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Factors influencing summer habitat use of black-tailed deer on South-Central Vancouver Island Morgan, Jeff 1994-02-25

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Factors Influencing Summer Habitat Use of Black-tailedDeer on South-Central Vancouver IslandByJEFFREY ALLEN MORGANB.Sc., Simon Fraser University. 1988A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTSFOR THE DEGREE OF MASTER OF SCIENCEinTHE FACULTY OF GRADUATE STUDIES(Departmeni of Forest Sciences)We accept this thesis as conformingto the required standardTHE UNIVERSITY OF BRITISH COLUMBIAAPRIL 1994©Jeffrey Allen MorganIn presenting this thesis in partial fulfilmentof 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 forextensivecopying of this thesis forscholarly purposes may be granted bythe head of mydepartment or by his or her representatives.It is understood thatcopying orpublication of this thesis for financial gain shallnot be allowed without my writtenpermission.Department of__________________The University of British ColumbiaVancouver, CanadaDate________________DE-6 (2/88)IIAbstractUse of seral stage, biogeoclfrnatic variant, aspect and elevation by 13radio collared black-tailed deer (Odocoileus hemionus columbianus Richardson)was examined at the ‘within home range’ level of habitat selection. Pooleddata sets were evaluated for patterns of habitat use in relation to timeperiods within the summer of 1991. The July/August period of peak clearcutuse coincided with the period of peak fireweed (Epilobium angustfolium L.) useidentified by other research. The percentage of locations within clearcutswas 67.9% in April/May/June, 78.2% in July/August and 69.2% inSeptember/October. Fireweed, an early successional forb, has been shownby other research to be the most important black-tailed deer summer foragespecies. Similar trends in fireweed use and the percent of locations withinclearcuts therefore, were predictable.The use of Montane variants of biogeoclimatic subzones was higher inthe latter portions of the summer. The percentage of locations withinMontane variants was 14.5% in Aril/May, 22.6% in June/July and 25.8% inAugust/September. Use of Montane habitat was lowest in the spring whenforage species within the Submontane were more phenologically advancedand more nutritious. Shifts in elevation and changes in the use of aspectwere not detected.Parturient deer did not change their use of seral stage within thefawning period. Home range size (minimum convex polygon) was greater inthe fawning period than in any other period. Increases in home range sizeduring the fawning period were caused by increases in the magnitudeand/or rate of movements to the peripheries of the home ranges. Thisfinding supports the idea that sites on the periphery of the dam’s home rangeare used for fawning, and is consistent with the ‘hider strategy’ model. Hidersliiare known to remove their young from the birth site to preventpredatorsfrom using olfactory cues to detect their young. If black-taileddeer were togive birth within their core areas of use, they would be forced to removetheirfawns from the habitats they mostIDrefer.Home range size was smaller in the immediate post-fawningperiodthan in all other periods except the pre-natal period. This behavioris againconsistent with the hider strategy. Immediately after fawning, the dammustremain close to her fawns, which are sedentary. Habitat use in theearlysummer was more concentrated in Submontane habitats wheregreen-upoccured first. Home range size in the pre-natal period therefore, should alsohave been small.ivTable of ContentsAbstractList of Tables viList of Figures viiAcknowledgments viiiChapter 1: General Introduction 1Rationale and Objectives 1Forage quality 1Habitat use during fawning season 3Study Area 4General Methods and Materials8Description of Study Animals 11Chapter 2:Changes In Habitat Use In Relation toForage Quality 14Introduction 14Seral stage 14Delayed phenology 17Methods and Materials 18Results 21Seral stage 22Delayed phenology 24Potential confounding variables 27Discussion 30Seral stage 30Delayed phenology 32Chapter 3: Changes in Habitat Use by FemaleBlack-tailed Deer During the Fawnihg Season 34Introduction 34Methods and Materials 37Results 39Habitat use 39Fawning site selection 39Home range size 46Discussion 50Habitat use 50Areaof use 51VChapter 4: Summary and Conclusions...........54Changes in Habitat Use in Relaticn to Forage Quality 54Changes in Habitat Use During the Fawning season 55Recommendations and Implications 56Literature Cited 58List of TablesTable 1.1. General information on study animals. 1989-199 1 ...12Table 2.1. Data collection periods for collared deer, 1991 19Table 2.2. Number of locations per deer per month, 1991 22Table 2.3. Number of locations within clearcut andforested habitats and percent of locations withinclearcut habitats,1991 23Table 2.4. Number of locations within Montane andSubmontane habitats and percent of locations withinMontane habitats, 1991 25Table 2.5. Percent clearcut habitat within the Montaneand Submontane variants within the 95% MCP homeranges of study animals 17701 and 18181 29Table 2.6. Percent use comparison of Montane andclearcut habitats for study animal 17701 29Table 3.1. Home range size (hectares) by time period 47Table 3.2. Paired difference test comparison of homerange size during the fawning period versus allother periods 49Table 3.3. Paired difference test comparison of homerange size during the immediate post-fawningperiod versus all other periods 50vilList of FiguresFigure 1.1. Location and major subdrainages of theCaycuse Study Area6Figure 2.1. Monthly pattern of use by black-taileddeerof forage species in forested and cutover areas(Rochelle 1980. p.64) 16Figure 2.2. Percent of pooled locations withinclearcuts, 199124Figure 2.3. Percent of pooled locations withinMontane, 199126Figure 3.1. The 1989 fawning movements of study animal1818141Figure 3.2. The 1990 fawning movements of studyanimal1818142Figure 3.3. The 1990 fawning movements of study animal1320143Figure 3.4. The 1991 fawning movements of study animal17991 44Figure 3.5. The 1991 fawning movements of studyanimal12801 45Figure 3.6. Mean summer home rapge size and standarderrorfor 10 female deer in five periodsduring the summerof 199148viiiAcknowledgmentsI am most grateful to my wire,Beth who was foolish enoughto marryme shortly after I had begun mygraduate studies. Her emotionalandfinancial support saw me through challengingtimes. My mother and father,Art and Lois, supported me likewise.Thanks also to my sister Shelleywho, bydemonstrating that university bouldbe an enjoyableexperience,encouraged me to advance. Otherpositive influences were DaveDunbar,Adam Lewis and Ken Budd. Manyother friends and family membershavesupported me at different levels overthe past few years. Theirhelp andpatience was appreciated.Dr. F.L. Bunnell supervised mypost graduate studies and research.Hiscommittment to science andecucation ensured that Iwas challenged.Thanks also to my committeemembers, Dr. M. Pitt, Dr. T. Sullivanand Dr. A.Derocher who helped fashionthe study design and/or providedvaluablecriticism. My ‘operational supeMsor,Scott McNay, arrangedlogistic,financial, and technical assistanceand was a constant soundingboard.Technicians who assisted inthis research were Janet Windborne,Dan Bate,Yves Giguere, Line Giguere andWayne Wall. Othermembers of theVancouver Island, wildlife biologycommunity that helped withthis researchwere Marvin Eng, Dr. R. Page, JoanVoller, Myke Chutter, DonDoyle, KimBrunt and Doug Janz. LesPeterson deserves specialthanks. Were it not forhis altruistic nature and analyticalabilities, I might still be writingup. Under histuition I learned many ofthe skills required to completethis thesis. DerekWilkinson provided timely assistancewith the the figures. JackieJohnson,office manager of the unmanageableHeader House, kept myacademiccareer in order, proof read manuscriptsand was a friend. Thanksalso to theHeader House folks fortheir encouragement, assistance,criticism andfriendship.This project was funded by a B.C.Ministry of Forests/ForestryCanada-South Moresby Forest ReplacementAgreement grant tothe ResearchBranch of the B.C. Ministry ofForests. Fletcher ChallengeCanada Ltd.,Integrated Resource Analysis Section,contributed finances,equipment andlogistical support. The B.C. Ministryof EnvironmentLands and Parks and TheB.C. Ministry of Forests(Integratec WildlifeIntensive Forestry Research)alsocontributed equipmentand logistical support. Mypersonal funding wasprovided by the Science Counciiof British Columbia througha G.R.E.A.T.award to Fred Bunnell. BobWillington, of FletcherChallenge Canada Ltd.,graciously cooperated in thearrangement of this funding.1Chapter One: ‘General IntroductionRationale and ObjectivesForage QualityIn the Temperate and Arctic zones of the Northern Hemisphere,availability of high quality winter range is often considered to be the mostimportant factor limiting conditioh and size of ungulate herds (Connolly1981). Consequently, the habitat ecology of ungulates during other periodsof the year has received relatively little attention. When habitat use andrange characteristics during the summer are documented, it is often doneonly to provide a comparison between winter and summer ranges.In many instances winter range quality is indeed the most importantfactor limiting herd productivity; there are, however, pitfalls in viewing thisconstraint in isolation of others. Although mortalities resulting frommalnutrition usually occur in late winter or early spring, they are a function ofboth winter forage availability and pre-winter conditioning (Short 1981). Thepre-winter conditioning that occurs while the animals are on their summerranges is often taken for granted. In many carrying capacity models, (e.g.,Errington 1945; Potvin and Huot 1 9S3; Bartmann et al. 1992), the quality of thesummer range is not considered. It is assumed that during the summer,ungulates have access to surplus quantities of forage. It follows, that theyenter winter in peak condition and that, with respect to over-winter mortality,the quality of the summer range is inconsequential.There is evidence that suggests this is not always the case. In aridregions, mule deer (Odocoileus hemionus hemionus) populations have beenfound to be limited by the quality of their summer ranges (Longhurst et al.1952; Russo 1964; Robinette et al. 1977; Smith and LeCount 1979).2In moister regions gradual migrations of ungulates suggest individualsèonstantly seek out fresh, high quality summer pastures to improve theircondition. Where such migrations occur, the spatial arrangement of differenthabitat types over the landscape, and the strategies that ungulates use toexploit them, are probably important factors affecting carrying capacity.Carrying capacity is defined as “the number of animals that a habitatmaintains in a healthy and vigorous condition” (Dasmann 1981: 151).Large herbivores such as mule deer, Rocky Mountain elk (Cervus elaphusnelsoni) and moose (Alces alces) have been observed to engage in gradualelevational migrations during summer months (Altmann 1952; Edwards andRitcey 1956; leResche 1974). Some authors have found mule deertoundertake such movements in response to plant phenology (Russell 1932;Leopold et al. 1951). Bunnell (1990: 61) observed that “migratory deer thatmove to alpine and subalpine areas as the season progresses are able toprofit from the higher quality forage found there during the late summerandfall”.Because most of the black-tailed deer (Odocoileus hemionus columbianusRichardson) research in the Pacific Northwest has focused on winter range,relatively little is known of the species’ summer habitat ecology. Inthemountainous habitats of this region, black-tailed deer populations usuallyhave a component of elevational migrators (Harestad 1979; Loft et al. 1984;Schoen and Kirchhoff 1985; McNay and Doyle 1987). Although differences inelevation, aspect, and vegetation types between summer and winter rangesare often presented, the data sets are seldom scrutinized for gradualchanges in habitat use that occur within the season. Seasonal home rangesare depicted as static entities Id which constant foraging strategies areemployed. Our knowledge of broad seasonal habitat requirementsis3extensive, but potential changes in habitat use in response to changes inforage quality within a season are not well understood. During the growingseason, black-tailed deer probably make constant adjustments to theirforaging strategies to maintain quality in their diet. Appropriate foragingstrategies could entail adjustments to the position of the home range inresponse to changes in vegetation quality (e.g., plant phenology).Alternatively deer might adjust their habitat use patterns within static homeranges.The primary objective of Chapter Two is to report the use of habitattypes within the summer range. A secondary objective, when appropriate, isto relate the changes in habitat use to gradual elevational migrations orrange expansions.Habitat Use During Fawning SeasonOne aspect of ungulate summer habitat use that has receivedconsiderable attention is birth site selection. Ungulate mortality rates areusually highest during the first few weeks after birth (Cook et al. 1971; Jacksonet al. 1972; Smith and LeCount 1979; Steigers and Fllnders1980; Ozoga andVerme 1986). Neonate mortality from predation is sometimes a major limitingfactor. Hatter (1988) estimated fawn mortality, within a northern VancouverIsland black-tailed deer herd, to be 63%, and concluded that wolf (Canislupus) predation was the primary factor limiting deer recruitment.Often, specific habitat types that confer anti-predator advantages tothe neonate and the mother are used during calving or fawning seasons.Steep, broken and rocky sites are often selected as natal sites by bighornsheep (Ovis canadensis) (Festa-Bianchet 1988). Rocky Mountain elk neonatesused sites with dense brush cover (Johnson 1951; Altmann 1952). In Utah,mule deer avoided giving birth ir sites with dry exposures and poor cover4(Robinette et al. 1977). Moose, caribou (Rangifer tarandus),and mule deerhave been found using small islands as birthing sites apparently toreducethe risk of predation (Steigers and 9inders 1980; Edwards 1983;Bergerud andPage 1987).Other authors have associated increasedintraspecific avoidance andantagonistic behaviours with calving and fawning seasons.Prior to calving,parturient mountain caribou (R. t. caribou)dispersed to high elevation sites toevade predation (Cichowski 1989). Bergerud et al. (1984)suggest that byspacing themselves out, caribou Increase the predatorsearching time forcalves and make it difficult for predators to subsist on a diet ofcariboucalves. Similarly, mule deer seek isolation from conspecificsbefore givingbirth (Robinette et al. 1977; Riley and Dood 1984) andpregnant black-taileddeer are known to be highly antagonistic during thefawning period(Dasmann and Taber 1956; Miller 1974).Despite the knowledge that has been gained onthe topic in general,the nature of fawning habitat requirements and birth siteselection in black-tailed deer is poorly understood. The objectives of ChapterThree are:1) to compare the use of seral stages during thefawning period versus therest of the summer, and2) to compare home range size duringthe fawning period with home rangesizes in other summer time periods. Minimum convexpolygon technique(Mohr 1947) is used as an index of peripheral movements.Study AreaThis study took place in the upper CaycuseWatershed, above andincluding Wilson Creek (Figure 1.1). Thissouthwestern Vancouver Island5system flows westward into the head of Nitnat Lake, which in turn emptiesinto the Pacific Ocean midway between the town sites of Bamfield and PortRenfrew. With its center at appro<imately 48° 48’ N latitude and 124° 29’ Wlongitude, the study area falls within the Windward Island MountainsEcosection of the Western Vancbuver Island Ecoregion (British ColumbiaMinistry of Environment 1988).Elevations range from 100 to 1160 m. Relative to many central andnorthern Vancouver Island watersheds, the Caycuse topography is gentleand rolling. Most ‘height of land’ ridge tops fall between 700 and 1000 m.The predominant biogeoclimatic zone is Coastal Western Hemlock(CWH) (British Columbia Ministry of Environment 1988). The subzones Very WetMaritime (Submontane and Montane variants), Moist Maritime (Submontaneand Montane variants), and Very Dry Maritime (Western variant) occurred.The latter however, was sparsely distributed (Lewis 1988). The Submontanevariants occurred from the valley bottom to approximately 600 m. TheMontane variants occurred between approximately 600 and 1000 m (Klinkaet al. 1984). The location of the Montane/Submontane transition wasafunction of elevation and aspect. Montane habitats occurred at slightlylower elevations when on east dnd north facing slopes. On thehighestmountain tops, at approximately 1000 m and above, the Mountain Hemlockzone occurred. This zone was well forested and did not approachsubalpineconditions.A cool mesothermic climate, chardcterized by coolsummers and mildwinters, prevails over the study area. The mean daily maximumandminimum temperatures for July are 21.5 °C and 10.1 °C, respectively. ForJanuary, the mean dailymaximuiLnand minimum temperatures are 6.7 °Cand 0.7 °C, respectively. Moist frontalsystems from the Pacific bring an62i.KilometersFigure 1.1 Location and major subdrainages of the Caycuse StudyArea7average 3649 mm of precipitation annually (Jamie McDuff pers. comm.based on data collected by Environment Canada at the Nitnat FishHatchery Weather Station: 1981-199 1). Snowfall and snow pack persistenceis highly variable between years and between sites. During the winter of1989-90 snow did not accumulate at a 700 m, south aspect snow station untilJanuary. A snow pack in excess of 40 cm then persisted from late January tolate March. In the winter of 1990-91 snow falls in excess of 40 cm began inNovember however, they ceased in late January. In this winter the snowpack was highly erratic and periods of low (<10 cm) snow accumulationsoccurred throughout. In mild wintrs persistent snow accumulations may benegligible on sites as high as 900rn(pers. obs.). In other winters the samesites could have snow accumulations in excess of two m (pers. abs.). In anywinter, snow accumulations within the study area are variable andinfluenced by elevation, aspect and proximity to Nitnat Lake. Low-elevationsites, close to Nitnat Lake usually remain snow free while high elevation,headwater sites often experience accumulations in excess of two m.Timber extraction within the study area began in the early 1940s(Wayne Wall pers. comm.) and continues to this day. Most subdrainageshave been progressively clearcut; thus, valley bottoms are dominated bysecond-growth stands. Most recent clearcuts are found on mid to high-elevation sites. Logged areas were slashburned. Because the study areawas logged from its headwater first, the pattern of logging along theCaycuse mainstem is different. Many of the recent clearcuts occur alongthe mainstem while second-growth stands dominate the sidehills. In 1990,the proportion, by area, of clearcuts, second-growth and old-growth standswithin the study area were 40%, 36% and 24%, respectively (Morgan, et aI.1990).8General Methods and MaterialsDeer capture operations were conducted between mid-Novemberand mid-April during the winters of 1988-89, 1989-90 and 1990-91. When theground was snow covered, Clover traps (Clover 1956) were set in, oradjacent to, winter ranges. Most trap sites were established in areas thatwere previously identified as winter ranges (typically old-growth standslocated on south facing slopes between 300 and 900 meters in elevation[Nyberg et al. 1986]). Others were established on an opportunistic basisaccording to the presence of deer sign. In the absence of snow, freeranging deer were immobilized with succinylcoline (Anectine®). Four to 12mg dosages were loaded into Pneu darts (Pnue Dart Inc., Williamsport, Pa.)and shot from a Palmer Cap-chure gun (Palmer Chemical Co.). Most dartingactivities took place at night in clearcuts. Spot lights allowed the deer to beobserved and darted from vehicles.Captured deer were fitted with a Loteck (Loteck, Aurora, Ont.) radiocollar and a numbered plastic ear-tag. Adult males, which experience neckswelling in the rut, were not collared because it was feared they might bechoked. Young males (8 to 10 months) were fitted with rot-away collarsdesigned to break off before the dnimal became 2 1/2 years old.A Telonics (Telonics Inc., Mesd, Ariz.) receiver and a Lotech, hand held,Adcock type antenna was usedtotrack the collared deer. Approximatelocations were based on the strength and quality of the signal and theamount of ‘swing’ experienced between trial positions. The precise(recorded) locations were then obtained using one of two techniques.The most common approach involved triangulation of three or fourcompass bearings taken from diiferent stations. Telemetry stations were9established every 100 meters along roadways within home ranges and wereidentified on a 1:20,000 planimetric map. To ensure that the bearingsproduced a ‘fix’, the angles were drawn on 1:20,000 scale field-maps. Time,date, and weather conditions were recorded for each location.The ‘Trimle’ program (White and Garrot 1991) was used to estimate theposition of the radio collaredde?r.This program calculates the UniversalTransverse Mercator (UTM) coordinates for the ‘maximum likelihood estimatorwhich is hereafter referred to as the location. In addition, a 95% error ellipsewas generated for each location.Because most collared deer could be approached by road to within afew hundred meters or less, error ellipse size was generally low. Of thetriangulated locations, 75% of the error ellipses were less than 1 ha. Twelvepercent were between 1 and 2 ha and 6% were between 2 and 3 ha. Only7% of the error ellipses were greater than 3 ha.All locations were accepted regardless of their error ellipse size. Iagree with Heligren et al. (1991) who argued that the detection of habitatpreference is conservative in the presence of lowered telemetry precision.Using the same rationale, the detction of differences in home range size isalso conservative in the presence of lowered precision.By comparison, the other technique for estimating locations wassimple. When in clearcuts, the study animals were often observed with theaid of spotlights and/or binoculars. These locations were recorded directlyonto 1:20,000 field maps. Visual locations constituted 30% of all locations.In the summers of 1990 and 1991 when the majority of the data weregathered, each collared deer was usually located three times per week.Locations were gathered according to a schedule that rotated through thefollowing daily time periods: 0000-0600, 0600-1200, 1200-1800 and 1800-240010hours. As a result, a 32-hour (24 hrs +8 hrs) time span usually separatedconsecutive location observations foreach deer. To reduce theautocorrelation of consecutive location observations,the minimumacceptabe time interval belweeh locations for an individual deer was24hours (White and Garrott 1991).Home range estimates for individual deer were based on the minimumconvex polygon (MCP) technique (Mohr1947). The MCP technique’ssensitivity to outliers made it a usefull meansof indexing peripheralmovements during the fawning season. Home rangecenters are arithmeticand produced by averaging the location coordinates(UTM).A geographic information system (Terasoft Ltd., Nanaimo, B.C.)program stored and manipulated several layersof data that were digitizedfrom a variety of sources. A forest cover typelayer, which identified standcharacteristics, was digitized from B.C. ForestService maps (1: 20,000).Another layer contained the BiogeodilmaticEcosystem Classificationinformation (BEC) (Meidinger and Pojar 1991) to the ‘site association level’(Lewis 1988). A topographical map (1: 20,000) was usedto identify anddigitize general aspect polygons which were classified according to thecardinal direction that most closely approximated the orientation of the hillside; north (315-45 degrees), east (45-135 degrees), south (135-225 degrees),and west (225-315 degrees). A 1:50,000 topographicalmap with 20 metercontour intervals was used to determine elevations.11Description of Study AnimalsThis section briefly describes the 13 deer involved in this study. McNayand Doyle (1987) identified three types of migratory behaviours in black-tailed deer: 1) obligate migrators move between seasonal home rangesevery year irrespective of snow accumulation, 2) facultative migrators moveirregularly and in response to snow accumulation and ablation, and 3)resident deer do not migrate.Five of the study animals were facultative migrators that occupieddistinct summer ranges and did not migrate unless forced by snowaccumulations. An additional deer migrated, but because it was notmonitored through more than one migrational cycle, its migratory status(obligate or facultative) was not determined. Three were resident deer. Fourother deer did not migrate, but bcause they occupied mid-elevation sitesand were monitored during the relatively mild winters of 1989-90 and 1990-91, their migratory status could not be determined. Table 1.1 providesgeneral information on the study ahimals.All of the migrators summered in headwater basins and wintered onside hills that were close to the Caycuse Mainstem. Four deer (18181, 17701,17912 and 15181) wintered on the southwest-facing, old-growth slopesnortheast of the Mystery Creek/Cedar Creek/Caycuse River confluence andsummered in Seventy Creek. Two others (12801 and 15301) wintered in southfacing, second-growth stands north of the Cedar Creek/Caycuse Riverconfluence and summered in Cedar Creek. The winter ranges fell between300 and 600 m while the summer ranges fell between 500 and 1000 m.12Table 1.1. General Information on Caycuse study animals, 1989-1991.Deer*Capture Mode of capture Site**Migratory SexID Date Capture Cover Type Status12801 90.03.14 dart 2nd growth facultative F13201 90.01 .19 dart clearcut resident F13891 90.04.10 dart clearcut undet. F15181 91.01.11 trap old growth undet. F15301 90.03.16 trap 2nd growth facultative M16781 90.04.17 dart clearcut resident F16781 90.04.17 dart clearcut resident F17581 90.03.2 1 dart clearcut undet. F17701 89.03.18 trap old growth facultative F17912 90.02.16 trap old growth facultative F17991 90.12.30 trap old growth undet. F18181 89.03.19 trap old growth facultative F18412 90.04.17 dart clearcut undet. M19212 90.03.22 dart clearcut resident F18412 90.04.17 dart clearcut undet. M*Collared deer were identified by the last four digits of their radio frequencies. A final digitwas added to distinguish deer that had used the same collar but at different times.**Clearcuts were 0-15 yrs, second growfh stands were 16-149 yrs, old growth stands were150+ yrs.All of the resident deer (16781, 19212 and 13201) occupied homeranges that were immediately ddjacent to the Caycuse Mainstem andbetween 100 and 400 m.The undetermined deer occupied intermediate ground between 400and 800 m. Three lived in the Wilson Creek drainage (1799 1,18412 and13891), the other lived near McLure Lake (17581).13Chapter Two: Changes in Habitat Usein Relation to Forage Qualitylnti’oductionPlant phenology refers to the periodic phenomena (growth stages) inthe annual cycle of a plant that are controlled by climatic factors(Daubenmire 1974). The nutritive value of a plant is influenced by itsphenology (Klein 1965; Rochelle 1980). Accordingly, the profitabilities (Krebsand McCleery 1984) of forage species vary within the annual cycle ofgrowth.Black-tailed deer selectively forage on a wide variety of plant speciesand adjust their preferences to account for changes in the nutritional qualityand availability of forage species (Miller 1968; Rochelle 1980; Hanley 1984).Preference, defined as the likelihood that a resource will be chosen if offeredon an equal basis with others (Johnson 1980), is dynamic and dependent onthe phenology of available forage species. In cool sites, such as those onshady aspects and/or at high elevations, plant phenologies are delayed(Geiger 1961; Klein 1965; Aitken 1974). By foraging on these sites later in thegrowing season, deer can extend the period of time that they have accessto high-value phenological stages.Vancouver Island’s mountainous habitats are very heterogeneous andbecause each plant species has its own growing site criteria, the abundanceof each forage species is often highly variable or ‘patchy’. In coastal forestecosystems, many herbaceous forage species such as pearly everlasting(Anaphalis margariracea L.), fireweed (Epilobium angustfolium L.) and hairy cat’sear (Hypochaeris radicata L.) are most abundant in the early successional14stages that follow logging, fire or other disturbances (Rochelle1980;Haeussler 1990). Other forage species, such asVacciniun spp. exist underforest canopies and in early successional stages (Rochelle1980; Haeussler1990).Given that plant species are distributed unevenly over most habitatsand that their nutritive values are dynamic; changesin the habitat usepatterns of black-tailed deer, while on their summer ranges, were expected.Use of habitat types at the ‘within home range’ level of selection wasexamined.Seral StageRochelle (1980: v-iv) stated, “fireweed was the most heavily usedspecies during the spring to fall period” and that “it displayed thehighestenergy content of the species (forage) examined”. He also offered that“deer appeared to select plants high in energy and other nutrients in springand summer; availability appeared to have a stronger influence on selectionin fall-winter” (Rochelle 1980:193). When analyzing rumen contents, Rochelleused Mealy’s (1975) percent importance value (IV%) to rank the dietaryimportance of several forage species.IV% is defined as:IV=Frequency of OccUrrence (percent) x Volume (percent)IV%=(IV of forage item/E IV all forage items) x 100Figure 2.1, repeated from Rochelle (1980: 64), clearly demonstratesthat, during the middle and late summer periods, fireweed was by far themost important forage species in the Nimpkish Valley.15ImportanceValue (%)4020Figure 2.1. Monthly pattern of use by black-tailed deer of forage species inforested and cutover areas. from Rochelle 1980,p.64).In the Caycuse and Nimpkish Valleys the Very Wet Maritime subzone ofthe CWH zone is predominant at elevations between 300 and 1000 m (BritishColumbia Ministry of Environment, 1988). In both areas, clearcut loggingbegan in the 1 940s and after logging the sites were slashburned (Harestad1979). Because of these similarities I am assuming that the food habits of the202020Fall-winter spring summerFall-winterVaccinium spp.Blechnum spicant20Cornus canadensisgo604020Epilobiumangustfoilum3 F N A N 3 3 A S 0 N 0(3)’ (5) (8) (6)(1) (6) (5) (Li) (2) (4) (9) (4)Month‘Number of rumensanalysed.16deer within both valleys are similar.Haeussler et al. (1990: 90) stqted that “In many areas of the province,succession is too rapid for pioneer conditions favorable to fireweed to persistfor as long as 20 years” and that “In coastal British Columbia, once a canopydevelops, fireweed dies out”. Gates (1968) studied post-logging seralsuccession in a central Vancouver Island drainage and concluded,“herbaceous plants reach their peak cover-densities about three years afterburning” and that “thereafter they declined gradually through competitionand shading and are of minor importance by the fifteenth year.” This iscorroborated by Dyrness (1973) who found that invading herbaceous speciesdominated post-logging burned sites from the second through fourth yearsonly. Gates also noted that in old-growth, forbs covered less than onepercent of the surface.Most Caycuse second-growth stands were devoid of fireweed. Itpersisted, sparsely and in lower vigor, in only a few of the stands with moreopen canopies (pers. obs.). In the recent clearcuts, fireweed dominated theplant community. During the July-August flowering season, most youngclearcuts exhibited a pinkish hue, owing to the plant’s high density.In the later portions of the summer, when the IV% of fireweed wasfound to be highest (Rochelle 1980). deer should maximize their foragingefficiency by increasing their use of clearcuts. By minimizing the amount oftime spent foraging deer are able remain inactive for greater proportion oftime. Inactivity allows deer to be more vigilant for predators and it mayreduce their risk of detection by predators. In examining the use of differentseral stages through the summer months, I proposed a null and an alternatehypothesis.17Ho Percent use of clearcuts inthethree periods: 1) April/May/June;2) July/August and 3) September/October does not differ.Hi Percent use of clearcuts correlates with Rochelle’s (1980)IV%.Clearcut use should be lowest in April/May/June, highest in July/Augustand intermediate in September/October.Percent use is defined as: (no. locations within a habitat type/total no.locations] x 100.Delayed PhenologyDuring the mid and late summer periods, deer should seek out coolersites where delayed phenology maintains many plants in their more nutritiousstages of growth (Geiger 1961; Klein1965; Aitken 1974, Bunnell 1990). Inexamining the use of cooler sites in the summer months, I proposed a nulland an alternate hypothesis.Ho The percent use of cooler sites does not change as the summerprogresses.H2 The percent use ofcooler sites increases as the summer progresses.Three temperature related habitat variables are tested independently.They are; BEC variants, elevation, nd aspect.18Methods and MaterialsLocation data were gatheredduring the summer months.The timeframe for ‘the summer’ was basedon the average phenologyof fireweed onthe south coast of BritishColumbia and on the periodover which Rochelle(1980) found it to beimportant to the diet ofblack-tailed deer. Fireweedbegins to develop aerialshoots in late March (Haeussleret al. 1990). By earlyOctober most of fireweed’saerial shoots have died back;however, fireweedcontinued to be animportant dietarycomponent until November.Accordingly, 1991 datacollection started on April1st, and terminatedonOctober 30th.In April 1991, snowaccumulations persisted inmany of the summerranges within the study area.Data collection wasdelayed for some studyanimals because of thepossibility that snowaccumulations, ratherthanforage quality, could beinfluencing habitat use atthis time. Locationdatawere not used if they werecollected before thedeer’s summer rangewasfree of snow. Traceamounts of patchysnow were acceptablebut aconsistent snow pack ofonly one centimeterin any portion of thehomerange was not.In regions where appreciablesnow accumulationspersist well intothesummer, this approach wouldnot be feasible. Theabatement of snowonsouthern Vancouver Islandhowever, is usuallyearly and rapid. Thespring of1991 was no exception.Table 2.1 summarizesthe 1991 datacollectionperiods.19Table 2.1. Data collection periods for collared deer, 1991.ID Start Date Finish Date16781, 13201, 17581 April 1st October 31st17991, 18181, 13891 April 10th October3lst15181, 12801. 15301 AprU 18th October 31st17912, 17701In 1990, data collection starfed on June 1St and ended October 31st.Those data are presented (though not statistically tested) to substantiate the1991 findings. Study animals 17991 and 15181 were not in the 1990 data set.Although study animals 19212 and 18412 were not in the 1991 data set, theywere in the 1990 data set.Four habitat variables; elevation, aspect, biogeoclimatic ecosystemclassification (BEC) variants (Meidinger and Pojar 1991), and seral stage weredeveloped from sources identified in Chapter One. The latter two requirefurther clarification. The BEC Montane variants of the Moist Maritime andVery Wet Maritime subzones were amalgamated to form the Montanehabitat type. The Submontane variants of the above subzones and thewestern variant of the Very Dry Maritime subzone were amalgamated intothe Submontane habitat type.Logged habitats were grouped into two seral-stage habitat typesbased on the dates of stand establishment after harvest, as they related tothe production of herbaceous forage. Stands were considered established ifthey were satisfactorily stocked(>500 stems/ha) with a commerciallydesirable (coniferous) species. The date of establishment was determined bythe age of the tree stock. Clearcut habitats were less than 16 years postestablishment in 1990. Second-growth habitats were at least 16 years post-20establishment in 1990. Old-growth habitats were at least 150 years old andhad never been logged. Because both second-growth and old-growthforests contained low quantities of herbaceous forage, they wereamalgamated to form the ‘forested’ habitat type.Most habitat use studies compare the use of particular habitat typeswith their availability, to develop preference ratings (Neu et al. 1974, Johnson1980, Alldredge and Ratti 1986). Because the intent of this study is todocument changes in the use of various habitat types through time, thesecalculations are not necessary. Instead the summer was divided into timeperiods which were, with one exception, two months long. Percent use ([no.locations within a habitat type/total no. locations] x 100) of habitat types orclasses for each period are then compared.Time periods developed to test for shifts in clearcut use wereApril/May/June, July/August, and September/October. These periods werebased on changes in Rochelle’s (1980) percent importance value (lV%) andnutritive values of fireweed. Fireweed’s IV% were relatively low during April,May and June (see Figure 2.1). The percent importance value rose sharply toa peak in July and remained high through to October. Because the nutritivevalues (digestible dry matter and caloric content) of fireweed rapidlydeclined after August (Rochelle 1980), the data were separated further intoJuly/August and September/October time periods.Time periods developed to test for increased use of cooler sites wereApril/May (spring), June/July (summer) and August/September (early fall).October’s data were not included in this test because: 1 )they were notrequired to test the hypotheses, 2) the possibility of a downward elevationshift in October in response to cooler weather existed and 3) by late August,any upward elevational migrations should have ceased.21Mohr’s (1947) minimum convex polygon technique (MCP) was used toestimate the home range boundaries of the study animals. The95% MCPhome range, defined as the smallest convex polygon containing95% of thesample locations (Akerman, Leban, Samuel and Garton 1989). was used.The G-test (Sokal and Rohlf 1981) was used to test for differences in theuse of the habitat types between established time periods. A one-tailedpaired difference test (Sokal and Rohlf 1981) was used to test for changes inthe elevation of home range centers and the average location distancefrom the home range center during different time periods. A type one errorprobability (p-values) less than 0.05 was considered to be ‘strong evidenceagainst the null hypothesis’ (Ho). A probability between 0.05 and 0.10 wasconsidered to be ‘moderate evidence against the null hypothesis’. Aprobability greater than 0.10 was considered to be ‘no evidence against thenull hypothesis’.ResultsIn the summer of 1991, 812 deer locations were recorded. Eleven deerwere monitored during this period; however, one (15301) was killed by wolvesin mid-July. Table 2.2 presents thenumber of locations per collared deerpermonth.In the summer of 1990, 11 deer were monitored and 542 locationswere recorded.22Table 2.2. Number of locations per deer per month, 1991.Deer April May June July Aug. Sept. Oct. Total13201 11 14 13 11 14 9 12 8416781 10 14 12 10 14 9 12 8117581 8 14 12 10 14 9 12 7913891 8 14 10 10 13 10 12 7717991 9 13 12 11 14 10 12 8118181 8 14 13 11 14 10 12 8217701 4 14 12 11 14 11 12 7817912 4 14 13 11 14 10 12 7815301 3 12 8 7 na na na 3012801 4 14 9 11 14 10 12 7415181 3 14 6 11 13 10 11 68TOTAL 72 151 120 114 138 98 119 812MEAN 6.5 13.7 10.9 10.4 13.8 9.8 11.9 78.2Seral StageIn 1991 the percent use of clearcut habitats was highest in the July!August period (Table 2.3). Pooled location data, for the nine deer which hadclearcuts in their home range (708 locations), reveal that the percent use ofclearcuts was 67.9% in April/May/June, 78.2% in July/August and 69.2% inSeptember/October (Table 2.3, Fig 2.2). The G-test (G=7.37, p0.025)provided strong evidence against the null hypothesis which predicted thatthe percent use of clearcuts would not change as the summer progressed.Over the entire summer, less than 3% of the locations were within oldgrowth.23Table 2.3. Number of locations withn clearcut and forested habitats andpercent of locations within clearcut habitats, 1991.*numberof locations within clearcut and forested habitatsAiril/Ma /June Julv/Auciust Set.IOct. Aoril-Octoberdeer Id. cc:for* cc cc:for cc cc:for cc cc:for cc%**% % %13201 38:0 100.00 25:0 100.00 15:6 71.43 78:6 92.8513891 17:15 53.13 21:2 91.30 12:10 54.55 50:27 64.9315181 0:23 0.00 0:24 0.00 2:19 9.52 1:67 1.4716781 33:3 91.67 21:3 87.50 17:4 80.95 71:10 87.6517581 22:12 64.71 23:1 95.83 21:0 100.00 66:13 83.5417701 28:2 93.33 25:0 100.00 22:1 95.65 75:3 96.1517912 31:0 100.00 22:3 88.00 20:2 90.91 73:5 93.5817991 20:14 58.82 21:4 84.00 15:7 68.18 56:25 69.1318181 10:25 28.57 14:11 56.00 11:11 50.00 35:47 42.68pooled 199:94E67.9j172:48 78.2 135:60 69.2 506:202 71.5**=percentof locations within clearcut habitats24807876C)74.$272.! 7068::62, IApril-June July/Aug. Se pt/OctTime PeriodsFigure 2.2. Percent of pooled locations within clearcuts, 1991.These data compare favorably with the 1990 data (464 locations)where the percent use of clearcuts for the July/August andSeptember/October time periods was 78.2% and 73.7% respectively.Delayed PhenologyFor the eight deer which had Montane habitat within their homerange487 locations) percent use (pooled data) of Montane habitat was 14.5% inApril/May, 22.6% in June/July and 25.8% in August/September (Table 2.4, Fig.2.3). The G-test (G=6.67, p=0.036) provided strong evidence against the nullhypothesis which predicted that the percent use of Montane habitat wouldnot change as the summer progressed.25Table 2.4. Number of locations within Montane and Submontane habitats andpercent of locations within Montcine habitats, 1991.August/Sept.(early fall)*=number of locations within Montane and Submontane habitats**=percentof locations within Montane habitatsApril/May(sDring)June/July(summer)April-Septemberdeerid. Mon:Sub Mon. Mon:Sub Mon. Mon:Sub Mon. Mon:Sub Mon.*% 70 7012801 0:18 0.00 0:20 0.00 3:21 12.50 3:59 4.8415181 6:11 35.29 11:6 64.71 14:9 60.87 31:26 54.3915301 0:15 0.00 5:10 33.33 no na 5:25 16.6717581 1:21 4.55 0:22 0.00 3:20 13.04 4:63 5.9717701 0:18 0.00 4:19 17.39 6:19 24.00 10:56 15.1517912 4:14 22.22 10:14 41.67 7:17 29.17 21:45 31.8217991 0:22 0.00 0:23 0.00 1:23 4.17 1:68 1.4518181 11:11 50.00 8:16 33.33 9:15 37.50 28:42 40.00pooled 22:130 14.5f38:130 22.6 43:124 25.8J103:384j21.22630 -to25-.S2015=oSpring Summer EarlyFeIlTime PeriodsFigure 2.3. Percent of pooled locations withinMontane habitats, 1991.These results compare favorably with the 1990data set which hadpercent use of Montane habitat valuesof 19.8% to 24.3% for June/July andAugust/September, respectively.The percent use of north and east (cooler)aspects was not higher inthe latter portion of the summer. Instead the1991 pooled data set for all ofthe 1991 study animals (693 locations) revealedthat the percent use of theseaspects combined was 26.5% for April/May, 26.1%for June/July and 24.2% forSeptember/August. When only location data fromthe eight deer which hadMontane habitats within their summer range (487locations) were used, thepercent use of north and east aspectswas not higher in the latterportion ofthe summer. The percent use of north andeast aspects combined was36.2% for April/May, 35.7%for June/July and 32.3% for September/August.The distribution of Montane variantsover the landscape is a function of0-27elevation and aspect. The higher use of Montane habitats in the latterportions of the summer without a similar trend in the use of east and northaspects suggested that the study animals were using higher elevations as thesummer progressed. The elevations of the home range centers of the deerwhich had Montane habitats within their home ranges were averaged andcompared. For the April/May, June/July and August/September time periodsthe mean elevations were 660.0 m (SE=33, n=7), 666 m (SE=51, n=7) and 674meters (SE=43, n=7) respectively (1530 excluded). The mean elevation gainper deer between the home range centers of April/May andAugust/September was 14 m (SE=11, n=7). Although the mean elevationchange was positive, a paired difference test provided no evidence againstthe null hypothesis (t=0.22, p=0.83) which predicted no change in elevation.An alternate hypothesis could be formulated to account for theincreased use of Montane habitats. If the majority of its home range waslocated below Montane variants, a deer could increase its use of Montanehabitats, without shifting its home range center (and therefore its averageelevation) by expanding its home range equally along the axisperpendicular to the Montane/Submontane ecotone. Such an expansiondid not occur. The average distance between the locations and theirrespective home range centers for the spring and early fall periods were 214m, and 205 m respectively.Potential Confounding VariablesThe pattern of logging in many Vancouver Island watersheds makes itdifficult to distinguish between increases in the use of Montane habitats andincreases in the use of clearcut habitats. Montane variants, were often thelast to be logged and as a result they usually contain a high proportion ofrecently clearcut habitats. Conversely, Submontane variants usually contain28a high proportion of second-growth habitats.Before any generalizationsonchanges in the use of seral stage and or BECvariants can be made, thepotential for these variables to be confoundingmust be assessed.Animals 12801 and 15301were not included in seral stageanalysisbecause their summer ranges consistedentirely of one seral stage (second-growth). Similarly, animals 13201, 13891,and 16781 were not included in theBEC variant statistical analysis becausetheir home ranges consisted entirelyof Submontane variants. The 6study animals used in bothanalyses wereanimals 15181, 17581, 17701, 17912,17991 and 18181. The Montaneportionsof the home ranges belonging toanimals 15181. 17581 and 17991 consistedentirely of second growth habitats;therefore, increases in the percentuse ofMontane and clearcut habitatscould not have been correlated.Similarly,the Submontane portions of the homerange occupied by animal 17912consisted entirely of clearcuts.Again, increases in the percentuse ofMontane and clearcut habitatscould not have been correlated.A breakdown of seral stage habitattypes within the Montane andSubmontaneportions of the 95% MCP home rangesof the remaining two studyanimals ispresented on Table 2.5.The percentage of clearcuthabitats within the home rangeof animal18181, was lower inthe Montane variants thanin the Submontane variants(Table 2.5). Thus, there was not a potentialfor increases in the percentuse ofMontane and clearcut habitats tobe correlated. Within thehome range ofanimal 17701 the percentageof clearcut habitats washigher in theMontane variants than in the Submontanevariants.29Table 2.5. Percent clearcut habitat within the Montane and Submontanevariants within the 95% MCP home ranges of study anImals 17701and 18181.Deer Id. Percent Clearcut Habitat Percent Clearcut Habitatwithin Montane within Submontane17701 100.0 90.918181 11.8 38.2The location data were separated by month and inspected forcoincidental increases in the use of Montane and clearcut habitats. Thesedata are summarized in Table 2.6.Table 2.6. Percent use of Montane and clearcut habitats for study animal17701.Month April May June July Aug. Sept. Oct.% Clearcut 100.0 85.7 100.0 100.0 100.0 100.0 91.7% Montane 0.0 0.0 25.0 9.1 21.4 27.3 0.0As Table 2.6 demonstrates, animal 17701 used clearcuts regardless ofwhether they occurred in Montane or Submontane variants.30DscussionSeral StageAn increase in use of clearcuts by deer during July and August is oftennoted by biologists who conduct census work on Vancouver Island withinthat period. The deer hunting sedson, which on Vancouver Island starts inearly September, may force deer into areas of greater visual screening andout of clearcuts. This however, would not explain the relatively low use ofclearcuts during the early summer.The observed trend in cledrcut use, for the Caycuse Study Area in1991, was similar to the trend documented by Rochelle (1980) for fireweeduse (IV%) in the Nimpkish Valley. Clearcut use and lV% were lowest duringApril/May/June, highest during July/August and of intermediate values inSeptember/October.Old-growth and second-growth habitats within the summer ranges ofthe nine study animals, supported very little herbaceous forage. In theadjacent clearcut areas howevei, large quantities of fireweed and otherpioneering herbs grew vigorously. It is not surprising therefore, that theincreased fireweed use (IV%) coincides with an increase in the use ofclearcut habitats.It is difficult to determine whether the increased IV% of fireweed andthe increased use of clearcuts are a function of forage availability or foragequality. Rochelle (1980) noted th9twhen availability is high for a variety offorage species, deer appear to e selecting the forage with the highestnutritive value. Klein (1965) suggested that unless forage quality in thesummer months is maintained, deer cannot consume enough to meet theiroptimum growth requirements.31Hanley (1984) found that it, the western Cascades of WashingtonState, the availability (measured as biomass) for all forage species peaked inJuly. The biomass of fireweed increases until after its flowering stage inAugust however, only the most digestible, top portion of the plant isconsumed (Moen 1981, pers. obs.). I believe that, between emergence anddie-back, the availability of fireweed tops at any given site is constant.Fireweed availability, therefore, does explain the percent use of clearcuts orthe lV% for fireweed which were highest in July and August.The nutritive quality (dry natter digestibility and caloric value) offireweed is at its peak in August. The percent dry matter digestibility (DDM) offireweed increased steadily from 60% in May to 80% in August then backdown slightly to 78% in October (lochelle 1980). Similarly, the caloric valueof fireweed rises sharply from 3 kccil/0.8g in May to 6 kcal/0.8g in August thenback to less than 3 kcal/O .8g in October (Rochelle 1980).A variety of non-herbaceous forage species do persist beneathforestcanopies. Rochelle (1980) found that, within forested habitats, Vacciniumalaskense and Vaccinium parvfolium followed trends similar to that of fireweed.However, peak DDM values, of approximately 60%, occurred muchearlier inMay. Caloric values rose sharply to a plateau of approximately 5 kcal/0.8gwhich lasted from April to July. Given their similar phenologies (Haeussler etal. 1990), it is probable that other important shrub forage species, such astheRubus spp. followed similar nutritive cycles.In the early summer therefore, the nutritive values (DDM and Kcal/g) ofseveral alternate forage species, found within forested habitats, arecomparable to that of fireweed. As summer progresses however,fireweedexceeds the alternate forage species in nutritive value and itdominates thediet (lV%). The trends in the percent use of clearcut habitats andthe lV% of32fireweed are similar.Clearcut habitats possess fireweed in its highest density and vigor(Haeussler et al. 1990). By selectihg clearcuts as foraging sites in July andAugust, deer maximize their foragihg efficiency. In doing so, they are able tomaintain quality in their diet and remain inactive for a greater proportion ofthe day. Inactivity allows deer to be more vigilant and it decreases theprobability of detection by predators.Delayed PhenologyDuring the spring months, forage phenology within the coolerMontane habitat is delayed (Geiger 1961, Aitken 1974). At this time theforage within the warmer Submontane habitat is more phenologicallyadvanced and more nutritious. Not surprisingly, the percent use ofSubmontane habitat is highest in the April/May period. As summer progressesthe more profitable phenological stages become available in the Montanevariants and the percent use of this habitat increases.Mackie (1970) found that mule deer used drier habitat types less oncethe forage plants became desiccated. Movements away from sites withovermature forage and into sites with new growth enable deer to maintainquality in their diet (Klein, 1965). In Caycuse however, the percent use ofMontane habitat increased between spring and early summer, well beforedesiccation would have occurred within Submontane habitats. Black-taileddeer summer forage species attain their maximum nutritive values in June,July and August (Rochelle 1980). The increased use of Montane habitatstherefore, was not in response to lower forage quality within Submontanehabitats. Instead, deer use of Montane habitat was minimal when theabundance of highly nutritious forage within it was low.33The distribution of Montane variants over the landscape is primarily afunction of elevation and aspect. It is surprising therefore, that gains inelevation and/or the use of easf and north aspects were not detectedcoincidentally with the increase In the percent use of Montane habitat.Failure to detect changes in elevation may have been a function of samplesize and/or the size of the elevational movements. The 1991 summer ranges(MCP) of 10 female study animals averaged only 55 ha (Table 3.1). Thus,movements into Montane habitats usually required only small elevationalgains.34Chapter Three: Changes in Habitat Use by FemaleBlack-Tailed Deer During the Fawning SeasonIntroductionHerding cervids such as caribou and elk often move to specific,1well-defined calving habitats (Altmann1952; de Vos 1960). The natal habitats ofthe more solitary members of the deer family such as moose, white-taileddeer (Odocoileus virginianus) and black-tailed deer are more difficult tocharacterize (Miller 1974; Stringham 1974). Parturient deer should seekhabitats that confer anti-preddtor advantages during this period ofvulnerability. Livezey (1991) and Mackie (1970) found that black-tailed deerand mule deer respectively, used the same areas during fawning each year.They did not, however, identify favored habitat attributes in such areas.To prevent predators from detecting their newborn fawns, black-taileddeer employ the ‘hider strategy’. When very young, ‘hiders’ have little scent,are cryptically colored, and crouch when threatened (Lent 1974; Geist 1981).During their first weeks of life hiders do not follow their mothers. Instead theyremain hidden, often for many hours, while their dams forage or bed in otherareas. Red deer dams infrequently visit their offspring to nurse them and tostimulate them to defecate by licking their anal region (Clutton-Brock et al.1982). Feces and urine are ingested by the dam to remove evidence thatwould betray the presence of the fawns. Geist (1981) noted that the hiderstrategy often includes ingesting the placenta and birth fluid soakedsubstrate, early removal of young from birth sites, conspecific segregationduring fawning and movements where possible, into escape terrain.Neonate hiders have been found to use habitats with dense groundcover (Johnson 1951, Altmann 1952, Huegel et al. 1986). Although forests35provide screening cover, they often lack the dense ground cover found inmore open sites. Johnson (1951) found that new-born Rocky Mountain elkcalves were likely to be found in sagebrush (Atemisia spp.) habitats and thattheir use of forested habitats decreased as the distance from the forest edgeincreased. Calving sites were associated with the dense ground cover foundin earlier successional stages, not beneath coniferous canopies. Huegel etal. (1986) found shrub and sapling densities to be higher at or around white-tailed deer fawn bedsites and Robinette et al. (1977) found that areas withdenser cover were used by mule deer for fawning. On Vancouver IslandGates (1968) found that “shrubs and young deciduous trees dominate thelow stratum of vegetation from the third year through to at least the fifteenthye&’. If black-tailed deer prefer to fawn in areas with dense ground cover,then their use of clearcut habitats in the fawning season should increase.In 1991, all clearcut habitat polygons used by collared deer, exceptone, were between three and 16 years of age and supported dense groundcover (herbaceous, shrub and seedling percent cover 2 m from theground). The adjacent second- and old-growth stands supported sparserground cover (pers. obs.). In examining the use of clearcut habitats duringthe fawning period versus other summer periods, I proposed a null and analternate hypothesis.Ho The percent use of clearcuts by maternal black-tailed deer is notdifferent during the fawning season than in the remainder of thesummer.Hi The percent use of clearcuts by maternal black-tailed deer is higherduring the fawning season than in the remainder of the summer.36During the fawning period, adult females appeared to move outsidetheir typical area of use with greater frequency. Often only a short time wasspent outside the typical area of use before the doe returned. Theseobservations led me to postulate that black-tailed deer give birth outside oftheir core areas of use to avoid contaminating their preferred sites withscents associated with parturition. After a few days, they lead their fawnsback into their core area and away from scents that might attract predators.If these postulates are true, then we should expect to measure an increase inhome range size during the fawning period. In examining this relationship, Iproposed a null and an alternate hypothesis.Ho Home range size of maternal black-tailed deer is not differentduring the fawning period than in other summer periods.H2 Home range size of maternal black-tailed deer is larger during thefawning period than in other summer periods.I also suspected that maternal females would reduce their area of usein the period immediately after fawning because of their tendency to remainnear their sedentary fawns. I again proposed a null and an alternatehypothesis.Ho Home range size of maternal black-tailed deer is not differentduring the period immediately after fawning than in other summerperiods.37H3 Home range size of maternal black-tailed deer is smaller during theperiod immediately after fawning than in other summer periods.Method and MaterialsThe summer of 1991 was divided into the following 29-day periods: thepre-natal period (PrNP), May 8 to June5; the fawning period (FP), June 6 toJuly 4; and three post-natal periods, (PoNP1) July5 to August 2, (PoNP2)August 3 to September 1, and (P0NP3) September 2 to September30.Because the average number of locations per deer was lower in the PoNP3than in the FP, an adjusted P0NP3 was also analyzed. To produce the‘PoN P3-adjusted’ data set, locations from early October were added in theorder of their chronology to the PoNP3 data set. The locations were added(or subtracted) until the number of locations for each deer was the same inthe P0NP3-adjusted as in the FP. In the case of 15181 and 12801, locationswere removed. The number of locations per period and per deer aresummarized on Table 3.1.During the summers of 1988, 1989, 1990 and 1991, an affiliatedresearch project captured a totdl of 246 neonate fawns in the CaycuseStudy Area and in the South Fork of the Nanaimo River. The Nanaimo RiverStudy Area is 30 km north of the Caycuse Study Area. The earliest and latestfawn capture dates within any summer were June 6th and July 4th,respectively. These dates were used to define the 29 day range of thefawning period (FP). Like white-tailed deer fawns (Heugel et al. 1985), blacktailed deer fawns, up to approximately 10 days old, could be captured by aperson on foot. The FP therefore, extends further into the summer than itwould have if it had been based on the time of parturition. The median for38parturition. The median for the FP fell betweenthe 19th and 20th of June.Thomas (1970) estimated the mean dateof birth for a central VancouverIsland watershed to be 4.5 days edrlier, June14th.The fawning period for this study was designedto produce informationon natal sites and habitat selection. Becauseblack-tailed deer neonatesare not immediately mobile (Miller1965) and because I could not be sure oflocating the study animals at the exact time when theygave birth, the lag inthe Caycuse FP was appropriate for this analysis.As in Chapter Two, forest age determined standclass. Forestedhabitats were old-growth or at least 16years post stocking in 1990. Siteslogged and restocked more recently wereclassified as clearcuts.The G-test (Sokal and RohIf 1981) was used to test for interactionsbetween the percent use of thestand class variable and the establishedtime periods.The second section of this chapter examines home rangesize duringthe time periods. The minimum convexpolygon technique (Mohr, 1947) wasused to estimate home range size foreach study animal in all time periods. Aone-tailed paired difference test (Sokal and Rohlf 1981)was used tocompare the home range sizes of different time periods.Type one errorprobabilities (p-values) less than 0.05 were considered tobe ‘strong evidenceagainst the null hypothesis’ (HO). Probabilities between0.05 and 0.10 wereconsidered to be ‘moderate evidenceagainst the null hypothesis’.Probabilities greater than 0.10 were consideredto be ‘no evidence againstthe null hypothesis’.Anecdotal information depicting the fawning movementsof somestudy animals is also provided. The data for this section were collected inthesummers of 1989, 1990 and 1991. Many of the locations usedfor the39‘Fawning Site Selection’ section were not separated in time by a 24 hourminimum. Many of the locations were separated by only a few hours whileothers were separated by a few days. In this section, the post-natal homerange was not subdivided into three post-natal home ranges. The95% homeranges (Hartigan 1987) were calculated by program Home Range(Ackerman et al. 1989). Pre- and post-natal home ranges were separated intime by the ‘fawning movements’. Fawning movements were those thatoccurred between June 6th to July 4th and that were known, or weresuspected, to be movements to, or from, the natal site.ResultsDuring the summer of 1991, ten adult (>1.5 years old) females weremonitored through five time periods producing a total of 563 locations.All of the study animals except 12801 and 17581 were observedinteracting with fawns that appeared to be theirs. Animals 12801 and 17581may have fawned, but because they occupied home ranges with heavyvisual screening, their reproductive status was not determined.Habitat UsePercent use of clearcuts did not increase during the FP. Pooledlocation data (563 locations) revealed that 67.9% and65.6% of locationswere within clearcuts during the FP and during the remainder of the summer(PrNP, PoNP1, PoNP2 and PoNP3 combined) respectively. The G-testprovided no evidence against the null hypothesis (G=0.20, p=0.66).Fawning Site SelectionExact fawning sites were determined in only two instances. In June of1989 and 1990 study animal 18181 was located once every three hours for athree-day monitoring period. Her natal sites were found in both years. They40were within 70 m of one another and both were beyond the animal’s typicalarea of use. The 1989 and 1990 natal sites were 250 m and 600 mrespectively outside of the 95% pre-natal home ranges (Figs. 3.1 and 3.2).Similarly, the 1989 and 1990 natal sites were 28 m inside and 35 mrespectively outside their 95% post-natal home ranges. The ‘fawningmovements’ shown on Figs. 3.1 and 3.2 were from all locations taken duringthe three-day monitoring periods.In 1991, 13201 made several movements beyond the northwesternperiphery of her 95% home range (Fig. 3.3). Although the natal site wasnever detected, the fawn was first observed on the northern periphery of the95% post-natal home range. The ‘fawning movements’ shown on Fig. 3.3were developed from all locations collected within the FP.In other instances, large erratic movements were made during the FP.In 1991, animals 17991 and 12801 were found 1.6 and 3.5 km respectivelyoutside of their 95% pre- and post-natal summer ranges (Figures 3.4 and 3.5).Since both movements were single events involving only one peripherallocation each, the fawning movements are the departures from, and returnsto, the 95 % home ranges.540860n395000413958005407500<Pre-natal 95% MCPFawning movements• Birth location• Post-natal locations1:7,000Post-natal 95% MCP*Pre-natal locationsFig. 3.1. The 1989 fawningmovements of study animal18181.Two locations that were out side thetypical home range area that occurredin July are not shown. Their UTM coordinatesare (393320, 5406590) and (394620, 5408850)..,•• •- • ••• •• ••• ••.- •• •..F •• ••-•• • •••SSSt•_42...I\.\.\ I• . \ ••• .... . ‘•.• I •.. .. ..1 •\\*•.•I•\• .\ .._••• • •1.. ..•.—•).•—--- •_,-. • —.-‘I•_•— )•_•/•‘ —•—•—.,•‘“• Brthbcation•- Pre natal95% MCPFawning movements<.I• Post-natal locahons!! Post-natal 95% MCP*Pre-natal locations1:7,000396000•407600•••I••I•• ••• •I••••540B60(I394800Fig. 3.2. The 1990 fawning movements of study animal18181.435405300.3922005406500390400<Pie-natal 95% MCPFawning• First lawn obseivatloni.-.• Post-natal locations1:10,000L.. - .!Post-natal 95% MCP*Pie-natal locationsFig. 3.3. The 1990 fawning movements of study animal13201.4454108003904• 8-.—..• f • •1••‘i a?.ft• II;-I.!.-.•*38930C’_______________________________5408400IJPre-natal 95% MCPFawning movementsFHSt fawn observationi -.• Post-natal locationsI •P t- ta195°/ MCPLU2,000L. —.0*Pre-natal locationsFig. 3.4. The 1991 fawning movements of study animal 17991.45‘392400.5410700--.-.- .‘,..•‘ •..4•_,I4.•**3I00n5406300Pre-natal 95% MCPFawning movements • Post-natal locations1Post-natal 95% MCP*Pre-natal locations1:22,000Fig. 3.5. The 1991 fawning movements of study animal 12801.46Home Range SizeHome range sizes (100%) for each study animal within all time periodsare summarized in Table 3.1. The size of each deers total summer range(occupied from May 8 to September 30) is also given. The mean homerange size for each period is alsopresentedgraphically in Fig. 3.6.Table3.1.Homerangesize(hectcires)bytimeperiod.DeerTotalnPrNPiiFPnPoNPIitPoNP2itPoNP3itPoNP3itIDSummeradjustedHome Range12801140.88556.081267.28911.931215.781220.601015.8191320177.505721.641125.531318.171216.871222.31951.46131389132.32547.541214.64105.851114.28119.27109.27101518173.54514.741216.11616.531216.521134.851010.9361678134.925517.161112.341216.101113.821217.33917.46121758119.36573.08137.14124.801110.61126.5296.52121770128.425912.651212.53128.44127.751210.761110.76121791230.335911.731212.06136.371212.151213.281013.28131799186.585716.911149.82125.471210.471213,221015.11121818128.025912.211217.191313.251210.25123.89104.4113mean55.1956.311.3711.823.4611.210.6911.712.8511.815.139.815.5011.2SE12.231.916.171.620.972.814.20==48302520015xI.F50 I I I IPrNP FP PoNP1 PoNP2 PoNP3Time PeriodFigure 3.6. Mean summer home range size and standard error for 10 femaledeer In five periods during the summer of 1991.The distribution of the home range size data set (PrNP, FP, PoNP1,PoNP2 and P0NP3 combined) was not normal (Sha,piro-Wilk normality test,W=0.71, p0.001 [SAS 1988]). A natural log (In x) transformation improved thevalue of the W-statistic to 0.97 (p=0.32).Home range sizes during the FP were compared to those of the othertime periods with a paired difference test. The test results of the transformed(In x) data are summarized in Table 3.2.49Table 3.2. Paired difference test comparison of home range size duringthe fawning period versus all other periods.Comparison FP-PrNP FP-PoNP1 FP-P0NP2 FP-P0NP3 FP-PoNP3adjustedMean Difference 12.1 12.8 10.68.3 8.0in Hectarest-value 2.54 2.71 1.89 1.53 1.70Probability 0.016 0.012 0.046 0.080 0.06 1(p-value)Home range sTze was greater in the FP than in any other period. Whenthe FP was compared to the PrNP, the P0NP1 and the P0NP2 the t-testprovided strong evidence against the null hypothesis. When the FP wascompared to the PoNP3 and the P0NP3-adjusted the f-test providedmoderate evidence against the null hypothesis.A migration in the FP away from an early-spring range might explainthe large home ranges in this period. The mean distance between the PrNPand the PoNP1 home range centers however, was determined to be only162 m (SE=30). This small shift was less than the 190 m mean difference(SE=30) between the home range centers of the PoNP1 and PoNP3. Shifts inhome range location therefore, were not larger in the early portion of thesummer than in the late portion and they can not be used to explain thelarger home ranges during the FP.Home range sizes in the immediate post-fawning period (PoNP1) werecompared to those of other time periods with a paired difference test. Theresults are summarized in Table 3.3.50Table 3.3. Paired difference test comparison of home range size duringthe immediate post-fawning period versus all other summer periods.Comparison PrNP-P0NP 1 FP-PoNP 1 P0NP2-PoNP 1 PoNP3-PoNP1Mean Difference 0.68 12.8 2.2 4.4in Hectarest value 0.088 2.71 1.95 1.58Probability 0.466 0.012 0.042 0.075(p-value)Home range size was smaller in the PoNP1 than in all other time periodsexcept the PrNP. When the PoNP1 was compared to the FP and the P0NP2the t-test provided strong evidence against the null hypothesis. When thePoNP1 was compared to the PoNP3 the t-test provided moderate evidenceagainst the null hypothesis.DiscussionHabitat UseThe hider strategy predicts that mother and offspring are together foronly brief periods. A fawning-related increase in the use of any habitat typetherefore, would be difficult to detect because the relationship between thesites occupied by the dams and those of their offspring is weak.It is probable however, that a relationship between clearcuts andfawning habitats did not exist. Forest canopy and shrub layer heterogeneitycould have allowed the parturint study animals to meet their coverrequirements within forested habitats. Although ground-cover was usually51less dense in the forested habitats, many pocketsof dense ground coverwere present along creeks, slides, wet areas,and in other areas of low standdensity. Depending on the site conditions, shrubs,such as red huckleberry,devil’s club (Oplopanox horridus (Smith) Miq.) andsalal (Gauliheria shallon Pursh),often thrive within second- and old-growthforests.Area of UseBecause the locations of fawning sites were rarely determined,theanecdotal evidence presented servesonly to illustrate the development ofthe alternate hypothesis (H2); ‘home range sizeof maternal black-tailed deeris larger during the fawning period than in othersummer periods’.Average home range size was larger in the fawning period thanin allother periods. Increases in home range size during the FP werecaused byincreases in the magnitudeand/or rate of movements to the peripheries ofthe home ranges. Similar peripheral movements of parturientblack-taileddeer and mule deerwere found by Livezey (1991) and Riley andDood (1984)respectively. Livezey reported that “three deer (of 16) fawnedin areas thatwere at least 150 m (range 150-450) higher in elevation than themaximumelevation of their ranges during the rest of the year”. Riley andDood statedthat “no relationship was evident between the point of captureand a fawnssummer home range”. They found that37% of the fawns were neverreobserved in their area of capture and that an additional22% werecaptured on the periphery of their summer range.My findings, and those ofthe others, support the idea that sites on the periphery of the dam’s homerange are used for fawning. This behavior conforms to the ‘hider strategy’model. Hiders are known to ingest the placenta and to remove theiryoungfrom the birth site to prevent predators from using olfactory cues to detecttheir young. If black-tailed deer were to give birth within their core areas of52use, they would be forced to remdve their fawns from the habitats they mostprefer. By giving birth on the periphery of their home ranges, female black-tailed deer avoid contaminating favorite areas and are able to move backinto them immediately after parturition.It could be argued that core areas are avoided during fawningbecause they are already laden with the scent of deer. This is not a likelyexplanation for two reasons. First, on Vancouver Island, black-tailed deer areubiquitous and their densities are usually high. By moving outside their coreareas to give birth, it would be difficult not to enter another deers core area.The known birth sites of animal 18181 were within an area that was heavilyused by animals 17912 and 177O1. Thus the dam’s movements did notremove her from areas with deer scent. Second, the study animals oftencontinued to frequent their core areas during the fawning period and theyusually brought their fawns into their core areas a few days after theirperipheral movement(s). It is doubtful therefore, that scent dissipation withinthe core area would have occurred before the return of the dam andoffspring.Average home range size was lower in the immediate post fawningperiod (PoNP1) than in all other periods except the pre-natal period (PrNP).Miller (1974) also found that “maternal black-tailed deer reduce the sizes oftheir home ranges during the post fawning period”. This behavior is alsoconsistent with the hider strateg’. During the period immediately afterfawning, the mother must remain close to her fawns which are sedentary.Large distances between foraging sites and the fawn’s hiding place wouldincrease the dam’s travel costs and risk of predation. By minimizing thisdistance, energy savings may be transferred to the fawns or used by thefemale. A reduction in travel would also allow the dam to remain motionless53and vigilant for greater periods and thereby reduce predation risks.Chapter Two suggested that in the earlier portions of the summer,habitat use was more concentrated in Submontane habitatswhere greenup occurred first. It is predictable therefore, that home range size should alsobe small during the pre-natal period (PrNP).54Chapter Four: Summary and ConclusionsChanges in Habitat Use In Relation to Forage QualityThe habitat use patterns of 13 radio collared black-tailed deer wereinvestigated in the summers of 1990 (June 1-October 31) and 1991 (April 1-October 31). Because the location gathering in 1990 starled late, these datawere not statistically tested. They were, however, used to substantiate the1991 findings. The summers were subdivided into time periods and thepercent use of various habitat types were compared. The use of seral stage,biogeoclimatic ecosystem varianls, elevation and aspect within the timeperiods was examined.The data sets of the nine study animals that had clearcut habitatswithin their summer ranges were pooled. Their use of clearcuts was highest inthe same period that Rochelle (1980) found fireweed to receive its highestuse. Percent use of clearcuts was 67.9% in April/May/June, 78.2% inJuly/August and 69.2% in September/October. Given that fireweed is anearly successional species and that it is the black-tailed deer’s mostimporlant summer forage species (Rochelle 1980), the relationship is notsurprising. I suggest that the availability of fireweed was not higher in theJuly/August period than in the other periods. Increased clearcut usetherefore, was probably in response to the nutritional values (digestibility andcaloric) of fireweed which peak in August.The use of Montane variants (BEC) was higher in the Later portions ofthe summer. The pooled data set for the eight deer that had Montanehabitats within their summer range revealed that the percent use ofMontane was 14.5% in April/May, 22.6% in June/July and 25.8%August/September. The use of Montane habitats therefore, was lowest when55the forage species within Submontane were more phenologicallyadvancedand of greater nutritional value. As the summer progressedand as foragephenology within the Montane habitats advanced, the useof Montaneincreased.Although the distribution of Montane variants over the landscapeisprimarily a function of elevation and aspect, changes in the useof the lattertwo habitat variables were not detected. With respect to elevation,the lackof a significant difference may have been an arlifactof sample size.Changes in Habitat use During the Fawning SeasonDuring the summer of 1991, ten adult females were monitored throughfive periods producing a total of 563 locations.The summer was divided intothe following 29-day periods: the pre-natal period (PrNP), May 8 to June5;the fawning period (Fr), June 6 to July4; and three post-natal periods,(PoNP1) July 5 to August 2, (PoNP2) August3 to September 1, and (PoNP3)September 2 to September 30.Unlike other researchers who have found relationships between natalsites and early seral stages, the use of clearcut habitats didnot changeduring the fawning period. Forest canopy and shrub layer heterogeneitycould have allowed the parturiént study animals to meet their coverrequirements within forested habitats. Many pockets of dense ground coverwere present in areas of low stand density and many species of shrubs, suchas red huckleberry, devil’s club and salal, often thrive in the second and oldgrowth forests of coastal British Columbia.Home range size was greater in the fawning period than in any otherperiod. Increases in home range size during the fawning period were causedby increases in the magnitude and/or rate of movements to the peripheriesof the home ranges. This finding supports the idea that sites on the periphery56of the dam’s home range are used for fawning and is consistent with the‘hider strategy’ model. Hiders are known to remove their young from thebirth site to prevent predators from using olfactory cues to detect theiryoung. If black-tailed deer were to give birth within their core areas of use,they would be forced to remove their fawns from the habitats they mostprefer.Home range size was lower in the immediate post fawning period(P0NP1) than in all other periods except the pre-natal period. This behavior isalso consistent with the hider strategy. During the period immediately afterfawning, the mother must remain close to her fawns which are sedentary.Large distances between foraging sites and the fawn’s hiding place wouldincrease the dam’s travel costs and risk of predation.In the early portion of the summer, habitat use was moreconcentrated in the Submontane where green-up occurs first. The homerange sizes in the pre-natal period therefore, should also be small.Recommendations and Implicatiofls1) In the logged watersheds of coastal British Columbia fireweed andother early successional species öre important summer forage species forblack-tailed deer. 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