thermal, and snow interception cover. Only 2 types of cover areaddressed in the habitat suitability models, security cover, and thermal/snowinterception cover. Security cover is defined as vegetative or topographic covercapable of obscuring 90% of a standing adult elk at a distance of <61 m from ahuman observer (Thomas et al. 1979), although this relatively arbitrary definition isopen to question (Rahme 1991). Nyberg and Janz (1990) defined the standard forthermal cover in British Columbia coastal forests as stands > 10 m in height with amean canopy closure of > 70%; stands > 10 m in height with a mean canopyclosure of 60-90% provide snow interception cover for elk in coastal BritishColumbia forests. Habitat plots established in the study area (n = 50), examinationof air photos, consultation with a vegetation ecologist, and a detailed knowledge ofthe area, formed the basis for decisions on which habitats satisfied elk coverrequirements. Cover suitability values are presented in Table 9.Similar to forage values, understory types form the basis for the assignmentof cover values in the 3 models (Brunt 1991), however, habitat polygons are notdifferentiated by their relative cover value; they either do, or do not, satisfy coverrequirements. Brunt (1991) felt that there is insufficient information currentlyavailable to rank the relative ability of different habitats to satisfy elk cover-83-requirements, and that given our present level of understanding, this "all-or-nothing" approach is the most realistic (and conservative). Habitats were assignedvalues of either 0 or 0.99 for security and thermal/snow interception coversuitability values depending on whether or not they satisfied these coverrequirements.InterspersionInterspersion of forage and cover is recognized as a critical component ofwildlife habitat suitability (Nyberg and Janz 1990). Numerous studies (e.g. WitmerIn al. 1985, Hunter 1990; see also Nyberg and Janz 1990) have noted the declinein elk use of both forage and cover areas as the distance from their common edgeincreases. Although elk are an highly mobile species, their use of a particularhabitat is strongly influenced by the forage and cover characteristics of nearbyhabitats. Because of this, one half of the total habitat suitability of a particularpolygon is derived from its forage and cover characteristics, and the other halfdetermined from distance from cover and high quality forage (forage suitabilityvalue 0.5) (Brunt 1991),Edge-to-edge distances from areas of high forage suitability (z 0.5) and fromsecurity and thermal/snow interception cover were determined for the entire studyarea, and the resulting interspersion factor was determined for each model usingthe GIS' corridor analysis function.Interspersion modifier values for the various distances from cover and highforage value areas (Table 10) were developed by Brunt (1991) from distance-to-Table 10. Interspersion modifier valuesa.Distance from Cover or PreferredForage Areas (m)Modifier ValueOb 1.0<140 1.0141 - 249 0.6250 - 300 0.4>300 0.1/0.01c 1aFrom Brunt (1991)bSite qualifies as cover or preferred forage areac0.1 for cover; 0.01 for food-85-forage/cover relationships generated in previous elk/habitat research (Brunt et al.1989, Nyberg and Janz 1990). Modifier values assigned to distances > 300 mfrom cover or from high quality forage differed because Brunt (1991) considered elkdistribution to be more affected by extreme distances from high quality food thanfrom cover. He argued that cover availability was high throughout his study area,and because cover availability was also high in the present study area, I used thesame modifier values.Aspect/ElevationThe limiting factor to habitat suitability in mild winter and severe wintermodels is a combination of site aspect and elevation, which together influencesnowpack depth and persistence, both of which are critical components of elkwinter range suitability (Brunt 1991). Snow buries forage and increases the costsof locomotion at a time of the year when food availability is limited and energeticdemands are high relative to those in summer (Nyberg and Janz 1990).Aspect/elevation modifiers (Table 11) are used in the models because highelevation, north aspect areas receive and retain more snow than do lowerelevation, south aspect areas. These modifier values were developed based onrelationships among aspect and elevation and snowpack accumulation and meltrates "...from [Brunt's] general experience and observations, and from consultationwith other deer, elk, and habitat biologists working on Vancouver Island" (Brunt1991:56).Table 11. Aspect/elevation modifier values used in the mild and severe wintermodels'.Elevation(metres)Aspect290-700(north)71-1100(east)110-250°(south)250-2900(west)Flat0-350 0.6 0.8 1.0 0.8 1.0351-550 0.4 0.6 0.8 0.6 0.8551-1050 0.2 0.4 0.6 0.4 0.6>1050 0 0 0 0 0'From Brunt (1991)-87-Habitat Suitability CalculationsTo calculate a habitat suitability value for each polygon in the study area foreach of the 3 models, I followed Brunt's (1991) methods. Habitat polygon datawere digitized into the GIS in a vector (line) format, and subsequently gridded intoa raster format for overlay analyses and area calculations. Data levels containingforage, cover, interspersion, and aspect/elevation values were overlaid, resulting inthe formation of 492 polygons for the entire study area. Each resultant polygonwas assigned summer, mild winter, and severe winter habitat suitability valuesbased on the respective model's algorithm. Within the model algorithms, therelative weight values applied to the variables and component relationships used tocalculate seasonal range suitability "...were judgement based estimates of therelative contribution of the various factors to habitat suitability derived from 10years of elk-forestry studies on Vancouver Island" (Brunt 1991:58).The basic assumptions used in the development of these weighting valueswere the following (Brunt 1991):i) forage quality/availability is more important than cover during summer andmild winters compared to severe winters;ii) snow interception cover is not required on summer ranges and is lessimportant during mild winters than during severe winters; andiii) the influence of aspect/elevation on snowpack accumulation andpersistence is more important during severe winters than mild winters.The algorithms (Brunt 1991) used to calculate seasonal range habitatsuitability values were:-88-Summer Season Habitat Suitability:exp (0.5*ln (1-exp(0.7*In (1-SUMM) + 0.15*In (1-THERMAL) + (0.15*In (1-SECURITY)))) + 0.25*In (SUMMDIS) + 0.125*In (TDIS) + 0.125*In (SDIS))Mild Winter Season Habitat Suitability:exp (0.5*In (1-exp(0.7*In (1-WINT) + ((0.2*In (1-THERMAL)) + (0.1*In (1-SECURITY)))) + 0.25*In (WINTDIS) + 0.125*In (TDIS) + 0.125*In (SDIS))*ASEL• 5Severe Winter Season Habitat Suitability:exp (0.5*In (1-exp(0.5*In (1-WINT) + ((0.4*In (1-THERMAL)) + (0.1*In (1-SECURITY)))) + 0.20*In (WINTDIS) + 0.20*In (TDIS) + 0.1*In (SDIS))*ASEL L°Where:^In = natural logarithm;exp(x) = e'x';WINT = winter forage value;SUMM = summer forage value;THERMAL = thermal cover value;SECURITY =security cover value;WINTDIS =distance to winter forage of value ....0.5;SUMMDIS =distance to summer forage of value ....0.5;TDIS = distance to thermal/snow interception cover;SDIS = distance to security cover; andASEL = aspect/elevation modifier value.A more detailed explanation of algorithm development and choice of mathematicalcomputations is presented in Brunt (1991:56-61).Home Range Suitability CalculationsThe cumulative ranges and core use areas generated from the home rangeprogram were imported into the GIS to generate habitat suitability values for theranges. The seasonal suitability values of the individual polygons within the rangeswere weighted by the area of the polygon in question relative to the total Elk Rivervalley study area. The sums of the weighed habitat suitability values representedthe overall suitability of the range.-89-Analyses of Modeled Habitat SuitabilityThree main analyses were conducted on the habitat suitability datagenerated from application of the 3 seasonal models. Where appropriate, thehabitat suitability of individual animal locations and/or the habitat suitability ofanimal home ranges generated by the home range program from the animal locationdata, were used for model validation tests.The 3 validation tests of the habitat suitability models' output were thefollowing:A) testing for random habitat selection with respect to modeled habitatsuitability;B) examining modeled habitat suitability of the total study area, unusedportions of the study area, cumulative range, and core use areas; andC) assessing which of the 3 seasonal habitat suitability models best reflectsthe range selection of non-migratory elk.A) Random Habitat Selection TestBrunt's (1991) habitat suitability models would have no predictive capabilityif the elk were selecting habitats at random with respect to modeled habitatsuitability. In order to provide "statistical evidence" that no random habitatselection occurred within the study area, a chi-square goodness-of-fit test wasapplied to the summer season animal location data. Because it was immediatelyobvious that random habitat selection was not occurring, and because modeledmild winter and severe winter habitat suitability values for the animal location datawere very similar to those of the summer location data, goodness-of-fit tests werenot applied to the 2 winter models.-90-Four classes of habitat suitability were defined in an attempt to follow thesuggestions made by Cochran (1954:417) that no expected frequency should beless than 1.0, and no more than 20% of the expected frequencies should be lessthan 5.0. Expected frequencies were determined by calculating the proportion ofthe entire study area having habitat suitability within the 4 classes chosenmultiplied by the number of locations obtained (n = 216) for the summer seasonanimal location data. Observed frequencies were identified as the number ofanimal locations occurring within each of the 4 suitability classes.B) Habitat Suitability Within Study AreaIf the 3 seasonal habitat suitability models accurately reflect elk habitatselection patterns, areas of higher use should have higher suitability values. Theprinciple validation test of the models' ability to predict elk seasonal range selectionwas to compare the habitat suitability of the following 4 calculated regions: thetotal study area; portions of the total study area unused by elk; the cumulative elkrange within the study area; and the core use area within the cumulative elk range.All 3 seasonal range suitability models were applied to each of the 4 rangetypes. The seasonal suitability values of the individual polygons within each of the4 regions were weighted by the relative area that the polygon represented of thetotal area of the region. The sums of these weighted values represent overallsuitability of the 4 regions.The total study area was somewhat arbitrarily delineated west-to-east by themaximum height of the Elk River valley watershed at the Drum Lakes, to Upper-91-Campbell Lake, and north-to-south by the approximate height of land of the studyarea. The cumulative range and core use area were estimated by Program HomeRange (Ackerman Dt DI. 1990) using all animal locations (n = 284). The portion ofthe study area unused by elk was determined by repeated ground and aerialsurveys of those regions of the total study area exclusive of the cumulative rangethroughout the season of interest. Although Janz and Lloyd (1977) reported thatthese areas supported a very small group of elk during more severe winter seasons,no elk sign could be found during the study period, and I feel confident that theseareas did not support any elk during the study period.Although comparisons of total study area, unused total area, cumulativerange, and core use area habitat suitability values cannot be accomplished in astatistically rigorous fashion (essentially because there is only one of each rangetype available for comparison with the other types), if the models adequatelypredict habitat selection, a trend should be evident, indicating habitat suitability tobe proportional to the elk's use of an area.A more rigorous statistical analysis of the habitat suitability values betweenanimal locations (n = 284) in the intensively used (core use area), and 284 randomlygenerated locations in the unused portions of the study area was accomplished byusing a one-tailed t-test. Habitat suitability was weighted by the area of thepolygon relative to the area of the core use area in which all of the animal locationsfell. Each animal location and randomly generated location was given the habitatsuitability value of the polygon in which it fell, for each of the 3 models. Means-92-and standard deviations of the habitat suitability values were then calculated foreach model. If the models adequately predict habitat selection, the mean habitatsuitability value of the core use area animal locations should be significantly higherthan that of the randomly generated locations in the unused portion of the studyarea.C) Comparisons Among Seasonal Model PredictionAs discussed earlier, if the models accurately reflect elk habitat selection,areas of higher use should have higher suitability values. But because the Elk Riverelk are non-migratory, it is expected that at least 2 of the models (summer andwinter) should be equally good at predicting year-round habitat selection; no onemodel should predict higher suitability values for areas of higher use.In order to determine if there is any difference in the abilities of the 3 modelsto predict habitat selection for a non-migratory elk population, habitat suitabilityvalues generated by all 3 models were compared using animal location data.Habitat suitability were weighted by the area of the polygon relative to the area ofthe core use area in which all of the animal locations fell. Each animal location(n = 284) was given the habitat suitability value of the polygon in which it fell, foreach of the 3 models. Means and standard deviations of the habitat suitabilityvalues were then calculated for each model.D'Agostino's test for normality (D'Agostino 1971 a, b) indicated that thedistribution of habitat suitability values for the animal location data was non-normal(D = 0.191495, critical values = 0.2705, 0.2866, a = 0.05). However, because-93-n = 284 was large enough to approximate a normal distribution, two-tailed paired t-tests were performed to determine statistical difference between mean habitatsuitability value output of each model (A. Kozak; pers. comm.).RESULTSHabitat Suitability Model Validation TestsFigure 7 presents the Elk River valley study animals' (elk #'s 120, 179, 218,and 259) cumulative range and core use area within the total study area asestimated by Program Home Range (harmonic mean estimator).A) Random Habitat Selection TestSummer habitat suitability values of locations used by elk in summer weresignificantly different than expected values based on availability (Table 12); elkused habitats in summer with the highest suitability values ( > 0.90) more oftenthan expected. This indicates non-random habitat selection with respect tosummer model output, and suggests that further analyses to test model validity arewarranted.B) Habitat Suitability Within Study AreaA consistent trend of increasing habitat suitability value is obvious in all 3models from the unused portion of the study area, to the total study area, to thecumulative range, to the core use areas (Table 13).Mean habitat suitability of animal locations (n = 284) in the core use areawas significantly higher than that of randomly generated locations (n = 284) in the0^ 2.5^5 kmTOTAL STUDY AREA ^CUMULATIVE RANGECORE USE AREA UNUSED PORTION OFTOTAL STUDY AREA-- — - - Logging Road___ _ _   Highway 28..—....„-------------._,„,-----,Figure 7. Total study area, unused portion of the study area, cumulative range, and core use area for elk #'s 120, 179, 218, and 259in the Elk River Valley, as delineated for seasonal habitat suitability model testing.Table 12. Frequencies of expected (given random habitat selection) and observedhabitat suitability class values for the summer model.Summer ModelSuitability ClassSummer' Animal Locations (n = 216)"Observed Expected'<0.60^t 24 36.120.60 - 0.87 0 79.540.87 - 0.90  60 62.71>0.90  132 37.64Chi-square = 320.28; critical value = 7.815; a = 0.05;degrees of freedom = 3aSummer = May 1, 1992 - September 15, 1992'Proportion of the study area in the suitability class multiplied by 216 locationsTable 13. Habitat suitability values for unused portion of the total study area, totalstudy area, cumulative range, and core use area for summer, mild winter, andsevere winter models.ModelTotal StudyAreaUnused Portion ofTotal Study AreaCumulativeRangeCore UseAreaSummer  0.76 0.71 0.81 0.88Mild Winter 0.73 0.67 0.77 0.86Severe Winter 0.76 0.71 0.81 0.89-97-unused portion of the study area (Table 14), for all 3 models (summer t =10.551,mild winter t = 9.056, severe winter t= 9.736; for all tests tcriti„, = 1.645, df = 566,a =0.05).C) Comparisons Among Seasonal Model OutputThe mean habitat suitability values of summer, winter, and cumulative(summer and winter) animal location datasets calculated by all 3 models showedlittle variation (Table 15). Of the 9 comparisons possible between mean habitatsuitability values of summer, winter, and cumulative animal location datasets foreach of the 3 models (Table 16), only that between summer and winter valuesfrom the mild winter model, was significantly different (t = 2.180, t-critical = 1.960,df = 282, a = 0.05) (Table 16).DISCUSSIONBrunt's (1991) models are models of understanding; they represent a firststep towards the development of a predictive tool capable of making realistic,quantitative predictions about the response of elk populations to habitatmanagement activities. While the results of my study validate the models'simplification of the real-world elk-habitat relationship, one must keep in mindBunnell's (1989:8) caution that "validity and simplicity are not synonyms ofveracity," and that although a researcher can quantify the accuracy of apredictive model for a manager, he cannot quantify the degree to which the modelhas been strengthened.Table 14. Mean habitat suitability values for animal locations in the core use area,and random locations in the unused portion, of the Elk River valley study area.ModelMean Habitat Suitability Value ofCore Use AreaAnimal Locations(n= 284)Unused Portion of StudyArea Random Locations(n= 284)Summer 0.87^(0.15)a 0.66 (0.30)Mild Winter 0.87 (0.17) 0.68^(0.31)Severe Winter 0.89^(0.19) 0.67 (0.33)"Sample standard deviationTable 15. Mean habitat suitability values of summera, winter b and cumulativeanimal locations estimated by each of the 3 seasonal models.ModelERV Group`Summer Locations(n= 216)Winter Locations(n= 68)Cumulative Locations(n= 284)Summer 0.88^(0.14)d 0.84 (0.17) 0.87^(0.15)Mild Winter 0.89^(0.16) 0.84 (0.18) 0.87^(0.17)Severe Winter^0.90 (0.18) 0.86 (0.22) 0.89 (0.19)'May 1, 1992 - September 15, 1992bJanuary 11, 1992 - April 31, 1992; September 16, 1992 - January 6, 1993`Includes elk #'s 120, 179, 218, and 259dSample standard deviation-100-Table 16. Results of all possible two-tailed paired Mesta comparisons amongsummerb , winter, and cumulative d animal location datasets using the 3 habitatsuitability models.Model Used Comparisons BetweenAnimal Location Datasetst-valuesummer and winter + 1.948Summer summer and cumulative + 0.760winter and cumulative -1.443summer and winter + 2.180Mild Wintersummer and cumulative + 1.336winter and cumulative -1.292summer and winter +1.512Severe Wintersummer and cumulative +0.596) winter and cumulative -1.133aln all cases a = 0.05, and t-critical =^1.960bSummer = May 1 - September 15, 1992'Winter = January 11 - April 30, 1992; and September 16, 1992 - January 6, 1993dSummer and winter animal location datasets combined-101-The models of seasonal habitat suitability tested in the present study werevalidated in that they appear to reflect elk range selection patterns. While no trueseasonal ranges were selected by the non-migratory Elk River valley elk population,modeled suitability of the habitats selected by elk were significantly different fromwhat would be expected if selection had been random. This was the case for all 3models, using both the animal location data and the home ranges estimated fromthose location data.For all 3 models, validation tests further indicated that the cumulative elkranges were of significantly greater suitability value that the total study area and,as a consequence, those portions of the total study area unused by elk. Inaddition, the elk core use areas had significantly greater suitability values than thecumulative ranges. Core use areas are especially important because they denoteareas of particularly high home range usage, and thus they often may provide amore clear measure of the changing pattern of range use than does total homerange area. Core use areas are often more useful than the more peripheralcontours delineating an animal's home range for understanding both intraspecificand interspecific patterns of home range use (Harris et al. 1990). As expected, theanimal locations within the core use area had the greatest mean habitat suitabilityvalues within the entire study area.The relative increase in predicted habitat suitability values from unused tocore use areas indicates the ability of the models tested in this study to reflect elkhabitat selection patterns. The assumption being that elk select cumulative areas-102-to satisfy certain requirements from all areas available to them (the total studyarea =the lower Elk River valley), and that within the cumulative range, the elk "keyin" on core use areas of particularly high habitat suitability. Similar consistenttrends in Brunt's (1991:132) study led him to likewise conclude that they provided"...what I consider the best corroborative evidence" of the models' validity.Habitat suitability values for both home range and animal location datasets inthe Elk River valley were notably higher than those calculated in Brunt's (1991)study of migratory elk, where cumulative range habitat suitability values rangedfrom 0.27-0.55; seasonal range values from 0.36-0.74; and core use area valuesfrom 0.24-0.85. The difference in habitat suitabilities calculated in my study andBrunt's (1991) is probably related to the fact that Brunt's study animals weretransplanted into a watershed with which they were unfamiliar, and consequentlyunaware of all areas of high habitat suitability value. In addition, Brunt's studyanimals were all migratory. One of the reasons the Elk River valley elk are non-migratory may be related to the higher habitat suitability values found in the areathey occupied.In the present study, no one model best predicted the habitat suitability ofthe home range selected by the non-migratory study animals. Studying migratoryelk, Brunt (1991) found that the 2 winter models predicted statistically highersuitability values for winter ranges than for summer ranges, as was expected.However, there was no statistical difference between the summer model'spredicted suitability value for the summer and winter ranges. Brunt postulated that-103-the winter ranges within his study area could provide areas of similar suitability tothose occupied during the summer (winter ranges had the highest suitabilitypredicted by each model), but elk introduced to the study area continue to migrateout of habit. In support of this concept that elk movement behavior becomesrelatively fixed and unchanged despite translocation, Brunt (1991) noted that inprevious transplants of non-migratory elk on Vancouver Island, the animalsremained non-migratory even though high quality summer range was available athigher elevations. Likewise, migratory elk may continue to migrate even whenadequate summer range is available without migrating.In the present study I found that the 3 models showed little variation incalculated habitat suitability values using summer, winter, and cumulative animallocation datasets. There was no statistical difference between model outputsexcept between summer and winter using the mild winter model. In view of theother 8 comparisons for which no differences exist (Table 16), I believe thedifference between the summer and winter model is unimportant, and that there isvirtually no difference in habitat suitability values predicted by the 3 models.Russell (1979) concluded that the amount of available forage in the Elk Rivervalley is the greatest limiting factor on the number of elk using this area. Inaddition to forage as a limiting factor, the vast majority, if not the entirety, of thestudy area has been significantly and routinely disturbed from its pre-colonial state(Tredger et al. 1980, Kellerhals 1992). Unquestionably, this must have hadnumerous negative effects on local fish and wildlife populations, including on the-104-Roosevelt elk. Two potential threats particular to the elk in the Elk River valleyinclude illegal hunting and accidental road kill, both of which are facilitated by the 2roads running through the area. In spite of these threats, I agree with Brunt's(1991) recommendation that the location and density of roads need not beincorporated into the models. Illegal hunting and accidental road kill areunpredictable factors, and thus difficult to incorporate as variables of habitatsuitability models on Vancouver Island.Eng et al. (1990) noted that Vancouver Island exhibits a cyclical pattern ofsnow depths; severe winters occur approximately every 18 years. Thus, long termmaintenance of elk populations requires the availability of both mild and severewinter habitats. Because the winter season during the study period providedconditions under which the mild winter model might be applied (MOELP; pers.comm.), the severe winter model remains untested with respect to both migratoryand non-migratory elk populations. Brunt (1991) noted that since the initiation ofthe IWIFR research project which provided the majority of information for thedevelopment of the models, winter weather conditions on Vancouver Island havebeen relatively mild. Until the habitat relationships that drive the severe wintermodel can be tested during winters of prolonged, deep snow accumulations, thesevere winter model cannot be considered more than preliminary, and as Brunt(1991:133) cautioned, "...applied with prudence."Brunt (1991) recommended conducting further model validation tests indifferent areas during years of different weather conditions; this provides the-105-necessary context for more intensive research in the future. As discussed earlier,most tests of wildlife-habitat models, including Brunt's (1991), the present study,and the ongoing study on northern Vancouver Island, examine only model output.Schamberger and O'Neil (1986) noted that additional tests of the individual modelvariables or assumptions of the model provide information for determining andimproving model reliability. Intensive research into the individual modelcomponents and relationships would be the next logical step in further assessmentsof the validity of the models, and to improve their predictive capabilities (Brunt1991). This is particularly appropriate in light of the findings of the present study,in terms of the extent to which the models apply to the habitat selection patternsof non-migratory Roosevelt elk. Brunt (1991) noted that previous transplants ofnon-migratory elk on Vancouver Island to an area where higher elevation summerrange was available did not result in the animals adopting migratory behavior totake advantage of these areas. Further model validation tests using such atransplanted population may provide additional insight into the dynamics of non-migratory elk habitat selection patterns.CONCLUSIONSAfter testing and validating the 3 habitat suitability models, Brunt (1991) stillconcluded that the models may not be embraced by forest and wildlife managers.The principle reason for possible reluctance to make extensive use of the modelscenters on the understory mapping which is necessary for determination of both-106-forage and cover suitability indices used in the models. Unfortunately, informationto generate an understory map is not readily available from existing map bases. Anextensive background is required in vegetation ecology, expertise in air photointerpretation, and extensive ground checking to prepare a validated understorymap (Brunt 1991). This will likely prohibit ready acceptance of the models intogeneral forest resource and wildlife-habitat planning.Bunnell (1989) considered 4 ways by which modeling can waste time,talent, and funding. They include misuse, where models are used for the wrongpurpose; disuse, where models are no longer used; lack of model evaluation; andlegal challenges to the validity of the model and its use in making managementdecisions. The models tested in the present study are most likely to fall victim tothe second fate: disuse.Disuse of a model is obviously wasteful if developing the model wasexpensive in terms of time and money, and the model is not used. Bunnell(1989:2) noted 4 broad, related reasons why wasteful disuse of models occurs: i)the model does not address the question or purpose of the intended user; ii) theintended user never existed, "just someone who might be convinced"; iii) althoughit asks the right question, the model is too complex to be used and/or requires toomuch data; and iv) model output is not sufficiently accurate. At the present timeand state of model accuracy, all 4 reasons apply to BC Parks potential disuse of themodels.-107-As was previously discussed, the primary purpose of initially developing themodels was to provide forest managers with a reliable means of assessing theimpacts of forestry development on elk habitat suitability. However, it has beensuggested that BC Parks might apply the models as a management tool with aslightly different goal: to make predictive use of the models in terms of identifyinghigh quality elk habitat throughout and adjacent to Park lands. Such identifiedareas would then be targeted for a particular management action such as specialprotection, reduced human access, transplant activities, etc.I believe that the cost-benefit ratio for this use of the model is notwarranted, given the time and money required to collect the information necessaryto generate an understory map of the entire Strathcona Park. A systematicidentification of high quality elk habitat in the Park by qualified elk-habitat expertscould be accomplished in less time, for far less money, and would not necessitatefurther ground verification. However, in addition to identifying high quality elkhabitat, the baseline data gathered in the completion of a Park-specific understorymap would allow for future studies of other species wildlife-habitat relationships.This study provides additional support for a correlation between predictedhabitat suitability and elk habitat selection. Like Brunt's (1991) initial study ofmodel development and testing, it is, in effect, an unreplicated experiment with nocontrols. Further research in watersheds throughout Vancouver Island and on non-migratory as well as migratory elk is needed to gain confidence in the results from-108-application of the models. The models still must not in themselves be consideredas decision-making tools.Romesburg (1981) stated that modeling was never intended to function as ameans to scientific knowledge, and that the use of modeling in science is limitedbecause it cannot predict to within established tolerances. However, its continualuse is assured as a planning tool that can integrate scientific knowledge andcommon sense, as well as theory, hunches, and expert opinion to forecastalternative future images: "Planning is inaccurate by the standards of science, butthat is no reason to abandon planning," (Romesburg 1981:310). 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