Spatial Behaviour and Habitat Utilbation of Wapiti (&ruus eirrph~s)in the French River and Burwash Regions of Ontario

by

Glen S. Brown

A thesis subrnitted in partial fulfillment of the requirements for the degree of Master of Science in Biology

School of Graduate Studies and Research Lauremian University Sudbury, Ontario

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The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fkom it Ni la thèse ni des extraits substantiels may be printed or othewise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. This thesis is dedicated to my fàrnïJy for their encouragement throughout the projed. Acknowltdgments

nianks to my supavisors Dr. Frank F. Maliory and Dr. Joseph Ham, Depariwat

of Biology, Laurentian University, for their support and guidance thrwghout the study

and Ivan Filion, Cambrian CoUege, for field assistance, manpower, apipment, and

financial support. 1 would also like to thank Dr. R Pitblado of rny Research Advisoiy

Comminee and my extemal advisor, Dr. J. FryxeU, Department of Zoo10gy, University of

Guelph- Randy Staples, OMNR, provided tccbmcd support and di- maps used to

perform spatial anaiyses. Dr. Naylor, OMNR, provided Forest Resourcc Inventory 0

data and ecosite classification software. Aerial radio tracking was enhriced by the

dedication and enthusiasm of pilot Roland Hemmer- ïhe Ontario Mimsay of Naturai

Reswrces, Department of National Defence, Ontario Federation of Angiers and Hunters,

Rocky Mountain Ek Foundation, Safan Club International (Ontario Chapta), Tembec

Forest Products, Parks Canada, and Sudbury Fish and Game Protectn-e Association provided finaricial support. Heli North Aviation, Atwood Island Lodge, and Hadey Bay

Marina provided aippon in kind. Fellow students providing assistance with the field worlc included; Monika Jost, Lynn Landriault, Andrea Tumer, Nelson Deschenes, Jeff

Zuchlinski, and Paul Jaworski). Sincere thanks to Marita and Seija Maiiory for their encouragement and hospitality in providing a borne away &om home. Fe,special thanks to my dear nieads, Sin1 Wade Wallace, Michelle Zander, Kristai Conroy, and my favorite Wushu kids at the local YMC4 for keeping me sane during tbc writing of my thesis.

List of Figures

Figure 1.

Figure 2. Annual and seasonal ranges for the Burwash adult mak Y6 during 1996 usbg the adaptive kemel(95 % annuai home range and 30 % seasonal core activity areas) ad minimum convex polygon (100 % annuai home range) methOd~...... 30

Figure 3. Annual and seasonal ranges for the Burwash aduh fernale Y3 in 19%using the adaptive kemel(95 % lumusl home range and 30 % seasonal core activity areas) and minimum convex pdygon (95 % amml bme mge) methods...... 3 1

Figure 4. Application of the adaptive kernel and minimum conva polygon methods to deliaeate the 1996 annual home range and seasonal wre activity areas for the adult male B9 of the fiver ~~~dd-.~..-..----*-dddddddddddddddddddddddddddddddddddddddddddddddddddddddddddddddddddd34

Figure 5. Application of the adaptive kernei and minimum convex polygon methods to deliwate the 1996 annuai home range and scasonal core activity areas for the yeariing femde B8 of the French River herd*...... 35

Figure 6. Proportions of annual and seasonal borne ranges of aduh females Y2 and Y3 of the Burwash herd which overlapped in conseutive years ...... 39

Figure 7. Ropodons of iill~~uaiand seasonal ranges which overLpped in successive years fÔr four fdes(BI, B3, B5, and B8) in the FmdI River .....-.-...... 40

Figure 8. Mean proportion of each habitat type in seasonal cm aaivity araof females (adult. yeariing) and a dedf Bmdraon...... 44

Figure 9. Proportions of each habitat type in semonal and uimiJ 1996 30% core activity areas (ADK)for the adult mak Y6 ...... 45 Figure 10. Mean proportions of different habitats in seasdnal areactivity areas of adult fedes and ahes in the French River region 47 List of Tabk5 Page

Table 4. Approkte dates of rnovement between summer ranges no& of the French River and wuiter ranges south of the rivq------.---.------38 Spatial behaviour and brbitat utilbation by dk, Chusdqphs, in fbe

French River .ad Burnash regioms of centrd Ontario.

Glen S. Brown, Department of Biology, Lawentian University, Sudbury7ON, P3E 2C6

Abstract

This study investigated home range characteristics, habitat sdection, and the

impact of snow accumulation on elk, Cervus e@h, spatial bebaviour in the French

River and Burwash regions of central Ontario. Results were inconclusive as to seasonal

and gender differences in the site of home ranges and core activity areais. In the French

River region, the mean annual home range size (95% adaptke kernd method) was 128.92

km2+/- 43 -40 SD (a = 3) for addt femaies, and 2 12.74 bn2 (~2)for adult males. The

mean annuai home range size at Burwasà was 34.12 km2 for two adult fernales and 43.28

km2 for an adult male. There was no evidence of migratory behaviour within the Burwash

herd. In contrast, French River anunals migrateci between surnmw and winter ranges.

Collared animals in both herds exhiiited fidelity to home ranges and migratory French

River animais rehinieed to seasonai ranges. Elk selected some habitats in a greater

proportion than avaiiaôk. Stands of poplar, white bkh, white spruce?balsam fk were commonly selected in the Buruash region Selection of oprock areas rnay have been of benefit in winter due to reduced snow depth. Conifr dommatad black spmce-white pine stands were also seiected and may have been important as sbdta, deavoidance of grass and meadow bab'rtats was apparent in the Burwash ara. Habhat seiection was les evïdent in French River animais. Bkkopnice-white pine stands, ad wbite plie-rd pine- white spruce-white buch stands were selected throughout the year. kasonal changes in aspect use and proximity to water ouggested thaî snow depth influaiaal spatiai beh.viour in this region The utilintion of soutb dopes was highest in mid-wins (January-March), when snow accumulation was greatest and declined to a minimum in hme and July. The relative fiequency of dk locations on south fahg dopes was signindy greater than on no* facing slopes durhg the *ter pend, but not in the aimmer. LFtilization of south slopes was probabiy associatexi witb greater exposure to the sua and horde snow conditions. Elk were found to be si@cantiy closer to water during hte winter than during early winter. Frozen water bodies were used as travei corridors and were of greatest importance duriag peiods of deep snow. Knowledge of the spatial behaviow of a species is basic to our understanding of its ecology and the implementation of sustainable management practices. Analysis of spatial behaviour at the landscape scde cornmonly includes delineating annuai home ranges, seasonai ranges, migratory behaviur, home range fidelity, and habitat selection. Burt

(1943) defined home range as %ut area traversed by the individual in its' normal activities of food gatherïng, mating, and caring for young". The most commonly used measure of home range is the minimum convex polygon method (MCP) which encloses a range area by comecting ail outlying point locations for an animal (Moh, 1947). Bun (1943) suggested that the home range shouid not include every are-the animal has ever roarned, but rather those areas it "nonnally" visits when perfonning &y-thy activities.

Consequently, visits to outiying areas shouid be excluded fiom home range estimation.

White and Garrott (1990) suggested that the home range should include only animal locations within a certain probability. Probability density fbnctions used to describe home ranges commonly contain 95% of the animai locations (Jemich and Turner, 1969; hm and Gipson, 1977). It should be noted that defining outliers as the 5% of animal locations which are firth- fiom the center of an estimated home range is stiil a subjective decision by the researcha and the potehtjai biological significance of these outlying movements should not be overiooked.

Research conducted on elk in the United States (Montana, Wyoming) indicate variability in the sUes of home ranges among dinerent populations. The wui uinuai home range size (minimum convex polygon) of 3 1 fedeelk in Chaniberiain Creek, Montana during two consecutive years was 45.06 km2 (Edge et al-,1985). The srnailest range was 16.12 km2 and the largest 100.10 hn2. Estimated home ranges of three fernale eik in Madison

Drainage, Wyoming, ranged between 15.61 km2 and 30.72 km2(Craighead er d.,1973).

Summer (21 June to 23 September) and fa (23 September to 22 December) seasonai ranges were similar in size to the total home range, while winter (22 December to 20

March) and spring (2 1 March to 2 1 lune) ranges represented only a smaiî portion of the total home range in this population. Summer and fa11 ranges varïed between 13.53 kmz and 21 -33 km2, winter ranges varied between 0.26 km2and 2.34 km2, while spring ranges varied between 3.38 km2 and 6.24 km2. Twenty-one adult fedes studied at Jackson

Hole, Wyoming, had a mean summer range size of 12.49 km2, with a minimum size of

2.60 km2 and a maximum of 32-47 km2 (Martinka, 1969), while the summer home range for a single adult male was 12-49km2. Edge et al- (1985) suggested that home range size was duenced by food availabiüty, ambient temperature, biting insects, and availability of wver. According to ZNvin and Peek (1983), plant composition and forage density were important factors detennining home range size and location, whiie social relationships and population dmplayed a secondary role. Redation and human-induced disturbances such as huntiag might ais0 be iduential.

Werences between maies and females in home range skmay be associated with foraging and reproductive strategies. Beier and McCullough (1990) hypothesized that male white-tailed deer, OdocoIleus Wginiams, used areas of Iowa forage quality than females end thenfore required Iarger home ranges to mmnutritional requirements. In contrast, Gcist (1982) suggcsted that bull elk shouid select srmima ranges, which provide high Quality forage to replenish fat resemes dedfor the large energy expendinires dunng the rut and wimer. Red deer stags cm travel long distances to rutting grounds

(Clutton-Brock et ai., 1982) and Beier and McCullough (1990) noted that male white- tailed darranged widely during non-rutting seamns to learn the distribution and reproductive status of females and rival males. BuU eik in Michigan also had signiticantly

Iarger home ranges than females during both the rut and non-M pefiods (Beyer, 1987).

White and Garrott (1990) defineci migration as "a regular, round-trip movement of individuals between two or more areas or seasonal ranges." The occurrence of migratory behaviour within a herd may be influenced by seasonal changes in food availability, and the avoidance of predators du~gcalving, rutting and Mnter periods (Alcock, 1993). Many eUc populations in North Arnerica migrate between seasonal ranges (Altmann, 1952;

Craighead et al., 1972; Morgantini and Hudson, 1988). One of the most extensive migrations documented occurs in the Jackson Hole ek herd, which migrates approximately 100 km nom summer ranges in Grand Tetm Narionai Park and Bridger-

Teton National Forest to winter ranges in the Natiod Elk Refuge (Boyce, 1989).

However, both migratory and non-migratory behaviour has been documented in tbis herd

(Martinka, 1%9). Boyce (1 989) mggesteci that the primary -ors idluencing the Miuig and pattern of migration within the Jackson Hole herd were snow accumulation and hunting. Brazda (1953) found that parturition was a more imporiant factor than plant phenology in infiuencing the spring migration of cows in the Gallatin River Drainage,

Montana-

Herds of migrating elk ofien show fidelity to seasonal ranges (Murie, 195 1; Altmann,

1952; Morgantini and Hudson, 1988). Fidelity refers to the teadency of an animal to retum to an area previousty occupid or to remah m an area for an extended period of time (White and Garrott, 1990). EU< are known to show fidelity to seasonal ranges*as well as, specific sites witbin the range, such as caiving sites or mtting gromds (Brazd*

1953; Craighead et al-, 1973)- Edge et ol. (1 986) found strong range fideiity among non- migratory groups of cows in the Chamberlain Creek area of western Montana Less than

5% of the cows dispersed nom the traditional herd range. Wapiti in BmfFNational Park exhïïited predictable movement patterns during migration and a tendency to return to the same ranges each year (Morsamini and Hudson, 1988). Smith and Robbins (1994) docurnented strong site fidelity among radio-coilared adult elk that sw~neredin

Yellowstone National Park. Of 11 coiiared elk, 10 retumed to the same drainage or mountain cornplex each iaimmer. One possible expldon for this behaviour could be fideiîty to matrilined ranges by offspring. Clutton-Brock et al. (1982) observed that fernale red deer (Ce= ewk)on the Island of Rhum often occupied ranges and core areas which overlapped those of their mothers.

Habitat utili;rrition by elk in western North America has been weil documented

(Collins and Umess, 1983; Marcum and Scott, 1985; Wydeven and Dahlgren, 1985;

McCorquodale et aL, 1986; Edge et (11.- 1987; Edge et al., 1988; Peck and Pe* 1991).

Ek bave been found in a variety of habitats, hclucüng coastal rain forests (W~trnerand decalesta, 1983; Schmer et d.. 1993), montane regions (Maraim and Scott, 1985;

Morgantini and Hudson, 1988). ad opai grarslaad plains (Skovlui, 1982; Peck and Peek,

1991). Habii selected in cornrl rain for- of the Olympic Perurinsula, Washiagton, varied fiom young deciduais fores&, dominaîed by rai alda (Alrms mbra) and willows

(Solix spp.), coniferouddeciduous forests, dominatd by Sitka spruce (Picea sitckms), black cottonwood (Popuhû trf~hacafpa)and bigleaf maple (AmnraaopbZIam), to dense coderous forests, dominated by Sitka spruce and western hemlock (Tsugu heterophyk). Montane regions selected by eik in BanffNational Park, Alberta, consisted of alpine and subalpine habitat, with forests dominated by Engelman spmce (Picea engehiz] and alpine fir (Abies iizsî~)).White spruce (Picea glatra) and lodgepole pine (Pimcs conforta) were cornmon at lower elevations. Fexue (Festuca scabreih) grasslands were comrnonly utïijzed dong montane river valleys. Grassland habitats used by elk at Jackson Hole, Wyoming, consisted of sagebnish/bunchgrasscomplexes

(Artemrrio tridentam, KmIena cristata, and Festuca idahamsis) and sedge/bluegrass

(Cmer spp. and Pm spp.) complexes (Maninka, 1969)- Although desert areas appear to be avoided, McCorqude et al-(1 986) documented the use of arid shrub-steppe habitats in the state of Washington. Boyce (1989) found that ekappear to be generalists in the use of plant comrnunities and wncluded that detaiied site specific information was unlikely to improve the species' habitat characterization.

Bryant and Maser (1982) presented information on the distribution of elk in North

America and noted that mos? populations were associated with coniferous forests extending across western Canada and the United States. Forest cover is thought to provide themregulatory benefits as well as, shelter fiom predation (BeaU, 1974; Skoviiq

1982). Factors that appear to inauence habitat selection by elk include food avaiiabiiity, food quality, shelter/escape cover, elevation, slope inclination and aspect, snow conditions, ambient temperature, avaiiability of water, as well as, rutting and calving sites

(Skoviïn, 1982; Collins and Urnes, 1983; Grova and Thompson, 1986). Edge et al.

(1987) stressed the importance of examinhg the interaction of îhese fictors, when attempting to daennine primary factors influencing eik habitat dection Although the spatial behaviour of elk in the United Statés and western Canada has been well documeented, Iittle is known about the elk populations in eastern Canada.

Historidy, the eastern ekCervus dnRsdlt~~s~ inhabited much of southeasteni Ontario and southern Quebec. However, due to hunting and habitat degradation, eUc were extirpated fiom the region. One of the iast Ontario animais was shot near North Bay in 1893 (Seton, 1927). The elk populations currently residing northeast of Georgian Bay, Lake Huron, are remnants of a feintroduction of Rocky

Mountain eliq Cemis dmsnelsoni, brought from Alberta in the 1930's. Many of these animals were destroyed in the 1950's by the Ontario Department of Lands and

Forests, due to the fact that they harboured giant liver fluke, F~ioIoidesmagnu, hannful to Livestock (Ranta, 1979). However, a dpopulation escaped and has persisted until the present. The total area inhabited by elk in French River anà Bunvash regions has not increased since the late 1950's. Little was known about the spatiai behaviour of these mimals, although interest in them has increased during the past 20 years. Ranta (1 979) provided an historical account of theû introduction in the 1930's and exarnined elkfmoose habitat relationships in the Bwash-French River area based on sightings, repons fiom hunters, trappers, and 104 residents and aeriaUground surveys conductecl fiom 1976 to

1978.

The high mortality of French River and Burwash ek due to Pain coilisions in winter and drowning during movement dong fiozen water bodies suggested that mow conditions iduenced spatial behaviour and habitat utilization (J. Hamr, pers. comm.).

Snow depths of 40 and 70 an caud elk in Colorado to move to areas of lower snow depth (Sweayr and Sweaiey, 1984) and suow deptbs ova 70 cm impaired movement. Peck and Peek (1991) concluded that msting and snow depths pater than 30 cm causeci

ek to rnove fiom grasslands to shmbdominsted habitats. Huggard (1993) found that

predation on elk by wolves increased 60m 1 animPV5.4 days wnh no snow, to 1/1.1 days,

when snow depth reached 60 cm. Snow depths in the French River and Burwash regions

fiequently exceeded 40 cm- Snow accumulation couid influence the rnovement of animals

to water bodies, as open windswept lakes and watenvays provide pater rnobility,

visibiiity, access to new forage and hcilitate escape 6om predators. El.were apected to

be associated with water bodies more in late wimer, than in eady winter and the snow-fk

pends. South-facing dopes were ai- expezted to be used at greater fiequency during winter, as snow depth wouid be reduced and snow compaction Ïncreasxi, due to greater sunlight and higher temperatures (Beall, 1974).

For this study it was hypotheskd that:

1. anwal and seasonal home ranges and core activity areas of

male and female elk in the French River and Burwash herds were similar in site,

2. ek in both herds migrate between winter and summer ranges,

3. elk exhibit fideiity to home ranges and areactivity areas,

4. elk select ail habitat types disproportionately to their adability, and

5. snow accumulation affects the location of ek relative to water bodies and dope

=pect. Materials and Metbods

Study Arta

The study area was located in the Great Lakes - St. Lawrence Ecotonal Forest

Region (Rowe, 1WZ), ~~rnrnodyassociateci with mixeci conifer and hardwood habitats

(Chambers et ai., 1996). The two populations of eik monitored in this study occupied two

distinct regions nie Bumash eik herd was found approàmateiy 35 km northeast of

Georgian Bay in Secord, Laura, Burwash, and Senos Townships, in Sudbury District i 1 The French River population occupied an area approxhtely 20 km northeast of

Georgian Bay, in Kiipatrîck, Stmthers, and Travers Townships and the French River

Provincial Park, in Sudbury and Pany Sound Districts.

Glaciation bas created a topography characterized by gdeslopes intersperd with outcrops of granitic bedrock (Vankat, 1979). Some areas, partidarly in the French

River region, have been completely denuded of soil, creating banai bedrock plains

(Clayton et ai., 1977). Soils in the region Vary between -01 (acidic with a dark layer of organic matter over a grey leached axa) and podsolic forms, which are less acidic, less susceptible to leaching, and contain les organic xnatter. Acidic podzols usually nippon coniferous forests, wbile les acidic podu,lic soils support deciduous stands (Vankat,

1979).

Climate in this region bas been descri'bed as continental with a moderating effax in areas near Georgian Bay @anta, 1979). Jdy is the warmest mamh of the y=, with Figwe 1 Burwash and French River elk ranges northeast of Georgian Bay, Lake Huron, Ontario.

a mean monthly temperature of 18.5 OC(Anonymous, 1994, 1995. 1996). January is the

coldest month, with a mean monthly temperature of - 15.5 OCh-een 1994 and 1997.

Total annual precipitation (snowfall and rainfdi) varied between 9 1.1 cm in 1994 and 97.9

cm in 1996. Saowfall typicaîiy commenced during the second to îast week of November

and continueci to rnid April. Snow depth was greatest in Febniar)-. averaging 4 1.6 1 cm

between 1994 and 1997 and the total mean annual snofiwas 237-5 cm between 1994

and 1997. Total annual snowfall was uousuaiiy high (300.9 cm) during 1996. Long term

records indicated that July was the wamest month of the year witb a mean temperature of

15 OC (Vankat, 1979), while January was the wldea month, with mean temperatures

commonly between 14 OC and 11 OC (Chambers et al., 1996). Mean annual precipitation

was 75-100 cm (Vankat, 1979).

The Burwash herd inhabiteci a forested region south of Sudbury, bordered in the east by a major highway (Hwy 69 South) and bisected centrally by the Wanapitei River.

Runnîng adjacent to the river and through the center of the range was a Canadian National

RaiIway he, which was used several times ddy. The northem part of the study area containeci a number of laLes (750-2500 m2), many dermanbes, and flooded land.

Elevations rang4 betwem 200 m dong the shores of the Wanapitei River and 282 m on adjacent hdis and ridges. The followïng descriptions of floriac communities in the region are based on the Forest Ecosystem Cldcation Ecosites for centra) Ontario

(Chambers et al., 1997). The study area was defincd by deiineating a 5 km buffer zone around the outer teiernetxy locations of aU elk monitored between January 1995 and

March 1997. Within this 324.7 km2areq Ecosites 12. 14, and 16 (Central Ontario Forest

Ecosystem Classification) ocarpied the greatm proportion of the land base. Ecosite 12

(1 5.3% of the study area) consisting of poplar and aspen (Populus tremuloicies, P. grandjrtentia, and P. balsamvera), white birch (Betula ppyïîera), white sprue (Picea glaum), and balsam fir (Abies baheiz) stands, ocairs mostly on xeric and mesic soils.

These stands characteristicaiiy had trembfing aspen and white birch in the canopy, with white spruce and balsam fir in the understory. Common understory species includeû northem bush honeysuckle (Diemilla lonicera), beaked hazel (Cory2u.s cormitu), low sweet bluebeq (Vaccinium angusrioIium), wild sanaparilla (Aralia mdcûuIis), large- leaved aster (Aster macrophyllus), star fiower (Tfientalis borealis), bluebead iiiy

(Clintonia borealis), and brackm fan (Pteridîum aquilimrm).

Ecosite 16, consibg of white pine ( Pims sb.obtcs), largetooth aspen (Poplus grmddentaza)),and rdoak (Queras mbra) ocaipied 14.40/0 of the study area. White pine was the dominant canopy species, with red oak, white pine, red maple (Acer rubnun), and balsam fir ocairring in the understory Other comrnon plants in Ewsite 16 included beaked hiud and fly honeysuckle (Lonicero dnns),whüe bracken fem, wild lily of the dey(Maianthemm mmdme), wild sarsaparilla, large-leaved aster, rice gras

(0gvropszs aqen~olia),star fiower, wintergnai (Grnitheria pruambens), and bunchberry (ComCuMCILtems) were present in the understory.

The third most common cover type, Ecosite 14, occupied 10.W of the study area. The mrin canopy -es were white pine, red pine, white spmce, and white birch.

White spmce, balsam and rdmple fiequentiy oaamed in the subcanopy and regeneration layen, while beaked hwl mountain maple (Acer viwîum), fly honeysuckle, low sweet blueberry, and twinflower (LimeaboreaIs) were the common shnibs.

Bluebead Uy, wild sarsapda, wild Wy of the dey, bracken fern, large-Ieaved aster, ground pine (Lycoporiium&n&oicieum), and goldthread (Coptis tnyoIia) were also present. A more detailed description of all ecosite types, includiag soil, shband ground layer species can be found in Appendix E.

Open rock ridges and plateaus supporthg rdoak and red maple were also cornmon throughout the Burwash study ana. Along Hwy 69 Sou* on the eastem edge of the study area were a senes of abandoned fmfields (approx. 7.4 km2) once used to maintain the former Burwash Comectional Centre. These fàcilities bad not been used since 1975 and the fields were occupied primarily by grasses (Pm spp., Deschampsia spp., Agrosîis spp.) and sedges (Cmex spp.).

French River Range

The French River herd occupied foreas bordering the northeast shore of GeorgiaB

Bay, Lake Hm.Much of this region was covered with sdto medium skdlaLes, and river channels, cretks, marshes, and beaver ponds. Elevations in the area ranged between

185 m in the lowland marshes to 236 m on upiand plaîeaus. The French River composed of many fàst, narrow channefs separated the northern fiom the southem parts of the range ig1 A tributary, the Wanapitae River, flowed through the wrtheastern seaion of the range and entemû the French River at the northeast enâ of Kkg's Island. Bauing advity was high on the French River in dationwith lodges and cottages. Although few paved roads existed within the French River area, the northem component of the range

was interspersed with a network of abandoned logging roads and traiis, commonly used by

recreationists and hunters.

The French River study area was dehed by delineating a 10 km bder zone

around the outer telemetry locations of aU elk monïtored between March, 1994 and Aprii,

1997. This bufFer was chosen rather than a 5 hbuffer, due to the fact that French River animais ranged firther than the Bwash animais- The study ara encompasseci 1 418.8 la2;however, 285.1 h2were occupied by Georgian Bay and were unavdable to eik.

Habitat south of the French River consisted of a series of islands separated by the narrow, fast flowing cbannels of the estuary. The Iargest island, King's Island, was characterized by sphagnum (Sphagmrm spp.) bogs interspersed with tamarack (Drrir bicim)and black spmce (Picea rnm-anq) in the interior. Bordering the island were ndges dominated by white pine and jack pine (Pinus ~~),while eastem white cedar (&ja occi&ntaizs) lined the shoreline. The northern end of the island was characterized by upland plateau of mkedwwd white pine stands. The oventory was cornposeci prirnarily of white piw, red maple (Acer mbmm), jack pine, and black spruce.

The region north of the French River was also dominated by white pine mixedwood stands. Stands of trembling as- containing white pine, red maple, white spruce, balsam fir, and white birch were also cornmon. Within the 1 133 -7 km2 Prea available to the ekEcosites 14, 16, and 17 ocaipied the greatest proportion of the land base. However, small laLes ocaipied the second kgest proportion of this ara (1 1.5% or

129.9 km2). Ecosites 14 (1 7?/. of the study area) and 16 (1 0.6%) were desctl'bed previously for the Burwash range. Ecosite 17 ocaipied 7.9 % of the French River range and was dominated by white and rdpine stands. Such stands were characterîzed by

white pine and red pine in the main canopy, the absence of a subcanopy and red maple,

white pine, and balsam fir in the regeneration level. Shbspresmt in Ecosite 17 included;

beaked hazel low swat blueberry, and northern bush honeysuckle, while wintergreen

(GuuItheriaprocumbem),twinflower, wild lily of the valiey, wiid sarsapariiia, bracken

fem, large-leaved aster, rice grass, bunchbeny, and starflower were cornmon in the

ground layer.

Capture Techniques

During the winters of 1994 to 1997, ek fiom both herds were captured and irnmobilized by darting with the use of a helicopter and a Simmons-MS CAP-CHURrifie

(J. Hamr, pers. comm.). The dosage and mixture of tranquiiizers was 1 000 mg of powdereâ Ketamine, dissolveci in 6 cc of Rompun (100 mghi) and 1 cc of Hyonate

(sodium hyaluronate, 10 mg/ml). Hyonate was used to speed up the action of the tranquilizers. The same dosage and mixture was effective in immobilizing calves of both sexes and female adults. Additionai injections had to be administered to large males once they were partiaîiy sedated. Eik were fitted with convanional VHF radio-wllars (Mode1

LMRT 4) rnanuIactured by htek Electronics Inc., Aurora, Ontario and flexible livestock ear tags (AUflex hc.). Dropoff radio-collan modified with surgical tubing or bicycle inner tube were used on males of ail rges to prevent restriction of breathing du~gthe rut, when the neck increases in size (Lincoin, 1971). Eik were hjezted with antibiotics

(Peniciliin) and Dystosel (seleniun and vitamin E) to prevent pst-capture myopathy. The effed of sedatives was reversed with Yohimbine (Antagonil) prior to release. In total, 16

ekwere fitîed with radio coiiars, 6 fkom the Burwash herd and 10 fiom the French River

Herd. These numbers were estirnated to represent approximately 20 % and 25% of the

Burwash and French River Herds, respedvely (J. Hamr, pers. comm.). The capture

dates, periods of surveillance, and causes of mortality of radio-coiiared animals are

presented in Table 1. Individual animais were monitored for periods ranging between 22

and 1,126 days, where monitoring period relers to the number of days between the first

and last radio location obtained for an animal. However, 4 radio-ailar losses and 6

mortalities precluded deteminhg annual adorseasonal ranges for some animals.

Monitoring Techniques

Each radio-coliared elk was located 2-3 timedweek by trianguiation, using a three- element directional Yagi antenna or a 2 element H-antenna, a portable receiver-scanner

(Model SRX40, Lotek Inc.), 1: 5û,OOû topographic maps and a Silva compass (Model

#15). High granite ridges and deep ravines often renilted in refiedon of the signai. As a rd~an attempt was made to obtah a minimum of thne bearings for each location to avoid the use of erroneous bearings and to estimate error (White and Garrott, 1990).

However, on occasion only two bearings were avaiîable due to technical and logistic problems. The time period baween each bearing varied Eom 10 to 60 minutes. Bearbgs were subsequentiy recorded on 1:5O,ûûû topographic mpps and converted to Universal

Transverse Mercator (UTM) coordinates, rncorded to an acawcy of 100 m. In addition to location, the identification number of each animal, date, and tirne were recorde-. Table 1. Surveillance periods, s~rvi'~al,and causes of mortality for radio-collnred clk of the Burwash and French River herds betwcen

1994 and 1997. l

Animal Sex Age(yrs) Capture Date Surveillance Period Mortality/Survival (ai end of study)

French River Herd BI F I adult 4 years alive B2 F I 8-9 1 month drowned B3 F 1 1.5 2.5 years drow ned B4 M 1 adult 7 rnonths unknown (dropped collar) 85 F i 1.5 3 months drowned B6 M 0.5 1 rnonth st or ved B7 F 2.5 1 .S months drow ned B8 F 0.5 I year starved B9 M adult 6.4 nionths unknown (dropped collar) BI0 M adult 9 months alive i Burwash Herd i i Y 1 M 0.5 2.5 years al i ve i Y2 F adult 2.5 years alive Y3 F adult 2.5 years dead (unknown cause) Y4 F 1.5 1.5 months train-killed Y5 F i 0.5 1.5 months starved Y6 M , adult 6 months unknown Ground Tracking

Radio-tracking of each animai occu~edweekly or biweekly, dependiq upon

iogistics and season. Locations of the Bwash animais were rnonitored fiom elevated

points along grave1 roads, power transmission lines, and ridges used year-round. Duruig

the summer, the French River herd fiequented the Bird Lake region. Access to coflared

animais was obtaïned by using an &-terrain vehicle or hiking along abandoneci logging

roads or trails. However, during the fa and winter months, access to these anùnals

required using motor bats and snowmobiIes in the French River estiand trachg was

accomplished fiom elevated locations along the shoreline of King's Island and the adjacent

maidand. During the spring, aumner, and fall of each year, 1-5 locations were obtaineà

for each animal every 1-2 weeks. Locations could not be obtained for the French River

animais during the edy winter (mid-December to eariy Febmary) and early spring (late

March to late April) due to the fkeeze-up and thaw periods, which precluded access.

Aerid Trackinq

Ground tracking was supplemented with aerial trachg during mid winter (January

and February) and the summer (June to August). Dunng January and Febniary (1995-

1997), a M

tracking of WWanimais. During the merof 1995, aeziai tracking was performed

once a month using a Muiistry of Naturai Resourcts fixed-wing Turbo-Beaver aircd and biweekly during the armmer of 1996, using a priva* owned Cessia 172. Each aircrafl was outfitted with two-element, unidirectional Yagi anteimas, which were attacheci to wiag stnits or helicopter skids and Med to a receiver-scanner and right-l& switch box

Duriog raial tracking, once a radio sipi had been detectcd the aircrafi was directeci towards the strongest signal. High density forest uinopy ofhmemt that visual

observations of animals codd not be obtained to ver@ location during the snow 6ee

period. As a result, locations were obtained by flying a circle around the animal, while

keeping îhe strongest signal in the center of the circle. Lunited flight tirne prevented the

researchers fiom terting the accuracy of the aerial location estimata using a fixeci-wing

aircraff; however, one instance occurred where a coiiar dropped by a bu1 was retrieved

approximately 100 rn fiom the location estabiished by aerial tracking- Winter tracking

with helicopter invariably resulted in visual locations of aliared animais.

AnalysQ OC Home Range Sizc and Movement Patterns

Annual and seasonal home ranges were calculateci for individual eik employing the

minimum convex polygon method @%oh, 1947) and the adaptive kemel method described

by Worton (1989). The home range cornputer program Calhome (Kre et al., 1996) was

used to caldate home ranges. The minimum convex polygon wtbod has been widely used for measuring home range size and was included in the present study for comparative pwposes. This method involves connecting the outer telemetry locations of each animal to create a convex polygon, the ana of which is subsequentiy calailatecl (White and

Garrott, 1990). In spite of the widespread use of this technique, a number of drawbacks

Limiting the interpretation of home nnge boundaries and areas oca~~The fïrst problern is that as the number of locations hcrease, the eStjmated skof the borne range increases, due to the fâct that this technique measures the entire area utilUrd by each animal (Jennrich and Turner, 1969). As a result, this method is Uifluenced disproportionately by outlying points. Secondly, an underlying assumption of the minimum convex polygon method is that the locations are nodydistri'buted within the home range. Such a non- biological approach does not wnsider the infiuence of the environment on the anunal.

Consequently, home range size may be greatiy overestimated by ïncluding areas which the animals never utilize. An example would be an animal whose home range extends around the penphery of a lake. Using the minimum convex polygon rnethod, one Mght fasely conclude that the lake represented a large proportion of the animals home range.

The adaptive kernel method of home range estimation is a non-pararnetric approach, which is free of the assunptions of the minimum wnvex polygon technique conceming a bivariate normal distri'bution (Mohr, 1947; Worton, 1989). As a remît, this method identifies the important non-cunvex features of the home range. The adaptive kemel estimator is a probability density Fundon, which is plad over each data location point. Consequently, areas with a high concentration of locations have a higher de* kernel estimate than those with fewer locations. This feature prevents outliers fiom beïng included in the home range polygon The smoothhg parameter of the kernel estimate controls the amount of variation in each kemel (Worton, 1989). Thus, kemel esbmates with a low h value have lesvariance and show the finer detail in the anunal location &ta, comparexi to kemel estimata with higher h values, which have pater variance and dow only coarser features to k obsewed (ie. the home range is "smoothed" more in these areas). The adaptive kdmethoci varies the srnootbhg parmeter causing areas with a high concentration of observations to have lower h values than those with fewer observations. In other words, areas with fewer animal observations are smwthed, white areas with a high concentration of observations show the berdetails of where an animal

spends most time (Worton, 1989).

To Merunderstand habitat utilizattion, home ranges were calculateà for each

animal thrwghout the entire year, as weii as each season. Annual home ranges (adaptive

kemel method) were calculated using telernetry data fiom the entire year, based on 95 %

of the telemetry locations of each Mimal. Seasonal home ranges were calculateci using the

95 % adaptive kemel rnethod, mcluding telemetry data coiiected each season (winter,

sp~g,summer, and auh>mn). Seasonal core activity area represented a range cdcdated

with the adaptive kernel method, which includeci 30 % of the telemetry locations of an

individual animal for each season. Fitty percent core activity areas were also caldateci.

Seasons were deheated as: Wuiter (1 November-3 1 March); Spdg (1 April-3 1 May);

Sumrner (1 June-3 0 Augusî); and Au- (1 September-3 1 October). Site fidelity was

also investigated. Due to SIMU sample size, fidelity was not measuced by statisticai

methods, but was assesseci by overlaying home ranges of individual eik du~gdifîerent

seawns and years and me-gthe proportion of overlap. Methods used to enimate

error in telemetry foilowed procedures descnbed by Macdonald et al. (1980) and hduigs

are presented in Appendices 4 B, C, and D.

Habitat Anrlysis

Digitid ûntario Base Maps (OBM)and Forest Resource Inventory Maps (RU) were obtained fkom the Ministry of Natural Resources to analyze the re1ationships between the raâio location datl and landscape-de habitat amies.Four spatial data sets of landscape variables were used in the analyses. These include; the original FR1 data

on canopy species composition, ecosite forest classification types, dope aspect, and

distance to water.

Prior to performing habitat analyses, the original digitized maps were transformed into a format which aliowed for manipulation in GIS software, ARCINFO and ARCVIEW

(ESRI, Inc.). The FR1 data on forest canopy composition were subsequentiy re- categorized according to the "ecosite classification system" developed for ~tralOntario

(Chambers et al., 1997). The "ecosite forest classi6cation7'for each forested stand was subsequently Wedtn the respective FR1 "forest coveragey' polygon ushg ARCINFO.

Forest coverage polygoas were digitai representations of forested stands and contained various aîtriiutes describig the composition of the stand (eg. canopy species, stand height, age, and stocking).

Habitat utilkation by coilareci animas was analyzed using two dinerent techniques.

The fist method involved using point locations of each animal to test for habitat selection.

The second method involved examining seasonal trends in habitat use based on both the proportions of difFerent habitat types within 30% seasonal core activity areas. Although

Johnson (1980) descni habitat usage as the quantity of a resorircc type utüized by an animal in a ked period, habitat usage as referred to in the second analysis was describeci in tenns of the proportion of a resource within a specified area, used by an animai. Habitat Selection

The first method of habitat analysis involved using point locati011~of each animal to descnie the seleaion of specinc ecasite types. Measuruig habitat deaion required quantifjhg the utilkation of dinerent habitats, as compared to availability. Amrding to

Johnson (1980), seleaion would be demonstrated if habitat types were used disproportionately to availability. Ek habitat selection was determined by overlaying the focations of ali collared elk ont0 the "forest wverage" polygons of the FRI maps. This methodology aliowed each eik location to be digitally linked to stand specific FR1 attniute data, including an ecosite forest classi6cation.

Assessing selection by elk for partidar habitat types required measuriag the availability and utiiization of each habitat by individuai animais. For this analysis habitat type was based on forest ecosite classitications. A Chi-square test (Naet ai-,1974) was used to determine whether elk selected habitat types equal to, grater than, or in proportion to availability. This method was preferred over other Chi-square tests

(Friedman, 1937; Johnson, 1980), due to the srnali sample of collared animals. In addition, the method of Neu et al. (1974) asrumed that exact meaSUTes of habitat adabiiity were possible. The use of digitized maps in the present study aliowed for precise delineation of available habitat. To reduce subjectinty, available habitat was delineated by placing a 5 km buffa around the overd range of coiIared Bwash elk and a

1O lan bufk around the overd range of coîiared French Riva ek Herd ranges were deterrnined by using the minimum convex polygon method and inchidcd al animai locations coiiected between 1994 and 1997. It shouid be noted, however, that using home range to define habitat âvaiiability introduced bias, as animals had already shown some

selection in choosing the area (White and Garrot- 1990).

Availability of each forest -site class~cationtype was determineci by calculating the proportion of the 5 km (Bwwash berd) or 10 km (French River herd) buffered herd range occupied by each ecosite type. Utilkation was determined by caldating the fkquencies and proportions of the locations for each animal in a spedc ecosite. The expected number of locations in each ecosite was caldated by muliiplying the total number of locations for an individuai animal by the proportion of the buffered herd range ocaipied by each ecosite. This number of locations would ocau in each ecosite type, if no habitat preference or avoidance was shown by the study animais.

A Chi-square goodness-of-fit test with k - 1 degrees of fieedom, where k was the total number of ecosite types was subsequently used to test the hypothesis, tbat habitat ufili7i1tion occurred in proportion to availability, when al1 ecosite types were considered simultaneously. Ifthis test proved signifiant, suggesting dection or avoidance of some habitats, Bonfernoni confidence intervals were caldated for the proportion of tune an animai utilized each ecosite (Neu et al., 1974). The intervais were dculated using the fo rmuia:

(3) p, - z- @&pi)lnlLn <=pl <=p, + zd@&p,)h]* wherepi was the proportion of point locations in each =site (r) ad z- was the

Bonferroni Z statistic- Determination of ecosite avoidance and seldon was accomplished by wmparing the utilkation confidence intavals of each ecosïte with the avaiiabiiity proportions of each ecosite in the Wered ovdherd ranges. Selection above availability for a particuiar habitat would k demonstrateci, $the proportion of the herd range ocaipied by that habitat (availabiiity propohon) was lesthan the lower Limit

of the utiiization confidence interval. Selection less than in proportion to availabiiity

(avoidance) would k demonstrpted, if the proportion was greater thn the upper iimit.

Summing the Chi-square statistïcs for al1 animais to test overd habitat selection by both

herds was not possible due to small sample size and lack of independence arnong coiîared

elk. Because an increa~ein the number of available habitats reduced the power of

analfis, the rmmber of habitat cetegories was decrdby combinïng ecosite types witb

similar compositions of canopy species. Ecosite types and species compositions are listed

in Appendix E.

To assess seasonal chaages in habitat utïhttion, annual and seasonal home ranges were calculated for individual eik employing the adaptive kemel mahod described by

Wonon (1989). Seasonal habitat utili;ration was examuieci by overkying the annual and seasonal core activity areas of each animal on the OBM and FRI aiaps and descnig the proportions of areas ocaipied by each mate type-

Im~actof Snow -th - Distance to Water. and Selection of Slom As-

The locations for each animai were aiso used to detmnM dection of dope aspezt, distance to water, and the respoase to saow depth. Eacb locaîion was ovedain on the contour coverage of the appropriate OBM mrp and the dope aspect of the animai location was vinialiy estimated. Six tategories were used to record siope aspect: south

(S), including southeast and southwest; north (N), including northeast and northwest; west

(W); east (E); O, ifthe dope of a location was not deteztable on the contour map; and X,

if aspect wuid not be detennined due to a higb concentration of contours oumounding the

location. Topography was sunilar in both the French River and Burwash regions, allowing data to be pooled. Monthiy changes in the relative use Gequencies of slope aspect were examined. In addition, the Wdwxon's signed-ranks test for two groups (Sokal and Rohlf

1995) was used to test for sigiilfiûuit dinerences in the use of south facing slopes in

surnmer (June, My,and August) versus winter (Novernber, December, January? February, and March). Usage was defined as the relative fiequency of locations on south-facing slopes (south, southwest, and southeast) out of ail summer or wuiter locations.

Wdcoxon's signed ranks test was also used to test for ciifferences in the use of south versus north slopes in winter and summer. A Kruskal-Waiiis One Way Analysis of

Variance was used to examine the relationship between slope aspect uthtion and snow depth during the periods of mow accumulation (November to April). Data were pooled for the yean 1994 to 1997.

To measure the distance of ek to water bodies, telemetry locations were overlain on the appropriate digital OBM drainage in ARCVIEW. Three distance meantres were calculated: (1) the distance to the closest open water (Me, pond, or river), (2) the distance to the closest mush or hg,and (3) the distance to the closer of the two. Animai location data wae divided into 6 subseasons: early winter (Novemkr 1 - Janq 15), latte winter (Jmuary 15 - March 3 l), spring (April 1 - May 3 l), eariy summer (June 1 - July

1S), late summer (July 16 - August 30)' and autumn (Sept- 1 - ûctober 3 1). To determine trends in the distribution of elk, a set of randorn locations was generated withui the buffered herd ranges. A Mann-Whitney U test was used to check for dinerences in the distances to water bodies of dompoints and the actual animai loatioas throughout ali seasons. Analyses were paforwd separately for the Bunvash and French River data; however, data for ail ariimals within each herd were pooled, Trends were descri'bed relative to changes in snow deph over the respective the perïods. Monthly snow data for the mtdy area were obtained fkom the Sudbury Airport weather station (Anonymous,

1 994- 1997).

hiring the three year study, 958 telemetry locations were obtained for 16 eik fitted with radio wiiars (6 in the Burwash herd and 10 in the French Riva herd). The radio coilared ek were monitored for peziods ranging between 22 and 1,126 days. However, it should 'oe noted that an unexpected high mortaiiîy due to drowning vain collisions, and starvation reduced sample sizes, such that hypotheses on home range characteristics, migratory behaviour, fidelity, and seasonal habitai utiihtion wuld not be tested. For this reason, the rdtsof these components of the thesis are by necessity descriptive. Home Range Size and ksond Corn Activity Aras

Burwash Herd

Annuai home range sizes and seasonal core activity areas for aU collared Burwash

animals are presented in Table 2. The srnailest annual home range (decaKY 1) was

27.34 km2,and the large* annual home range (adult male Y6) was 43.28 km2 (Fig. 2).

During 1995 and 1996, seasonal core activity areas were srna representing between O. 1 -

12 % of tbe annuai home ranges. The smallest core activity area (female calf YS) was less than 0.01 bnt during the 1996 winter (Table 2). ïhe largest core activity area (adult female Y3) was 4.14 km2,during the same der(Fig. 3). Due to smaii sample sue, statistical cornparisons of home ranges based on swonand gender were not possible.

French River Herd

SufEcient radio locations were obtained to deiineate annual home ranges of adult females BI, B3, and B5, ywhg female B8, and adult male B9. Annual and seasonai ranges of coiiared eik in the French River region between 1994 and 1997 are presented in

Table 3. The mean annual home range sïze of duit females in the French River herd between 1994 and 19% rnegsu~ed128.92 km2 (n = 3, SD = 43 -40). The annd home range of adult male B4 measwed 220.87 km2and that of ad& male B9.204.6 1 km2(Fig.

4). These were the largest home ranges rewrded in either bad. In 1996, yearling fede

B8 had the destannd home range (1 9.17 kmZ)of any inimal (Fig. 5). Due to niiall sample size seasonal trends in home range site couid not be determuied. Among aii Table 2. Amal and seasonal ranges (kmz) of collared elk in the Bwash region between 1995 and 1997. Ranges include 95% estimates and 3W and SC% core aaivity areas obtained by the adaptive kemel meîbod (ADK), as well as minimum convex polygon (MCP) eshates. Blaak ceils indicate periods during whkh animals lacked a radio coiiar.

Range (km') Locations (%) Animal Y2 Y3 Y4 YS Y6 Annual95 ADKQS% ADKSO% ADK30°r6 MCP100% Annual96 ADK9S% ADKSO% ADK30°h MCPlW% ADK9S0h ADKSO% ADK30°h MCPl00% AQK95% ADKSO% ADK30% MCPl 00% ADK95% ADKSO% ADK30% MCPl00% ADK95% ADKSO% ADK30°h MCP1OOOh ADK9S0r6 ADKSO% ADK30% MCPlOOOh ADK95% ADKSO% ADK30% MCPlOO% ADK95% ADKSOOh ADK30% MCP100% Table 2. Continued

Range (kmz) Locations (%) Animal Y1 Y2 Y3 Y4 Y5 Y6 W96 ADK95% 37 28 17 1 14 ADKSO% 3 7 1 cl 1 ADK30% 1 4 cl cl 4 MCPl00% 34 25 7 4 5 W97 ADK95% 28 5 ADKSû% 1 <1 ADK30% 4 <1 MCPl00% 15 3

Annual - Range estimate based on the whoie year (January - December). Sp - Range estirnate based the spring period (1 April - 3 I May)- S - Range estimate based on the çummer period (1 June - 14 September). A - Range estimate based on the autumn rutting perd (1 5 September - 3 l October). W - Range estimate based on the winter period (1 November - 3 1 March). Figure 2. Annual and seasonal ranges for Bwash aduh male Y6 duhg 1996 uMgthe adaptive kd(95 % annuai home nnge and 30 % seasonal wre active areas) and minimum convex polygon (100 % annual home range) methods.

Figure 3. Annuai and seasonal home ranges for Burwash adult femde Y3 in 1996 using the adaptive kernel(95 % annual home range and 30 % seasonal core activity areas) and miaiaiun convex poiygon (95 % anaual home range) methods.

Table 3. Annual and seasonal ranges (km2)of collami eik in the French River region betw- 1994 and 1997. Ranges include 95% estimates and and 50% core activity areas obtaiaed by the adaptive kernel method (ADK), as weU as minllnum convex poiygon (MCP) estimates. Blank ceiis indicate perïods during which animals lacked a radio collar, NA indicates an insuilïcient number of point locations.

Range (km2) Locations (%) Animal

AOK95% ADKSO% ADK30% MCPlûû% ADK95% ADKSOOh ADK3O% MCPI 00% ADK95% ADKSO% ADK30% MCPl00% ADK9S% ADKSO% ADK30% MCP100% ADK95% ADKSO% ADK30% MCPl00% ADKQS% ADKSO% ADK30% MCPlOW ADK95% ADKSCW ADK30% MCPl ûû% ADK9S% ADKSO% ADK30% MCPlW% ADK95% ADKSO% ADK30% MCPl00% ADK95% ADKSO% ADK30% Table 3. Continued

Range (km2) Locations (96) Animal BI A95 AOK95% 104 AûK50% 1 ADK30% 4 MCPl00% 41 ADK95% 33 ADKSO% 13 ADK30% 2 MCPlOW! 17 ADK95% 14 ADKSO% cl ADK30% cl MCPlOWh 10 ADK9S% 22 ADKSO% 7 ADK30% 2 MCPl00% 12 ADK95% 43 ADKSO% 8 ADK30% 2 MCP100% 30 W97 ADK95% 8 1 ADKSO% 43 ADK30% 21 MCP100% 53

Annual - Range esbmate based on the whole year (Jmuary - December). Sp - Range estimate based on the spring period (1 Aprd - 3 1 May). S - Range estimate based on the surnmer penod (1 June - 14 September). A - Range estimate based on the autumn mtting penod (1 5 September - 3 1 October). W - Range estirnate based on the winter period (1 November - 3 1 March) . Figure 4. Application of the adaptive kemel and minimum cornpolygon mahods to delineate the 1996 muai home range adseasonal core activity areas for adult deB9 of the French River kd.

Figure 5. Application of the adaptive Lerne1 and minimum convex polygon methods to deiineate the 19% mual home range and seasonai core activity areas for the yearling fernale B8 of the French River herd. animais, core actiMty areas ranged between 0.05 % and 14 % of the annual home range.

The smallest areactivity area was less than 0.01 km2 for the 1996 spring range of yearling female B8 (Fig. 5). The largest con activity aru was 27.1 1 km2for the 1995 sp~grange of the adult female B 1. Due to the srnall sample size and high variance, no significant Merences in the size of seasonal core activity areas were observed.

There was no evidence of migratory behaviour in the Burwasb herd, however, srnall sample site precluded rnaking soiid conclusions. Considerable overlap existed in seasonal home ranges and the longest movement was 4.3 1 km between surnmer and winter in 1996, by adult male Y6. This was the only Burwash animal for which winter and summer home ranges did not overlap. There was a tendency for aii animais to utilire different core activity areas among seasons.

Distriions of core activity areas and home ranges and the timing of seasonal rnovements indicated that most animas in the French River region migrated between distinct merand winter ranges. An exception wu yearling female B8, who spent the spring, surnmer, and autumn of 1996 on a peninsula at Crombie Bay. Six of the mual home ranges were separated into hwo distinct seasonal polygons: a *ter home range south of the French River and summer home range north of the French Riva, aear Bi.

Lake (Fig. 4, e.g. for aduh maie B9). None of the winter home ranges overiapped with sumwr ranges. The mean distance banaumner and winter COBactivity areas was 10.4 +/- 3.2 km (N = 3) for adult femaies. The mean distance betweetl winter and sumrner core activity areas was 12.3 km for two adult males.

The Iocations of spring and auturnn core activiîy areas indicated indirect movement between surnmer and winter ranges. Sp~gcore activity areas of Mtfernales were either on iate winter ranges or near surnmer ranges. Five out of eight autumn ranges were located in the central portion of King's Island, where rutting activity was observed. Dates of movement between summer and winter ranges varied among anhds and years.

Autumn movements to islands south ofthe river occured between September 1 and

November 20 (Table 4). Spring movements to summer ranges north of the French River omedbetween Mar& 12 aad June 1. The estimated timing of spMg movement was less accurate due to logistic problems causeci by spring thaw (Iate March to late April), which Limiteci access to the river and resulted in longer pends between successive telemeîxy locations.

Overlap in home ranges iadicated that coilareci animais in both herds had fideiity to annuai and seasonal ranges (Figs. 6,7). Seasonal dinerences in fideiity couid not be determined due to dsample size. However, French River uiimrls exhiiited notable patterns in the spatiai distribution of seasonal ranges. AU lomipls wintered on isiands south of the French River, primuily King's Island and ~ummerexiwrth of the river in the vicinity of Bird and Loon Lakes. The majority of au- core activity areas were located Tab!e 4. Approximate dates of movement between summer ranges nonh of the French River and winter ranges south of the river.

- -- - Animal Year Movement soutb of river Movement north of river B1 1994 Sept.17 - Oct, 2 Apr. 19 - May 2 1995 Sept. 1 - Sept. 15 Apr. 5 - Apr. 20 1996 Sept. 26 - Oct 4 Apr. 28 - May 9 B3 1995 Oct. 13-Oc~2O Apr. 5 - Apr. 20 19% Sept. 26 - 0ct 4 Apr. 28 - May 9 B5 19% Sept. 26 - 0ct- 4 Apr. 28 - May 9 B8 1996 Nov. 13 - Nov. 20 March 12 - apr. 10 B9 1996 Apr. 28 - May 14 Figure 6. Proportions of annual and seasonal home mges of PUlt fdesY2 and Y3 of the Bunuash herd which overlapped in wdveyears. Each range represents a 95% home range estimated dngthe adaptive kernel method.

Annual - Range estimate based on the whole year (January - December). Sp - Range estimate based on the spring period (1 Apd - 3 1 May). S - Range estimate based on the surnmer period (1 June - 14 September). A - Range estimate based on the autumn ruttïng paiod (1 5 Septernber - 15 October). W - Range estimate based on the winter period (16 ûctober - 3 1 March).

Figure 7. Proportions of annuai and seasonai home ranges whid~overtapped in successive years for four females (Bi, B3, BS, and B8) in the French River region. Ranges represent 95% home ranges estimateci using the aàaptive Lerne1 method. Proportion estimata showing no ovdap between ranges were given a value of O. 1 to differentiate them firom cases where no annual cornparisons could be made due to a lack of data. instances of the latter were given a value of zero.

Annuai - Range estimate based on the whole year (January - December). Sp - Range estimate based on the spring period (1 Apd - 3 1 May). S - Range estimate based on the summer period (1 June - 14 September). A - Range estimate based on the au- rutting period (1 5 September - 1 5 October). W - Range estimate based oo the winter period (16 October - 3 1 March). at the nitting site on King's Island. Rutting activity in this area was confirmeci by the occurrence of tree scraping and bugiing.

Habitat Anaiyscs

Habitat Utili;ration and Selection

Resuits of the Chi-square analyses supported the hypothesis that ekin the region selected some habitat types disproportionately to availability. Habitat selection was more pronounced in Bucwash than in French River animals. Due to the mure of the andytical procedure and the limited nwnber of telemetry locations, data for fidualanimals were pooled for all seasons and years. In the Burwash herd, adequate daut were avdable to pefionn the -ses for two addt females (Y2 and Y3), the addt de(Y6), the yearling fernale (Y4), and the male calf(Y1) (Appendix F). In the French River herd, analysis was perfomed for the three adult fedes (B 1, B3, and BS), two addt males (B4 and B9), and one yeariing fede(B8) (Appendu G).

Burwash animais selected ecosites 1 I (popladwhite birch) d 12 (popladwhite birch, white spruce, balsam iir) significantly more than expened hmtheir availaôility.

This was dernonstrated by two addt females (Y2 and Y3), the addi de(Y6). and the male caif(Y1). Seleztion of black sprucdwhite pine stands (eco9tc 22) and rock outcrop habitat (MNR code 3 13) was also cxhiibited. Ecosite types which orne Eigmficaatly avoided by individuals inciuded; merbodies (MNR des62/64/152), rmclasJified bufk areas adjacent to water or otha habitat types ( MNR codes 302/303), trecd or open muskeg (MNRcodes 3 10/3 1 l), bmh and alder (MNR code 3 12), grass and meadow habitats (MNR codes 3 19316), lowland hardwoods with eastern white cedar (Ecosite

5/6), white pine-red pine-white spnice-white birch stands (Ecosite 14), eastern white cedar-white birch-white spnice-white pine habitats (Ecosite 1S), and white pine-red pine- jack pine stands (Ecosites 171 18/19). Many more habitat types were avoided than are desm'bed (Appendix F). These habitats were not used by coiiared animals due to low avdability, which resulted in zero values for the Bonferroni confidence intemals and were omitted fiom the discussion-

In the French River are% an addt female @ 1) and an adult male (B9) selected black spmce-white pine stands (ecosite 22), and yearling fernale B8 setected white pine- red pine-white spruce-white birch stands (ecosite 14) significantly more than expected from their availabiity (Appendk G). Two adult femdes and an Mt male avoided water

(MNR code 62/64/152) and one adult fde(BS) showed no selection or avoidance of any ecosite types. Habitat selection andyses are presented in Appendices F and G. Due to smaii sample size, it is unknown whether seledon of these hab'itat types was common among other members of these herds.

Seasonal habitat utilization was analyzed by overlaying each animais' annual and seasonal core activity areas on the relevant OBM and FRI maps and cornputhg the proportion of area occupied by each -site type. Among ail laimrls in the BurwaJh region, ecosite 12 @oplar/white birch, white spruce, balsam fir) was utilized wnsistently

in the core activity areas (Fig. 8). Amang winter areactjvity areas of females (adult

and yearling) and calves, ecosite 12 represented the hi- proportion of range area

utilized (A= 0.24, SD = 0.18, N = 5). Ecosite 16 (wbite pine-largetooth aspen-red oak)

occupied the second highest mean proportion of winter core activity areas among females and calves (3 = 0.14, SD = 0.09, N = S), followed by open rock areas (3 = 0.1 1, SD =

0.1 1, N = 5). Notabiy, open rock areas dominateci the winta care activity area inhabiteci by Y 1 in 1995. Exceptions to the dominance of ecosite 12 within winter core activity areas of individual aninials included lakes (MNR code 64) and black spruce/white pine stands (ecosite 22). Eastern white cedar-white birch-white spruce-white pine (ecosite 15) occupied the highest proportion of the winter core activity ara of adult male Y6 in 1996

(Fig. 9).

Sp~gcore activity areas of females and calves (2= 0.62, SD = 0.18, N = 5), as well as, an adult male were dominated by ecosite 12 (Figs. 8, 9). Ecosite 12 occupied the greatea proportion of surmer core activity areas for all animals (mde caif Y 1, addt females Y2 and Y3, and adult male Y6) between 1995 and 1996 (Fig. 8, 9). Autumn core activity areas were characterized by ecosite 12, as wdl as, black spruce - white pine stands

(ecosite 22). Among -sites which were present in &vdy bigh propanious, no signifiant composition Merences were found between seasons, wbm all animais were pooled.

In the French Riva region, ecosites 14 (white pner'd pine-white spnice-white birch stands), 19 (white pine-red pine-jack pine), and 22 (bkk spnice-white piw) Figure 8. Mean proportion of each habitat type in seasonal core activity areas of femaies (duit, yearbg) and a male calfin the Bwasb region. Refa to Appendix E for descriptions of each habitat type. Habitat type descriptions were based on MNR codes for non-forested habitats and ecosite types for forested habitats.

Figure 9. Proportions of each habitat type in suwnal and annual 1996 300/. core activity areas for the adult mate Y6. 'Avaiiable' refers to the proportion of the 3 24.743 ldBurwash study area occupied by dinerent ecosites. Habitat type descriptions are given in Appendix E and based on MNR codes for non-forested habitats and ecosite types for foresteci habitats.

were utiiized consistently in the seasonal core activity areas. Wmter core activity areas of

adult fernales and calves in the French River region were doxninated by ecosite 22 (black

spmce-white pine) (proportion: 2 = 0.27, SD = 0.30, N = 5) and -site 19 (white pine-

red pine-jack pine) was second most dominant (3= 0.20, SD = 0.27, N = 5), foUowed

by ecosite 17 (white pine-red pine) (2 = 0.11, SD = 0.18, N = 5) (Fig. 10). Among three

addt males, open rock habitat occupied the greatest proportion of winter core activity

areas (3 = 0.38, SD= 0.40), followed by lakes (MNR code 64, 2 = 0.24, SD = 0.3), and

white pine-red pine-jack pine stands (-site 19, 3 = 0.22, SD = 0.4) (Fig. 1 1). Open

rock habitat was utiiized most by yearling fernale B8 during 1996 (Fig. 12); however,

ecosite 14 (white pine-red pine-white spruce-white birch) ocaipied the greatest propodon

of her 1997 winter core activity area.

Among spring core activity areas, ewsite 14 (white phe-red pine-white spnice- white birch) occupied the greatest mean proportion of adult federanges, foiiowed by ecosite 22 (black spruce-mixedwood) (Fig. 10). The sp~gcore activity area of yearling fedeB8 was al- dominated by ecosite 14 (Fig. 12). Ecosites 5 (lowland hardwoods- black ash-poplar-mixed hardwood) (2 = 0.24) and 19 (white pine-rd pine-jack pine) (7

= 0.17) occupied the highest mean proportion of spring core activity areas of adult males

B4 and B9 (Fig. 11).

Summer core activitty areas of adult femaies were dominated by ecosite 14 (white pine-red pine-white spruce-white birch) (proportion: 3 = 0.62, SD = 0.12XFig. 10). Both ecosites 14 and 22 ocaipied bigh man proportions of ~nmmcure activity areas of aduh maies (Fig. 1l), whüe yculuig fdeB8 utüwd white pheAargetooth aspcn/red oak stands (ecosite 16) the most (Fig 12). Figure 10. Mean proportions of dinerent habitats in seuod wre activity anas of adult fernales and calves in the Fr& River region Hab& type descriptions are givm in Appendix E, based on MNR codes for non-forested habitais and ecosite types for forested habiis. LLE

B LE

ZLE Figure 1 1. Mean proportions of mirent habitats in seasonal core activity areas of three adult desin the French River region. Aunimn habitat utilization could only be described for aduit male B9 (1996). Habitat type descriptions are givm in Appeiidix E and bwd on MNR codes for non-forested habitats and ecosite types for foresteci habitats.

Figure 12. Reportions of different habitat types in sawd and annual 1996 30% core activity areas for yeariing fernale B8. 'Available' refers to the proportion of the 14 18.769 kmz French River study area ocaipied by dinerent ecosites. Habitai type descriptions are given in Appendix E and were baseci on MNR codes for non-forested habitats and ecosite types for forested habitats.

Autumn core activity areas of adult females were dominate. by ecosite 22 (black spruce-rnixedwood) (propohon: 2 = 0.44, SD = 0.34, N = 3) (Fig. 10). Ecosite 14 occupied the greatest proportion of habitat utiIized by yearling fernale B8 and aduit male

B9 in 1996 (Figs. 11, 12). Ecosites 14 and 22 occurred in relatively high proponions within core activity areas. No significant merences in proportions of each ecosite were found between seasons however, whea data were pooled.

Impact of Snow Depth Distance to Water. and Seledon of Slope As~ect

Elk locations were used to determine selection of dope aspect and distance to water in relation to snow depth and habitat u-tion (Table 5). Monthly changes in utiiization of different slope aspects are presented in Figure 13. As topography was similar in both the French River and Burwash regions, data were pooled. The majority of locations were in areas lacking distinct aspect detectable on the 1:20, 000 OBM rnaps.

No relationship existed between aspect types on which animais were found between

November and April, and the mean snow depth on days when locations were obtained

(Kruskai-Wallis, = 6.03, p

Mean Snow Oeph Month 1994 1995 1996 1997 Depth N SD Depth N SD Depth N SD De@h N SO

Jan Feb Mar AP~ May June July AuS SePt Oct Nov Dec Figure 13. Monîhly changes in dope aspect utilization based on the relative fiequency of animal locations @ooled for both hads).

T = 2, p<0.005), but not the summer pend (Wiicoxon's signed ranks test: T = 15,

p>0.05) (Appendix H). Due to high variance, no relationship was found between daily

snow depth and selection of dope aspect-

Mean distances of random and telernetry locations to water bodies are presented in

Tables 6 and 7. Kolmogorov-Smimov test statistics revealed that the distance to water

data were not distributeci nody. As a result, a Mann-Whitney U test was used to

determine whether point ldonswere distributcd randomly with respect to water. AU

herd data were pooled for the analyses. in the Burwash region, telemetry point locations

were found to be significamly closer than random to water in early winter (U = 1873, p <

0.05) and spring (U = 6437, p < 0.05), significantly closer thaa random to marshes in late

winter (U = 760 1-5, p c 0.05), spring (U = 653 8, p < 0.05), and late summer (U = 1465.5,

p < 0.05), and significantly closer than random to either water or marshes in late summer

(U = 15 17, p < 0.05). No signifiant Merences were found between seasons in the distance of telemetry point locations to water bodies (Mann-Whitney U test, p<0.05).

In the French River region, telemetxy point locations were found to be significantly farther fiom marshes than random in spring (ü = 915, p < 0.05) and from water bodies (ü

= 1406, p < 0.05) in autumn. Locations were found to be significantly closer to water in late winter compareci to early winter (U = 1972.5, p < O.OS), early summer (U = 23 79.5, p

< O.OS), and late summa (U = 2158.5, pc0.05). Point locations were signScantly closer to marshes in earîy wimer than late summer (LJ = 1491, F0.05). Point locations were signincantly doser to the nearest water or manh in eariy wimer compared to early swimer (U = 1688.0, -.OS), early winter campareci to late summer (U = 139 1-5, Table 6. Descriptive statistics for the seasonai distances to water bodies for a random set of point locations in the Burwash (324.7 km2) and French River (1418.8 km2) study areas. Ln the HerdfSeason/Drainage column, herd identificaîion is listed as B (Burwash) or F (French River); Seosons are identified as Sp (spring), S1 (eariy summer), S2 (late summer), A (autumn), W 1 (eariy winter), and W2(late winter); and drainage types are identified as C (closest of either water or marsh), M (rnarsh), and W (water). Asterîcs indicate a sigrdicantly non-normal distribution (Kolmogorov-Smirnov test, p

Herd/Season/Drainage N Mean (m) Std. Deviation Minimum (m) Maximum (m)

BAC ' BAM ' BAW BS1C ' BSlM' BSlW BS2C ' BS2M ' BS2W BSPC ' BSPM ' BSPW BWIC* BW1M' BW1W BW2C ' BW2M ' BW2W FAC ' FAM FAW FSlC FSlM FS1W FS2C * FS2M ' FS2W FSPC ' FSPM * FSPW FWIC* FW1M RN1W' MC' FW2M ' MW' Table 7. Descriptive statistics for the seasonal distances to water body masures for teiemetry locations collecteci for animals in the Bwash and French River regions. In the HerdlSeason/Drainage wlumn, herd identification is tined as B (Bunuash) or F (French River); Seasons are identifid as Sp (spring), S 1 (eady summer), S2 (iate aunmer), A (auhunn), W 1 (early winter), and W2 (late winter); and drainage types are identifid as C (closest of either water or marsh), M (marsh), and W (water). Astencs (*) indicate a significantly non-random distribution (MW test, p<0.05). (") indicate a significantly non- normal distribution (Kohnogorov-Smhov test, p

Herd/Season/ürainage N Mean (m) Std. Deviation Minimum (m) Maximum (rn)

BAC A 71 95.6 102.2 O 558.9 BAM BAW BSlC A BS1M A BS1W BS2C * A BS2M * BSZW BSPC A BSPM 'A BSPW ' BW1C A BW1 M BW1W ' BWCA BW2M * A BW2W FAC FAM FAW ' FS1C A FS1 M A FS1W FS2C FS2M FS2W FSPC A FSPM ' FSPW Fwic FW1M A FW1W FW2C A FW2M A FW2WA p<0.05), late winter cornpared to early summer (W = 2379.5, p<0.05), and late winter compared to late summer (U = 2365.0, p<0.05).

Discussion

Home Ranges and Core Acîivity Areas

Due to smdi sarnple size solid conclusions could not be made regarding seasonal home range characteristics and the hypothesized Merence between male and fernale home range size. Preliminary results suggested that annuai home ranges ofelk in the French

River herd were generaüy larger than those reporied in the iiterature, whiie ranges of

Burwash animais were ssimilar to those of other studies (Cnighead ez ai., 1973;Edge et al., 1985).

Mimatory Behaviour

Migratory bebaviour is dehed as an annual round-trip movement between

Summer and winter ranges (White and Garrott, 1990). There was no evidence of migratory behaviour for adult fdesof the Bmash herd and the adult dewas the only animal for which winter and summer home ranges did not overiap. Alrnost d French

River animais made seasonai movements to a winter range south of the French River and summer range north of the river near Bird Lake. The one exception was a yeariiug fernale who spent the spring, summer, and autumn of 1996 on the penimda at Crombie Bay. Martida (1969) also documentcd migratory and non-migratory behaviour within a single

herd. Factors which may have caused the development of migratory behaviour in French

River anirnals include hunter activity?snow conditions, food availability, avaiiabiiity of

rutting and calving sites, and predation. Hunting and related actMties on fores roads

have been shown to affect elk mowment @ktbb,1969; Lyon, 1983; Edge and

Mar- 1985; Morgantini and Hudson, 1988; Smith and Robbins, 1994). Lyon (1983)

found evidence that vehicle tr&c on forest roads induced an avoidance response in elk.

Edge and Marcum (1985) obsaved that eUc moved away fiom logging disturbance and

returned to these areas during non-active periods. Martinka (1 969), Knighî (1WO), and

Morgantini and Hudson (1988) reported instances of elk moving Eom hunted areas to park and refûge areas, at the onset of the hun~gseason. During the autumn in the

French River region, the network of abandoned logging roads in the Nmmer range was used extensively by bear (Sept. 1 - Oct. 15) and moose (Oct. 5 - Nov. 15) hunters. Trac involving four wheel drive trucks and d-tenain vehicles was heavy during these periods and hounds were aiso used for bear hunting. The presence of humans within the summer range during this period may bave resulted in the development of migration behaviour in this population. Autumn movement of animals away from the aunmer range occuned between September 1 and October 20. During this period, the animais were most often found south of Bird and Loon Lakes, in more isolated areas close to the French River. In the Burwash region, limited bmtbg activity occw~edon the West side of the Wanapitei

River and the absence of dogs during the bear hunt may have comiuted to the sedentaq nature of local ek Snow accumulation may also have infiuenced migratory behanour in the French

River region. Moran (1973) suggested that elk in Michigan restricted movernents under conditions of deep snow. Adams (1982) concluded that snow depth was one of the most important factors iduencing the timing of autumn migration in this species. Roosevelt ek inhabiting mountainous regions in Washington (Schwartz and Mitchell, 1945) and Alaska

(Troyer, 1960) were thought to migrate fkom sunmer ranges to winter ranges in response to snow accumuiation. In my study, it was unLikely that migration was initiated by snow accumulation, as soowfd did not occur und mid November. Howwer, the island system provided access to fiozen waterways and marshes 4th more wmpacted, windblown snow during the winter pend The greater number of these openings on the French River islands, compareci to the forested areas on the summer range, wodd give ek greater mobility and access to forage. The majority of French River elk dersightings were dong waterways or openings.

In the Bwash region, snow accumulation may have contnbuted to the absence of elk migration. The ranges in the Burwash region contained a greater number of open rock ridges than the sumnier ranges of the French River animals. Examination of core activity areas of the French River animais indicated that 6 out of 7 winter ranges conuimd open rock areas, while only 1 out of 8 summer ranges contained similar habitats. In addition, the average proporiion of open rock areas in French River winter core activity areas was

15 % (N = 10, SD = 0.29), while in summer ranges it was ody 0.3 % (N = 8, SD = 0.0 1).

In the Burwash region, al1 &ter and summer core acbvity areas contained Mar proportions of oprock areas: winter, 10 % N = 10, SD = 0.1 1 and summer, 1 1 ./. N =

6, SD = 0.05. Nthough not sigaincant, the above proporbons suggested that open rock areas in the Burwash summer and winter ranges were more extensive than in the sumrner range of the Freùch River anirnals. nie higher availability of wind swept areas lacking deep snow in the Bunvash area may have eliminated the need for miption in this population. The majonty of wimer sigùtings at Burwash were on open ndges.

Dinerences in the availability of open rock habitat between French River and Bwash regions niggest that snow accumulation may be another cause of migration in the French

River herd. The moderathg efféct caused by Georgian Bay would ahreduce mow depths in the winter range of this herd.

Forage availability was another factor that rnay have influenced migration, partjcularly during the spring. Jost (1997) concludecl that plant communities inhabited by both French River and Burwasb herds were similar, however, forage was less dense on

French River winter range than on nimmer range. These observations suggest that seasonal changes in food avaiiabiïty may also have been a factor kfluencing spring migration in the French River animals.

Brazda (1953) found that partmition was a more important factor inûuencing the spring migration in cows than plant phenology in the Gallatin Riva Drainage, Montana.

The need to find deand secludecl sites for caiving could induce migration fiom winter ranges and possibly reduce the risk of predation. Movement of adult fernale eik north of the French River occureci between Aprii 5 and May 9 (1 994- 19%). Taber et ol. (1 982) observecl that peak partaon of severai Rocky Mountain ekpopulations occamed between mid May and eariy June. Smith and Rob- (1994) found that calving in the

Jackson Hole, Wyoming uea occurred during or after migration to the summa ranges.

Sirnilar observations were made for animais in the French River region; however, as calving did not initiate migration in the Bwash population, this fmor remains questionable. Even so, adult fernales in the Bwash herd did move into different areas for calving foilowing the winter perïod.

Predation may have influaid migration in the French Riva animals and minimized it in the Burwash popdation. Reports f?om French Riva trappers and hunters indicated that woif and bear populations were pater in the eUc sunmer range than the whter range. These observations suggest that predation may have iduenced migation of these animais. In contrast, wolfnumbers were considered to be ton-er in the Burwash region, which might explain the absence of migratory behavior in this herd.

Migration behaviour rnay also have evolved as part of the dispersai of ek fkom original release sites (Morgantbi and Hudson, 1988). According to Baker (1982), initial exploratory rnovements fiom original release areas rnay rdtin seasod ret.to these areas. Such seasonal movemems might subsequentiy be passed on to future generations.

Martïnka (1969) observed that movements of yearling male ek were restricted to the areas ompied by fende-caif goups, with which they were aswciazed in surmner.

Morgantini and Hudson (1988) concluded that dispersai and seasonal migration in inaoduced elk may be fàcilitated by associations with remnant groups of îndigenous elk.

Although it has been hypothesued that the eastem elk origidy inhabiteci King's Island during the eariy period of the ek reintroduction program in the 19303, no genetic evidence was found to substsntiPte this possibiiity. Genetic an- of the eik population in the French River and BurwsJh areas indiaitecl Sunüanty to the Roc@ Mountain sub@es of ek introduced in 1932 (Polziehn et d. in press). These 6ndings suggest that French River animale may have deveioped migratory behaviour der dispersal fkom the release site of the original population; however, it does not explain the absence of migration in Bmhmimals or the fàct that the French River population has no association with the Bwash popdation.

A number of studies on elk have reporte.fidelity to seasod and annual home ranges (Brazda, 1953; Knight, 1970; Edge et ai., 1985; Morgantini and Hudson, 1988).

Results supponed the hypothesis that animais fiom the Burwash ad French River herds exhibited fideiïty to annual home ranges. Results were inconclusive as to seasonai changes in range fidelity. Annual home range and seasonal range fidelity is probably advantageous to an animai, as daded knowledge of habitat and landscape charaaenstics, especiaily food, water, and cover, wouid increase survivorship (Edge et al., 1985; Beier and

McCullough, 1990). Knowledge of winter forage distribution would be especially beneficial during periods of deep snow and limited movement. In the French River winter range, white cedar on the shores of King's Island were excessively browseû over successive years and Jost (1997) concluded that white cedar was an important late winter browse associated with the edges of waterways where snow depth was reduced.

Habitat Utüization and Selection

Ek in the Bunuash adFrench River regions tended to sd&t certain habitat types more than in proponion to availabiiity. Peck aad Peek (1991) obtmied simiiar results for ekin British Columbia. Analyser of habitat selection in Burwash ammals suggmed that ek seiected stands containhg poplar, white birch, white spruce, and balsam tir. Open rock outcrops and biack spruce-dite pine stands were dso selected above availability.

Analysis of seasonal habitat utiliration indicated that core activitty arcas were composed primarily of poplars, white birch, white spruce, and baisam fir (ecosite 12). The selection of this forest ecosite was consistent throughout each season. Jost (1997) found that elk in the Burwash region foraged on myiv of the species present in ecoshe 12, including white birch, trembling aspen, rdmaple, northern bush honeysuckle, beaked hazel, large-leaved aster, and bluebead My. Beaked hazel, large-leaved aster, and bluebead iily were selected more than in proportion to availability in al1 mixed forest habitats. However, it should be noted that these food items were also present in many of the other forest habitat types in the study area. Factors other than food availability may have influenced the use of ecosite

12, such as protection fkom predation or human disturbance.

Open rock areas were selected by adult females in the Buma& region. The proportion of open rock areas selected witbin the seasonal core acthity areas did not indicate a distinct pattern in utîiization. It was initidy suspected thwind swept open rock habitats would benefit animais during derby providing gream mobEty due to reduced snow deph (Raata, 1979). Opem rock habitats ocaipied 30% of the 1997 winter core activity are. for adult fede Y2, a year when snow accumulation was greater than in

1995 and 19%. However, this habitat ocaipied a pater proportion of surnmer wre activity areas than the winter core activity anas during 1995 and 19%. This relaîionship also existeci for Mtfde Y3. The beaefits of nimmer utihation of open rock areas might inchde close proximjty to shdter, inadavaiiability of spccific food item, and possibly relief 60m bithg insects. Open rock areas were wmmonly associateci 4th ridge

habitat in the Bunvash region Jon (1997) concluded that ridge habitats were important

to Ontario eik in late sumrner and auturnn due to the availability of the selected food items

such as common hairgrass (Deschampsiaflexuosa), acorns, and mushroorns of the genus

SuiIIils. Ridge habitats were dso used in arly summer due to the presence of selected

browse items, such as balsam poplar and largetooth aspen. Juiander and JefEey (1964)

came to similar conclusions and found that ndge tops were the prefd sumrner range of

eUc in Utah. Moran (1973) indicated that ek in Michigan were clorly associated Mth

openings of ali types (old farm fields, abandoned lurnber camps, orchards, air strips) and

attributed their use to the availability of new browse and close proxhity to cover.

Black spruce-white pine stands (ecosite 22) were selected by elk du~gauhunn and winter. Peck and Peek (1 991) dso documented the use of conifer dominateci stands by elk during periods of severe weather and attnbuted their use to changes in forage availability related to snow accumulation Knight (1970) found that conifer dorninated stands, primarily Douglas fi (Pseuabtsuga ta#dia), were used in extreme winter weather conditions in Montam

Habitat selection was les cornmon in the French River region. One expIanation may be the fact that the French River animals migraîe Ween nimmer and winter ranges.

This resulted in the use of a wider varie of habitats, increasing the variance and reducing the power of the Chi-square and Boafenoni analyses. Among collared animals in the

French River area, one aduit fdeand one adult male selected black spruce-white pine stands (ecosite 22), while a yeariing fdeselected white pine-red pine-white spruce- white birch stands (ecosite 14). It should be noted that although the aduit fdedid not show signincantly greater selecrion for white pine-red phe-white spruce-white birch

stands (ecusite 14), Bonferroni confidence intervals hdicated a trend towards greater use

than availabile (Appendk G).

Resuits of the habitat analyses, based on seasonal core actRity areas suggested

that eik in the French River region utilized coder dominated stands (-sites 14 and 22)

throughout the year. Yearling female B8 was the only French River animal who selected

white pine-red pine-white spruce-white birch stands (ecosite 14) merthàn in proportion

to availability. Figure 11 indicates that ecosite 14 ocaipied a large proponïon of the study

area available to ek Statistidy, the greater the avaiiability of a habitat type, the less

Likely an animal wiii show selection greater than availability. Searonal range use indicated

that this animal was highly sedentary in nature! and had high fidelity to her annual home

range. As the majority of forested land on this range consisteci of ecosite 14, selection for

this habitat may be related to sedemary behaviour. This soiitary behaviour appeared to be

due to the animal's separation fiom its mother during the initial coüaring procedure. The

anunal never regaineci contact and remained close to the capture site for the rest of the

winter.

UnWre ecosite 14, aosite 22 occurred in a smaiier proportion of the study area,

suggesting that its selection by elk was important. As suggested previously, selection of

conifer domiaated stands in winter mybe beneficial during harsâ weather. Mom (1973) found that conifêr stands were of greater importance to elk during Jcvae winters in

Michigan. French River animais also used openings dong watemap rad marsh systems during winter. Mult deB9 was the only animal in the French Riva region which selected ecosite 22 as the greatest proportion of the summer cmdvity area. This bu11 resided in an area intersperseci with open and treed bogs. The periphery of these openings

was presumably used for browsing with the adjacent forested land providing shelter.

Forage species within black spnice-white pine stands were likely important in tbe

selection of ecosite 22. The fact that males and fdwselected different habitats during

different thes of the year may refiect diffbrent foraging strategies. Geist (1 982)

suggested that to maxi+ reproductive success, males shouid avoid competing with

femdes by seiecting dinerent feeding sites. In addition, males must feed on highiy

nutritious and digestibIe forage in order to store fat reserves in preparation for the autumn

rut. Of the species available as forage in black spruce-white pine stands, only white birch

and eastem white &were found to be selected more than available by elk in the French

River and Bmash regions (Jost, 1997). However, white birch and white cedar were

probably of lower nutritional value than new forbs and grasses (Robbins et al. 1987). The

availability of lichens in ecosite 22 may have aided in the digestion of white cedar and

other woody browse, as lichens improved fermentation efficiency and digestion in white-

tailed deer (Jenks and Leslie, 1988). Jost (1 997) found that the use of lichens ocairred in

Iate winter and coincided with the increased use of conifers, partiailarly white cedar. It remains unclear whether the availabiîity of preferred forage items was the most important hctor iduencbg selection of black spnice-white pine stands (-site 22); however, forage composition within these stands, suggests that these stands were more important as shelter.

Habitai avoidance (seiection les than available) was also documented in French

River and Burwash sfiininis. Avoidance of water bodies was demonstrated in collared animals fiom both h&. This was expected considering that the majority of wata Mes consist of open water througbout mon of the year. In addition, the numerous lakes and

waterways redted in a relatively high proportion of the study area king occupied by

water, relative to other non-forested habitats. However, the importance of water bodies

to elk as travels routes shouid be recognized- Elk in both regions were wmrnonly

observed on fiozen laJces and rivers during the winter period.

Elk in the Bucwash region appeared to avoid grass and meadow habitats. Many

studies in western North America have demonstrateci that ek utilize grasslands and

meadows extensively (Colluis and Umess, 1983; Wydeven and Dahlgren, 1985; Peck and

Peek 1991). Herding behaviour in open areas can serve as an antipredator strategy

hproving the ability of prey species to escape (Krebs and Davies, 1987). Franklui et al.

(1975) studyiag Roosevelt eik obsewed that group ske was iargest on open prairies and

srnaIlest in brush-land and forests; however, this phenornenon may be influenced by the

abiiity of grasslands to support higher densities of animals, within the same unit of area

(Clutton-Br* 1974). The apparent avoidance of gnssland habitat by local elk rnay have resulted fiom low population demities limiting the value of herding as an antipredator strategy or may be due to the fact that ek in tbis region avoided grasses of old world origin (Jost, 1997). In addition, telemetry data wae collecteci primarily during diumai hours, wbich would not idethe possible noaumil use of grasslands. A remote cament located m a meadow in the Bucwash region indicated that most activity by eik at this site occurred during aoaunial and crepuscular hours (J. Harnr?pen. comm.).

However, it should be noted that utilidon of this site was influenad by the presence of a sait block. Although seasonal changes in habitat seiection wuld not be determined, the composition of core activity areas Vaneci between seasons. Among dared anunals in both herds, winter core activity areas were most variable in terms of habitat type. Surnmer core activity areas had the lowest variance and autumn and spring ranges were intermediate. Dwing summer, the majority of core activity areas of French River animais were found in ecosite 14, containhg white pine, red pine, white spruce, and white birch.

Burwash animais consistently utilized ecosite 12 on summer ranges, dominated by popiar, white birch, white spmce, and balsam fir. CoUared anunals typically had iarger winter ranges than summer ranges, while spring and autumn ranges were intermediate. The size of seasonai core activity areas wouid duence the diversity of habitat types. Moran

(1973) suggested that Michigan ek made use of a greater diversity of habitat types in winter than in sumwr. hiring summer, Michigan elk utiiized openhgs containing hi& quality forage, while 4available food and cover types were used during winter. îhe use of a greater diversity of habitat types in spring and autumn would be expected in animais which migrate between summer aud winter ranges. Although coUared Buxwash anids did not Mgrate, they moved between seasonal core activity areas. These findiags do support the hypothesis that animals &iited habitat selection, as aninuiis would be expezted to show seasonai changes in the location of core activity areas. Monitoring ofa gnater number of animas would k required to determine if this relationsbip is common among elk in the region The lm~actof Snow on Distance to Water and Slo~eAspect Use

Seasonal changes in slope aspect utihtion suggested that snow accumulation may

have infiuenced elk spatial behaviour. Although significant differences were not found

among months, the fiequency of south slope use was highest in mid-winter (January-

March), dropped to a minimiim during June and July, and increased again between

September and DeCernber. Snow depths were highest between January and March., with

Febmary having the highest mean snow accumulation (up to 62.82 cm in 1997). Snow depths between 40 an and 70 cm in Colorado resulted in eUc moviag to areas of less snow

(Sweeney and Sweeney, 1984).

Although the resuits suggested an increased use of south facing slopes in winter, no signifiant merenas were found in south slope use be-n summer and winter. The relative fkquency of locations on south facing slopes was found to be signïficantiy greater than on north fhgslopes during the winter period, but not the summer period. Maraim and Scott (1985) obsetved a positive correlation between increased use of south and south-west slope aspects by elk in Montana and grrater annual cumulative precipiîation.

Skovh (1982) suggested that southerly aspects were aïticai to eik during winter, becaux of reduced snow accumulation and increased availability of forage. Beail(1974) noted that timbered north slopes in Montana were consistently colder with deeper snow than south faciag slopes, which received grtater atposure to sunlight. He also obsemd that elk moved to Merent aspects and cbangd position on dopes, based on meteorologicai conditions. Ek were obmed to use lower devations during conditions of dap sww, but moved to higher devations when snow accumulation was limited. EU( were also observed ushg south slopes exposeci to direct sunlight under extremely cold conditions and moving upslope and avoiding shadows by staying in diuect Sunlight during the evening sunset (Beall, 1974).

Ln spite of the above hdings, it shodd be noted that in the present study the majority of locations were found in areas lacking any distinct aspeet detectable on the

1:20, 000 OBM maps. This was meboth in summer and &a- No relationship existed between asped types on wbich animais were found between November and Apd, and the mean snow depth on days when locations were obtained. The absence of high elevations and steep slopes in the region may have reduced the thermoregulatoxy value of slope aspect.

The distance of point locations to water bodies was expected to be inauenced by snow accumulation. The signifjcantly non-random association of ek with water bodies in both regions suggested that selection was occurriag. Seasonai Merences in the proamity of animals to water bodies were more pronounced in the French River region than the

Burwash region. French River animais were sigdicantly closer to water bodies during winter, tha.during summer. The most obvious explanation for tbis difference wouid be the observe-migration to islands for the winter, placing anirnais closer to water.

However, locations were found to be significantiy closer to wata ciuring late winter, the period of greatest snow depth, than early winter. Locations of Freach River animnls were also signrficantly closer to marshes in edy winter than in late summer. Even the non- rnigrating Burwash animais were found sgnifïcantiy closer than domto wata in eady winter and spring, but not during ?ciimm~.. Other reasons for seasonal dinerences in elk proximïty to water may be annual

changes in precipitaîion, forage avaiiabïlity, parturition, and biting insects. During dry

summers, elk in Chamberiain Creek Montana, moved to mesic &es eariier than durùig wet sumrners (Matcwn and Scott, 1985). Although animais in the Bucwash region were sigruficantly closer than random to marshes in late as cornpareci to eariy nimmer, thae was no correlation with weather data. Paxturition might also influence the distance of animals to wata. Skovlin (1982) suggested that lactating cows may have an increased dependence on water during the spring calving period. Burwash animals were significantiy closer than random to water in spring, but not in summer or late wimer.

It is also possible that biting insects influence the distance of elk to rnarshes.

Johnson (1 95 1) aigge~redthat ek in the Gallatin Drainage Basin in Montana moved to higher, windy elevations to avoid high insect densities. Brazda (1953) noted an inverse relationship between insect and elk numbers in the same area. In the Bunvash region, elk were significantly closer than random to marshes in late summer, but not early summer.

Black fiies, SimuZim nigncolcum, and mosquitos, Anopheles spp., were common in the region and emerged in large numbers in June and July. Aithough the habitat analyses did not indiaite a seasonai selection of wata bodies based on the proportion of water within core activity areas, the &ta on distance to water indicated the importance of this ecologicai feature to ek. Summary

Although the rdîsof this study were linmeci by the small sample size caused by the high mortality in the two herds, the data support the wnclusions tbat:

(1) migratory behaviour was minimal among coked aNmds of the Bucwash herd as ody one adult male had discrete winter and summer home ranges, while most coilared eik in the French River region migrateci between non-overlapping summer and winter ranges;

(2) animals fiom the French River and Bunuash herds exhibited fideiity to annual home ranges.

(3) Bwash animais selected stands of poplar, white buch, white sprue, balsam fir (ecosites 12) and open rock habitat more than available;

(4) French River animais selected black spruce-white pine stands ( ecosite 22), and white pine-red pine-white spruce-white birch stands (ecosite 14) more than available throughout the year,

(5) gniss and meadow habitats were selected less than available by adult females in the Burwash regioa;

(6) south slop utibtion increased during winter, aiggesting that snow conditions Muence spatial behaviour of elk in the region; and

(7) snow accumulation increased the proximity of anhaîs to water bodies in late winter, when fiozen water bodies were used as travel corridors; Ab,A W. 1982. Migration. Pg. 301-321, in Eik of North Amaica: ecoiogy and management (J. W. Thomas and DE. Toweill, eds). Stackpole Books, Harrisburg, PA

Alcock, J. 1 993. Animal behaviow: an evolutionary approach. Siriauer Associates, Inc., Suderiand. 625 pp.

Altmann, M. 1952. Social bebaviour of eik, Cervtrs dnszsneLsonii, in the Jackson Hole are-of Wyoming. Behaviour 4(2): 1 1 6- 143.

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Beyer, D.E. 1987. Population and habitat management in Michigan PhD. Thesis. Michigan State University, East Lansing. 160 pp.

Bosveld, K J. 19%. A review of doaimenteci occunences of native eik (Cems el

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Boyce, MS. 1989. The Jackson eik had: intensive wiidlife management in North Ameriui. Cambridge University Press, Cambridge. 306 pp.

Brazda, AR 1953. Etk migraiion patterns, and some of the fàctors affêcting movmients in the Gallatin River Drainage, Montaas. J. WdI. Manage. 17(1): 9-23. Bryant, L.D. and C. Maser. 1982. Class'ication and distribution. Pg. 1-59, in. EU( of North America: ecology and management. (J. W. Thomas and D.E.Toweül, eds) Stackpole Books, Harrisburg, PA

Burt, W.H. 1943. Temtoriality and home range concepts as appüed to mammals. J. Mammal- 24: 346-352-

Chambers, B., K Legasy, and C.V. BentIey. 19%. Forest plants of central Ontario. Lone Pine Publ., Edmonton 448 pp.

Chambers, B.A, B.J. Naylor, J- N~eppola,B. Merchant, and P. mg. 1997. Field guide to forest ecosystems of central Ontario. Ont. Min. Nat. Res.?SCSS FieId Guide FG-O1. 200pp.

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Clayton, J.S., W.A Ehrlich, D.B. Cam, J.H. Day, and I.B. Marshall. 1977. SoiIs of Canada. Cam Dept- of Ag., Soil Res. Inst., 1: 1-243.

Clutton-Brock, T.H.,F.E. Guianes, and S.D. Albon. 1982. Red deer: the behaviour and ecology of two sexes. Chicago University Press, Chicaso. 378 pp.

Collins, W.B., and P.J. Umess. 1983. Feeding behaviour and habitat selection of mule deer and elk on northem Utah surnmer range. J. Wrlài. Manage. 47(3): 646-663.

Craighead, Jr. J.J., G. Atweii, and B. W. O'Gara. 1972. Elk migrations in and near Yellowstone Nationai Park. Wddlife Monograph 29: 1-48.

Craighead, Jr. J.J., F.C.Craighead, Jr., RL.Re and B.W. O'Gara- 1973. Home ranges and activity patterns of nonmigratory elk of the Madison drainage herd as determineci by biotelemetry. Wddi. Monogr. 33 : 1-50.

Dunn, J.E., and P. S. Gipsoa 1977. Anatysis of radio telemetry data in studies of home range- Biornetrics 33: 85-101.

Edge, W.D.,and CL.Marcun 1985. Movements of elk in relaîion to loggiag distnirbmces. J. WiIdl. Manage. 49(4): 92-30.

Edge, W.D., CL.Marcum, and S.L. Olsoa 1985. Effkcts of log& activities on home-range fidelity of dk. J. Wddl. Manage. 49(3): 741-744. Edge, W.D., C.L.Mar- and S.L. Olson-Edge. 1987. Summer habitat seleaion by elk in western Montana: a mdtivariate approacb J. Wddl. Manage. 51(4): 844- 85 1.

Edge, W .D., CL.Marcum, and S.L.Olson-Edge. 1 988. Sumner forage and feedùig site selection by ek J. Widi. Manage. 52(4): 573-577.

Franklin, W.L., AS. Mossman, and U Dole. 1975. Social organization and home range of Roosevelt ek J. Mammai. SiS(1): 102- 11 8.

Friedmaq M. 1937. The use of ranks to avoid the asnimption of mrmaiity implicit in the anaiysis of variance. J. Am. Stat. Assoc. 32: 675-701.

Geia, V. 1982. Adaptive behaviowal strategies Pg. 219-277, in. Ek of North America: ecology and management. (J.W. Thomas and D.E.Toweili, eds.) Stackpole Books, Harrisburg, PA

Grover, KE., and MI. Thompson. 1986. Factors influencing spring feeding siîe selection by elk in the Ekhom Mountains, Montana J. Wddl. Manage. 50(3): 466-470.

Huggard, D.J. 1993. Effêct of snow depth on predation and scaveaging by gray wolves. J. Wildl. Manage. 57(2): 382-388.

Lrwin, L.L., and J.M Peek. 1983. Elk habitat use relative to forest succession in Idaho. J. Wildl. Manage. 4'7664672.

Jenks, J.k, and DM. Leslie, Jr. 1988. Effect of lichen and in vitro methodology on digestibiiity of winter deer diets in Maine. Can. Field-Nat. 102(2): 21tS220.

Jennrich, RI., and F.B. Turner. 1%9. Mea~u~ernentof non-cirdar home range. J. Theor. Biol. 22: 227-237.

Johnson, D.E. 1951. The biology of the elk ca& Cervrrs dl~~tsnehoni. J. WU. Manage. 1x4): 3-10.

Johnson, D.H. 1980. The coinparison of usage and availabïiity measurements for evaluations of resource preference. Ecology 61 : 65-7 1.

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Accuracy of a radio-telemetry system used to monitor collareci elk, Centur eiaphus, in the French River and Burwash Regions of central Ontario. Sampling errors and variance were measured by placing 12 VHF test collan (Mode1 LMRT 4, Lotek Inc.) throughout the Burwash study area in a variety of habitats and terrains frequented by the elk. Five replicate bearing estimates were obtained for each collar. The radio telemeuy system was

found to have a small positive bias (1 .O 1O). The standard deviation in bearing error for aU test transmitters and receivers was +/- 5.01. The average area of the 95% confidence ellipse was 137 700 rn2,with a mean axis length (major and minor) of 204.86 m (SD = +/- 122.82). Raw data and calculations of the sarnpling error (standard deviation) and probability error eiiipses are presented in subsequent appendices. Major causes of error in the system were transmission lines and granite ridgcs, and the time lag between recording bearings. The average area of a 95% confidence eiiipse was larger than the average area of habitat polygons based on the Forest Resource Inventory @RI) classification sy stem. However, it was smaller than habitat pol y gons delineated by the ecosite classification system. Al1 conclusions on habitat utilization by French River and Burwash ek using this type of telemetry system should be viewed with caution.

Descriptive statistics for bearing error data c 10' collected to detennine the accuracy of bearings obtained using a hand held Yagi antenna. The tme bearing from each receiving location to each test collar was calculateci using trigonometrïc formulas desmbed in White and Garrott (1990). Error was calculated by subtracting the estimated bearing from the mebearing. Bearing error data are in degrees.

Bearing Enor Receiver No. of bearings Minimum Maximum Mean SD Variance Location recorded Appendix A continued

Total 292 Mean Test results determining the accuracy of bcarings obcained using a hand hcld Yagi latcnna Five replicate karings were recordcd at eacb nceiving location for 12 diffkrent test collar (transmiucr) locations. nit me bearing from each receiving location to each test coiiar was calculated using aigonomctric formulas desdbed by White and Garroa (1990). Error was caiculated by subaacting the estimated bcaring hmthe mebearing. Only enor data bctween -IV and +IV were uscd for aaalysis- [Tx=UIM cast-west valuc (metered units) of a transmitter location, Ty = URvI n~southvalut for a pansmiacr location, Rx= UTM tast-west valuc for a rcceiving location, Ry = UTM aorth-swth value for a rccciving location]

1Oi3 I/% Full doud 9:47 a Blowing aow

1 1/1/96 Paniai doud 430 -5 Wind Parrul cId -5 Wüod

Partial Cloud O

FUclod 4

fullcld,4 IV?. Para;ilcIwd O

1 lBC96 Full dood 4 Appcndu B ConOnued 17: 16

1739

18:IO

18~43

1 1/3/96 Full Cloud 1 O:46 -4

1 Ii3196 Full clouri 1 O:B 4

Il35

1 118/% Full doud 14~45 6

15:14

1945

16a 1118196 Full clod 14:45 6

11/8/96 Fuil cloud 14:45 6 Appcndix B Continued

1 1/ 18/96 Full Cloud 13:18 -3 1 1/18/96 Full cloud 13:18 -3

Calculation of the standard deviation of pooled sampling enon which reflects thc rznpling error inherent in hearing estimation. The mean sampling error was calculated using the formula . Mcrn e = cii / nr, =-here n is the number of test collar locations, and eq is the enor for each replicate baring j at each location i Bearing standard deviation (SD = [ (eu - Man e)'l (nr - 1)11") was calculated in two ways: "all data" I error data for all bearings (0and locatioor 0;and "<= 100; only error data - 100<= eq x +100 was used in the calculation . SD ( <= 10" ) was used in tbc calculation of 95%confidtnce cllipscs for each test collar r White, 1985). Bearing standard deviations were reported in relevant chapters as kypertained to home range size and habitat utilization- Appendix C Continucd Appendix C Continued

Transmiacr No.

I Cj 3 i 5 6 7 a 9 IO II IZ DIr I(Y31/96 1111B6 11/1/96 1 IR196 11M 11M 11/81% 1 Il11/17/96 11/17/96 11/17/96 Cdlar 606 606 513 606 513 544 606 513 544 . 606 513 544 Tdtttrla. Eantx) 511250 512250 51m 512900 513200 511900 511450 511350 S1M50 512100 514190 511080 Nd(y) 5125750 5124900 5125250 5126200 5126600 5124400 5127050 51272ûû 5127200 5121730 51î5160 5122330

959 îonfidtnœ tllipsc utî (Ha) 6.08667 332556 4.4U29 5.18358 6.96143 652734 155093 182S17 3ZCü513 16.85 36.9246 13-0326 Appendix D continued intermediate measurcs in the calculation of the Iebgths of the major (M) and mina (nt) axes Tm39 -1.4049 -02572 216482 351435 1.74534 1.72074 -1.4415 -1.58742 -1.1153 -10-48 0.17188 -1.016 Td -0.5157 4.1265 0.63% 0.75515 037955 03546 -03233 455193 4.4465 4-9091 OB8531 4.4189 S.' 4034.74 329204 339555 3280.47 406058 1034.94 15024.7 18839.8 399202 1- 7528.88 W.64 s,' 2593% 527588 1673.85 231293 337022 116tS3 4520.94 4993.16 727137 4828-73 511382 536226 Sur O O O O O O O O O O O O Formikr Ta 39 = 2Sl SI - SI') s,' = st2 + Si2 Tane s,'=s~'-SizTdl s,=o

Majar adminor axes M 155591 140543 142735 140295 156.088 78.8013 30.246 336213 489.- 315.632 21- 231.663 m 124.731 17.7919 100215 117.803 142202 264.105 164.699 173.W nJB.874 170213 553922 17937 Fonmlar loclusiorn probability. P(r).stt to 0.95. Mth 3 = 6. wbar P(r) = 1 - e4= hi = 6" m=rS, Mum M luigrh = 224.97978 m standard kviatioo = 1 16.13957 a = 12 Mean m lenM= 184.75094m srandarddeviaioa=131.03123 n= 12 Majœ and minor poolal. mern km= 204.86536 m mmdwd dcnr9;oa = 12281898 m n = 24 Ralength = 102.43268 m sMdud deviaion = 12281898 m Appendix E

Forest Ecosystem Classification Ecosites for centrai Ontario (Minisuy of Natural Resources). Vegetation demiption includes es,shnibs; herbs, fems and allies, grasses, mosses and Iiverworts. and lichens. Site features pertaining to the ground iayer arc also provided. The number of herbs is a measure of ecosite diversity and ranged from, few (<8 species), moderaîe (8-15 species), and herb rich (>15 species). In the vegetation description, species are listed in order of frequency of occurrence (>50%), with those in brackets occurring from 30-508 of the tirne. (Chambers et al., 1997)

Ecosite 1 Sugar Maple-Basswood stands on fresh to moist soils. Moderate number of herbs

Vegetation Description

Trees Main Canopy sugar maple, (basswood) Sub Canopy sugar maple Regeneration Sugar maple (American elm, white ash, balsam fu, ironwood, basswood)

Shmbs alternate-leaved dogwd, swarnp black currant, fly honeysuckle

Herbs, Fems and Allies, Grasses spinulose wood fern, (helleborine, red trillium, wild sarsaparilla, foamflower, wild lily of the valley, blue cohosh)

Mosses and Liverworts Bruchythecium sdebrosum, (Calliciudiwn haldanianum, Brachythecium reflexm)

Site Features Occurs primanly on morainal Iandforrns, aiso on glaciofluvial and lacustrine landforms. Forest floor dominated by broadleaf litter and &ph of organic matter less than 20 cm. Humus forms range frmmulls to modem to fibnmors. Coarse fragments abundant to fcw.

Ecosite 2 Sugar Mapie-Amencan Bcech-Basswood stands on dry to moderatcly fkhsoils. Moderate number of herbs.

Vegetation Description Trtes Main Canopy sugar maple, (red oak, Amcrican kh) Appendix E continued

Sub Canopy sugar maple (American beech) Regeneration sugar maple (ironwd, red oak, American beech, white ash)

Shrubs stripped maple

Herbs, Fems and Allies, Grasses spinulose wood fern, wild My of the valley, starflower, red trillium (trout lily, false Solomon's seai, wild sarsaparilla hairy Solomon's seal, rose twisted-staik, nce grass, marginal shield fern)

Mosses and Liverworts Brachythecium reflexwn, Brachythecium salebrosum, (Dicranum scoporium, Callicladium haldanianum,

Lichens cladi~coniocraea

Site Featurts Occurs rnainly on morainal landforms, occasional on glaciof luviai landforms. Forest fioar dominateci by broadleaf litter, and depth of organic matter less than 10 cm. Humus fomis range from fibrimors to rnaàers. Coarse fragments moderate to few.

Ecosite 3 Sugar Maple-Red Oak-Basswood stands on dry to moderately fresh soils. Moderaie nurnber of herbs.

Vegetation Description

Trees Main Canopy (sugar maple, red oak) Sub Canopy sugar maplc (ironwood) Regeneration sugar maple, ironwood, white ash, red oak, (balsam fir, basswood, Amcrican beech)

Shb (Maple-leaved vibumum, Stripped mapk. Fly honeysuckle. Leathewood)

Herbs, Fems and Allies, Grassts luge-lcaved aster, wild ssrsppprilla, wild lily of the vdley, hairy Solomon's Ka, falsc Solornon's seai, (Rd trillium, Appendix E continued

rice grass, round-lobed hepatica, helleborine, spinulose wood fern, white trillium, downey yellow violet)

Mosses and Liverworts Brachythecium reflexum, Dicranum flagellare, Brachythecium salebrosum, (Mnium cuspidanun, Bracytkcium velutinum, Callicladium haidanianwn)

Lichens CladUza coniocraea, Cladonia chlorophPe4)

Site Featurts

Typically occurs on morainai landfonns. Forest floor dominated by broadleaf litter, and depth of organic matter les than 20 cm. Humus forms range from fibrimors to modes. Coarse fragments abundant to few.

Ecosite 4 Red Oak-Sugar Maple stands on dry to mderately fresh soils. Moderate number of herbs.

Vegetation Description

Trees Main Canopy red oak (sugar maple) Sub Canopy sugar maple (ironwood, red oak) Regeneration sugar maple, ironwood, balsam fir, red oak, white ash, (black cherry)

Shbs (strippcd maple, fly honeysuckle, serviceberry spp., wintergreen. partridgeberry)

Herbs, Fems and Allies, Grasses wild iily of the valley, wild srnaparilla, large-leaved aster, hairy Solomon's seai, rice grass (false Solomon's seal, spinulose wdfem, staxflower, bracken fem)

Mosses and Liverworts (Brachythecium refzexwn,Dicronum flagelhre, Braehytheciwn salebrosum, Polytrichumjuniperinum) Appendix E continued

Lichens Cladina coniocraea, Chdonia chioropliaeu

Site Features

Occurs pnmarily on morainal landforms, occasional on lacustrine and glacioflivial landforms. Forest floor dominated by broadleaf litter, and depth of organic matter less than 20 cm. Humus forms range from moâers to fibrimors. Coarse fragments abundant to few.

Ecosite 5 bwland Hardwoods-Black Ash-Popiar-Mixed Hardwood stands on wet to dry sites, in flats or pockets, telluric slopes and fluvial sites.

Vegetation Description Trees Main Canopy (black ash, trembling aspen) Sub Canopy (black ash, sugar maple) Regeneration black ash, balsarn fÎr, (sugar maple, white elm)

Shrubs dwarf raspbemy, beaked hazel, mountain maple, choke cherry, fly honeysuckle,(wild red raspberry, red currant, al temate-leaved dogwood)

Herbs, Fems and Allies, Grasses spinulose wood fern, wild sanaparilla, (large-leaved aster, fragrant bedstraw, lady fern, starfiowcr, wild lily of the valley, blue bead My, hairy Solomon's seal, rose twisted- stalk, meadow-rue, nodding tnlliurn)

Mosses and Liverworts (Brachythecium refexum, Callicladim huldaniruupn, Brachythecium salebrosron)

Site Featurits

Occurs on glaciofluvial, fluvial and 1acustrine landforms, occasional on morainal landforms. Forest floor dorninated by bf0adlea.f litter, and depth of organic matter less than 20 cm. Humus forms typically muils and modes. Coarse fragments typically few.

Ecosite 6 Eastern White Cedar-Lowland Hardwoods. Eastern White Ctdar-Black Ash-Ycllow Birch-Poplar stands on wet to dry soils, in flats or pockets, telluric slopes and fluvial sites. Appendix E continued

Vegetation Description

Trees Main Canopy eastern white cedar,(hardwoods 10-3096 frequency) Sub Canopy eastern white cedar, black ash, balsam fir Regeneration balsam fir, (black ash, eastern white cedar, sugar maple, red maple)

Shmbs dwarf raspberry, mountain maple, fly honeysuckle, swarnp black currant, (beakcd hazel, twinflower)

Herbs, Fems and AUies, Grasses fragrant bedstraw, spinulose wood fem, wild sarsaparilla, starflower, blue bead My, bunchberry, wild lily of the valley, large-leaved aster, oak fern, goldthread, lady fem, (nakod mitrewort, rose twistad-stalk, wood sorrel, kidney- leavcd violet)

Mosses and Liverworts (Plagiotheci~unlaetum, Pleuro~iwnschreberi, CallicLadium haldonianu)

Lichens (Chdonia coniocraea)

Site Feanires

Occurs on lacustrine, organic, morainal and glaciofluvial landfoms. Forest floor a mixture of broadleaf and conifer litter, and depth of organic matter occasionally 40 cm. Humus fomranging from mulls to moders to mors, to peatymors. Coarse fragments typically few.

Ecosite 7 Sugar Maple-Yellow Bhrh stands in dq to moderately fresh soils. Moderate number of herbs.

Vegetation Description Trces Main Canapy sugar maple, yeiiow birch (white spruce, eastem white &) SubCPwpy sugarmapk Regeneration sugar maple, baisam fi Shnibs mou~ltainmaple, fly honeysucklc, choke cherry, ground Appendix E continucd

hemlock, (beaked hazel)

Herbs, Fems and Allies, Grasses spinulose wood fem. rose twisted-aalk, starflower, false Solomon's seal, blue bead lily, shining clubmoss, (wild safsaparilla, hairy Solomon's seal, bunchkrry, large-leaved aster, ground pine, wild lily of the dley, northern beech fern)

Mosses and Liverworts Callicladium haldonianu, (Brach?tizecium salebrosum, Mniwn marginurum, Dicranun scopariwn)

Lichens (Cladonia coniocraea)

Site Featutes

Occurs on glaciofluvial and morauial landfom. Forest floor a mixture of broadleaf, and depth of organic matter typicaliy les than 10 cm. Humus fonns ranging from fibrimors to moders to mulls. Coarre fragments typically abundant.

Ecosite 9 Sugar Maple-Yellsw Birch-Eastern Hemlock on dry to moderateiy fresh soiis- Moderate number of herbs.

Vegetation Description

Trees Main Canopy eastern hedock, sugr maple, (yellow birch) Sub Canopy sugar maple, (eastern hemlock) Regeneration sugar maple, balsam fir, red maple, eastern hemlock, yellow birch, (Amcrican beech)

Shrubs stripeà maple, 8y honeysuckle, (kaked ha&, hobblebush, mountain maple)

Herbs, Ferns and Allies, Grasses spinulose wood fern, wild îiiy of the valley, rose twistai- stalk, starflower, wild sarsaparilla, -und pine, (haïry Appendix E continutd

Solomon's seal, shining clubmoss, Wian cucumber root, rtd tnllium, blue bead lily)

Mosses and Liverworts Brachythecium reflexum, (Callic~holdanianu, Plagiothecium hem,Dicranum moritanwn)

Lichens Clodonia coniocraea

Site Feanires

Occua on glaciofluvial and morainal landfonns. Forest floor a mixture of broadleaf and wnifer litter, and depth of organic matter typicaliy less than 20 cm. Humus foms ranging from fibrimors to humimors. Coarse fragments few to abundant.

Ecosite 10 Eastern Hemlock-Yellow Birch on dry to moderaiely fresh soils. Moderate nurnber of herbs.

Vegetation Description Trees Main Canopy eastern hemiock, (ycïlow birch) Sub Canopy (eastern hemlock, eastern white cedar, balsarn Fu) Regeneration balsam fir, red maple. eastern hemlock, yellow birch, (AmeRcan beech, eastem white cedar)

Shmbs striped maple, fly honeysuckle, inornitain maple, beaked hazel, (hobblebush)

Herbs, Ferns and Allies, Grasses spinulode wood fem, wild My of tbe valley, starflower. wild sarsaparilla, blue bead lily, rose twisttd-stalk, wcmd sorel, goldthrcad, shining clubmoss, bunchbeq, (ground pine)

Mosses and Livexworts Bauonirr tnïobata, Pkùgiothecium ktum, Polytrichtan commrrnc,(Dicrmuun figeilare, Dicranum montanum, Br4chythccim teflenan, Brotkreh recunans, Ptilidium pulchctrurwn) Appendix E continued

Lichens Cladonia coniocraea

Site Featwes

Occm on glaciofluvial and morainal landfom. Forest floor a mixture of conifer and hardwood litter, and depth of organic matter typically less than 20 cm. Humus fomis ranging fiom fibrimoa to humimors. Coarse fragments few to abundant

Ecosite 1 I Poplar-White Birch stands on dry to moderately fresh soils. Moderate number of herbs.

Vegetation Description Trees Main Canopy white birch, (largetooth aspen) Sub Canopy white birch, red rnaple, (sugar maple) Regeneration balsam fw, red maple, white birch, sugar maple (trembling aspen)

Shmbs beaked hazel, low sweet blueberry, fly honeysuckle, northem bush honeysuckle, mountain maple, (velvet leaf blueberxy)

Herbs, Fems and Allies, Grasses blue bead My, wild sarsaparilla, large-leaved aster, wild lily of the valley, starflower, ground pk,bracken fern, rose twist&-staik, bunchberry, (goldthrcad, shining clubmoss)

Mosses and Liverworts Pleuroziwn schreberi, (Callidadim haldanionum)

Lichens Cladonià coniocraea

Site Featurcs

Occurs on glriofluvial and morPinal landforms. Forest floor dominated by ôroadlepf Litter, and depth of organic matter Iess than 10 cm. Fibrimors m the dominant humus forms. Coarse fragments typically abundant. Appendix E continued

Ecosite 12 Poplar-White Birch-White Spnice-Balsarn fir stands on dry to rnoderately frcsh soils. Moderate number of herbs.

Vegetation Description Trees Main Canopy trembling aspen, white birch, (white spruce, balsam fir) Sub Canopy white spruce, (baisam fir, white birch) Regeneraiion balsam fir, (trembling aspen, red maple, white spmce)

Shntbs northern bush honeysuckle, beaked hazei, mountain maple, low sweet blueberry, fi y honeysuckle, (twinflower, northern wild raisin)

Herbs, Fems and Allies, Grasses starflower, wild lily of the valley, large-leaved aster, wild sarsaparilla, blue bead lily, bunchberry, ground pine, bracken fern, rose twisted-stalk, (spinulose wood fem, goldthread, wolf s claw clubmoss)

Mosses and Liverworts Pleurozium schreberï, (Plagiotheciron laenun, Ptîlidium pulcherrimum, Calliciadiwn haldanianum, Brachythecim salebroswn, Dicranum scopariwn, Sanionia unincltus)

Lichens Cladonia coniocraea, (Cladonia chlorophaea)

Site Features

Occurs on gia~iofluvialand morainal landform. Forest floor dominated by a mixture of bmadleaf and conifer litter, and de@ of organic matter typically less than 10 cm. Humus forms predominantly fibrimors. Coarse fragments abundant ta few.

Ecosite 13 Poplar-Jack Pine-White Spnrce-Black Spntce stands on dry to modcraîcly fresh soils. Moderate number of herbs.

Vegetation Description Trees Main Canopy trcmbling aspen, jack pine Sub Canopy (balsam fu, white sprue, white birch) Regenedon balsam fir, red maplt, white birçh, white Appendix E continued

spruce (black spruce)

Shmbs northem bush honeysuckle, low sweet blueberry, velvetleaf bluekrry, beaked hazel, twinflower, mountain maple (fly honeysuckle, showy mountain ash)

Herbs, Fems and Allies, Grasses wild lily of the valley, wild sarsaparïlla, bunchberry, blue bead lily, large-leaved aster, bracken fern, starflower, ground pine, goldthread, (spinulose wood fem, rose twisted-stalk)

Mosses and Liverworts Pleuroziwn schreberï, Dicranum polysetum, Dicranum fuscescens

Lichens (Cladonia coniocraeu) Site Feanue~

Occurs on glaciofluvial and morainal landform. Forest floor a mixture of bmadleaf and conifer litter, and depth of organic matter iess than 20 cm. Humus fomis predominantly fibrimors. Coarse fragments abundant to few.

Ecosite 14 White Pine-Red Pine-White Spruce-White Birch stands on dry to moderately fresh soils. Moderate number of herbs.

Vegetatioa Description Trees Main Canopy white pine, (white spruce, red pine, white birch) Sub Canopy (white spruce) Regeneration balsam fir. red mapie, (trembling aspen, white pine, white spruce, white birch)

Shmbs beaked hazcl, mouritain maple, northem bush honeysuckie, fly honeysuckle, low swcct blueberry, twinflower, velvetleaf bluebcny Appendix E continued

Herbs, Ferns and Allies, Grasses blue bead lily, wild sarsaparilla, wild Iily of the valley. bunchkrry, starflower, bracken fem. large-leaved aster, ground pine, goldthread, (spinulose wood fem, rice grasS. rose twisted-staik)

Mosses and Liverworts Pleurozium schreberi. (Ptilidium pulcherrimum)

Site Feahucs

Occun on glaciofluvial and morainal landforms. Forest flwr a mixture of conifer and hardwood litter, and depth of organic maner less than 20 cm. Humus fomrange from fibrimors to moders. Coarse fragments abundant to few.

Ecosite 15 Eastem White Cedar-White Birch-White spruce- White stands on dry to moderately fresh soils. Moderate number of herbs.

Vegetation Description Trees Main Canopy white birch, (white spruce, white pine, eastern white cedar) Sub Canopy balsam fir (easteni white cedar, white birch) Regeneration balsam fir, red mapie. eastem white cedar, white birch

Shmbs fly honeysuckle, mountain maple, Iow sweet bluebcrry, beaked hazel, showy mountain ash, northern bush honeysuckle, twinflower

Herbs, Ferns and Allies, Grasses wild lily of the valley. blue kad My. wild sarsapda, starflower, goldthread, spinulose wood fem. bunchberry, rose twisted-stalk, ground pine, (large-leaved aster, bracken fem)

Mosses and Livenivarts Pteuroziun schrtberi ,(PtiIidium pulckrrimrmr) Appendix E continued

Lichens Claddnia con iocraea , Clczdonià chlorophuea

Site Features

Occurs on glaciofluvial and morainal landforrns. Forest floor a mixture of conifer and broadleaf litter, and depth of organic matter lesthan 20 cm. Humus fomrange from fibrimors to moders. Coarse fragments abundant to few.

Ecosite 16 White Pine-Largetwth Aspen-Red Oak stands on dry to moderateiy fresh soils. Moderate number of herbs-

Vegetation Description Trees Main Canopy white pine, (red oak, largetooth aspen) Sub Canopy (red oak, white pine) Regeneration red maple, red oak, balsam fir, white pine (sugar maple)

Shrubs wintergreen, late low blueberry, beaked hazel, fly honeysuckle, (bush honeysuckle, velvet leaf blueberry, arnelanchier spp.)

Herbs, Fems and Allies, Grasses wild Iily of the valley, bracken fern, wild sarsaparilla, large- leaved aster, nce grass, starflower, bunchberry)

Mosses and Livtrworts (Dicrrmmflagelklre, Ptilidium pukhemmum, PleumziMt schreben]

Lichens Cladonia coniocraea, (Cladonia chlorophaea)

Site Feanires

Occurs on glaciofluvial and morainal landfoc~lls. Forest floor a mixture of conifcr and broadlcaf litter, and depth of organic matter lcss than 10 cm. Humus forms range hmfibrimors to moders. Coarse fragments abundant to few. Appendix E continued

Ecosite f 7 White Pine-Red Pine stands on dry to moderaiely €resh soils. Low number of herbs.

Vegetation Description Tnees Main Canopy white pine, (red pine) SubCahopy - Regeneration red maple, balsam fu, white pine (white spruce, white birch)

Shmbs late low blueberxy, beaked hazel, wintergreen, twinflower, velvet leaf blueberry, northern bush honeysuckle, (fly honeysuckle)

Herbs, Fems and Ailies, Grasses wild Lily of the valley, bracken fem, large-leaved aster, rice grass, starflower, bunchberry, (blue bead Iily)

Mosses and Liverworts Pleurozira schreben', (Dicronum polysem)

Site Features

Occurs on glaciofluvial and morainal landfom. Forest floor a mixture of conifer and broadleaf litter, with mosses, and depth of organic matter less than 10 cm. Humus forms range from fibnmors to rnoders. Coarse fragments abundant to few.

Ecosite 18 Red Pine stands on rnoderately fresh soils, bwnumber of herbs.

Vegetation Description Trees Main Canopy reà pine, white pine Sub Canopy - Regeneration balsam Tu, white pim, red maple, (red oak, white birch, white spruce)

Shbs beaktd hazel, northem bush honeysuckle, Iow swect blueberry, wintergreen, twinflower, (velvetleaf blueberry, fly honeysuckle. Amekhier spp.) Appendix E continucd

Herbs, Ferns and Allies, Grasses wild lily of the valley, bracken fem, wild sarsaparilla, large- leavcd aster, nce grass, (stamower, bunchberry, blue bead Iily)

Mosses and Liverworts Pleuroziwn schreberi, (Dicranum polysetwn)

Site Features

Occurs on morainal, glaciofiuvial and lacustrine Iandfonns. Forest floor a mixture of conifer litter, with some broadleaf litter and mosses, and depth of organic matter less than 10 cm. Humus forms range from moders to fibnmors. Coarse fragments abundant to few.

Ecosite 19 White pine-Red Pine-Jack Pine stands on dry to moderately fresh soils. Low number of hehs.

Vegetation Description Trees Main Canopy (white pine. jack pine) Sub Canopy (white pine) Regeneration red maple, balsam fu. white pine (white birch)

Shmbs low sweet blueberry, win tergreen, aorthern bush honeysuckle, (velvetleaf bluebeny, twinflower)

Herbs, Fems and Allies, Grasses wild lily of the valley, bracken fem, wiid sarsaparilla, (large-leaved aster, bunchberry, starflower, cow-wheat, hairgrass)

Mosses and Liverworts Pleurozïum schreberi, Dicranum pdysetwn, Dicranwn figellare, (Ptilidium pulcherrimum, Dicranutn onîarkme)

Lichens Cladonia coniocraeu, Cladina ranggenna, (Clodonu chlorophaea) Appendix E continucd

Site Features

Occurs on morainal and glaciofluvial landforms. Forest Boor a mixture of conifer and broadleaf litter, with mosses, and de@ of organic matter less than 10 cm. Humus forms predominantly fibrirnon. with some moders. Coarse fragments few to abundant.

Ecosite 20 Eastern white cedar-Black spruce-Tamarack on wet organic and fresh to moist minerai soils. Moderate number of herbs.

Vegetaîion Description Trees Main Canopy black spmce, (tamarack, eastern white cf=dar) Sub Canopy (black spruce, eastern white cedar) Regeneration balsam fir, eastem white cedar, red maple, black spruce, (yellow birch)

Shrubs creeping snowberry, northern wild raisin, low sweet bluebeny, showy moun tain ash, labrador tea, (speckled alder, mountain-holly, velvetleaf blueberry, dwarf rasbeny, twinflower)

Herbs, Fems and Allies, Grasses goldthread, starfiower, bunchbeny, blue bead lily, wild lily of the valley, (spinulose wood fern, three-fmited sedge, cinnamon fem)

Mosses and Liverworts Sphagnum girgenstohnii, Sphagnum mugellanicum, Sphagnm capilliofoZium, Pleurozim schreberi, (Dicranumfigellare, Plagiotheciwn laenan, Bazzunia trlobata) Lichens Chdonia coniocraeu, (Cladonu chbrophaea)

Site Features

Occurs on organic, giaciofluvial and lacustrine lamKorms. Forest fioor dominateci by mosscs, with some conifer litter, and dcpth of organic matter typically greater than 40 cm. Humus forms range hmpeatymors, to fibrimors and modtrs on upland sites. Coarse fragments typically few. Appendix E continued

Ecosite 21 Eastern white cedar-other conifer on dry to moist mineral soils. Moderate number of herbs.

Vegetaîion Description Trees Main Canopy (black spruce, white birch, eastem white cedar, balsam fir) Sub Canopy balsam fir, eastem white cedar, (white birch) Regeneration balsarn frr, eastern white cedar, red maple, (white birch, black spruce)

Shmbs mountain maple, fly honeysuckle, twinflower, showy mountain ash, (low sweet bluebeny, velvet-leaf blueberry, beaked hazel)

Herbs, Ferns and Allies, Grasses starflower, bunchberry, goidthmaci, blue bead My, wild lily of the valley, wild sarsaparilla, spinulose wood fern, (ground pine, oak fem, rose twisted-stalk, bracken fern)

Mosses and Liverworts Pleurozium schreberi, (Bazzmia tnIrtZobata,Dicranum flagellare, Plagiothecim laenun, Dicranwn fuscescens, Holyocomim splenderzs, Dicrunum polyserwn)

Lichens (Ciadonia coniocraea, CWOMchlorophaea)

Site Features

Occurs on morainal, glaciofluvial and lacustrine landforms. Forest floor a mixture of conifer and broadleaf liner, and depth of organic matter typically Iess than 20 cm. Humus forms range from fibrimors and humimors on upland sites- Coarse fragments few to abundant.

Ecosite 22 Black spnict-Pine on dry to moderately frcsh soiis. Low number of herbs.

Vegetaîion Description Trets Main Canopy black spnice, (white pine) Sub Canopy black spnice, (balsam Tir) Regeneration black spmce, baisam fir, white pine, Appendix E continued

(white birch, eastern white cedar)

Shmbs low sweet bluebeny, creeping snoa-beny, velvet-leaf blueberry, twinflower

Herbs, Ferns and Allies, Grasses bunchberry, wild lily of the valley, (starfiower. goldthread, bracken fem)

Mosses and Liverworts Pùuroziwn schreben, Dicranum pofysenon (Dicr~um scopariurn, PtiIidiwn ciiiare)

Lichens

Occurs on morainal and glaciofluvial landforms. Forest floor a mixture of mosses and conifer Iitter, and depth of organic matter typically less than 20 cm. Humus forms predominantly fibnmors. Co- fragments few to abundant.

Ecosite 23 Jack Pine-Black Spruce on dry to moderately hrbsoüs. Low number of herbs.

Vegetation Description Trees Main Canopy jack pine Sub Canopy black spmce Regeneration black spruce, baisam fu, (white birch, red maple)

Shnibs low sweet blueberry, velvet-leaf biaebemy, creeping snowberry, twinflower, (winterv, northern bush honeysuckl)

Herbs, Ferns and mes, Grasses wild lily of the vailey, bunchberry, @lue bead lily, starflower, bracken fern, moccasin fiower) Appendix E continued

Mosses and Liverworts Pleurozium schreberi, Dicranwn polysenrm (Sphagnwn girgensohii, Ptilidium pulcherrimun)

Lichens Cladim rangvenira (Ciadonia caniocruea, Cladonia chiorophoeu)

Site Features

Occurs on moraiaal and glaciofluvial landfoms. Forest floor a mixture of mosses and conifer litter, with some broadleaf litter, and depth of organic matter less than 20 cm. Humus forms typically fibrimors. Coarse fragments few to abundant-

Ecosite 24 Black Spruce-Tamarack stands on wet organic and moist to dry minerai soils. Low number of herbs.

Vegetation Description Trees Main Canopy black spruce (tamarack) Sub Canopy black spruce Regeneration black spruce, (balsam fu)

Shnibs low sweet blueberry, creeping snowberry, labrador te* sheep laurel, vel vet-leaf blueberry, northern wild raisin, (mountain holly ,speckled alder)

Herbs, Ferns and Mes, Grasses bunchberry, goldthfead, blue bead Iily, (three-leaved srnilacina)

Mosses and Liverworts Pleurozhm schreberi, Sphagnum girgensohnii, Sphagnwn capillï$olium, Dicranum poiysetwn, Sphogntmi magelhicwn, Ptilidium ciüare

Lichens Clcrdinrr rangiferina (Clarloniia ccmiocraea, Ciudoniia chlarophoco) Appendix E continued

Site Feanires Occurs on organic, glaciofluvial, morainal ,and lacustrine landfoxms. Forest flmr dominated by mosses, with sorne con& and broadleaf litter, and &pth of organic matter typically greatcr than 40 cm, or less in peaty phase and upland sites. Humus foms range from peatyfonns to fibrimors. Co- fragments typicaily few. Multiple Chi-square analysis and Bonferroni confiidence intervals for habitats utilized in the Burwash Study area. Tbe habitat available to coIlared ek was delineated by placing a 5 km baer around the 1W/o minimum convex polygon containing location data coUected for ail coiiared animals between 1995 and the winter of 1997. Tbe study area encompasseci a total area of 324.7 km2. Refm to appendix E for descriptions of each habitat type. Habitat type descriptions wae based on MNR codes for non-forested habitats and ecosite types for forested habitats (m = habitat type selected significantly greater than in proportion to availability, based on Bonfèrroni Confidence Intervals; 1 = habitat type sdected significantly lesthan in proportion to availability; e = hab'rtat type selected in proportion to availab'i. Habita? type "

Animal: Y1 Habitat Type Observed Expected Obsewed Available Bonferroni lntervals Frequency Frequency Proportion Proportion pz P+z

Total 61

Animal: Y2 Habitat Type Observed ExQeded Observed Availabie Bonferroni lntervals Frequency Frequency Proportion Proportion pz PZ

Total 208 Animal: Y3 Habitat Type Ob6erved Expected Observed Availabk Bonferroni lntervak Frequency Frequency Proportion Proportion pz Pz

Total 225

Animal: Y6 Habitat Type Observed Expected Observed Availabk Bonferroni Intervals Frequency Frequency Promrtion ProgoRiori pz Pz

Total 38 Multiple Cbi-square analysis and Bonferroni confidence intervals for habitats utilued in the French River study ana. Tbe habitat available to cdlared eUc was delineated by plafuig a 10 lan buf5er arwnd the 1W?%minimum wnvex poiygon containing location data coiiected for al1 coflared animals between 1995 and the winter of 1997. The study area availabie to the eik encompassed a total of 1 133 -7km2. Refêr to appendix E for descriptions of each habitat type. Habitat type descriptions were based on MNR codes for non-forested habitats and ecosite types for forested habitats (m = habitat type selected signülcantly greater than in proportion to availabdity, based on Bonferroni Confidence Intervals; 1 = habitat type selected sigdicantly less than in proportion to availabiîis., e = habitat type selected in proportion to availability. Habitat type "

Animal: 61 Habitat Type Ohserved Expecîed Obsewed Available Boflfemni lntervals Frequency Frequency Proportion Proportion pz P+z

Total 169

Animal: B3 Habitat Type Observed Expeded Observed Available Bonferroni lntervals Frequency Frequency Proportion Proporüm pz PZ

Total 100 Animal: 85 Habitat Type Observed Expected Observed Avaiiaôie Bonferroni Intervals Frequency Frequency Proportion PropoRiori pz P+z

Total 49

Animal: 88 Habitat Type Observed Expeded Observed Availabk Bonferroni Intervals Frequency Frequency PropoRion Proporüon pz P+z

Total 38 Appendix H

Wilcoxon's Signed Ranks Tests to analyze for ciifferences betwœn summcr and wintcr in the relative frequency of locations for each animal found on south-facing aspects, as weli as differcnces in south vs. north vpct use during summr and winter. D = difference ktwcen relative frequency values when comparing summer vs. winter, and south vs. north.

South Slopes

Animal Sumrner Winter O Rank Bi 0.05 0.29 -0.24 10 83 0.1 9 0.33 -0.14 8.0 B4 0.1 O 014 -0.W 5.0 85 OZ6 OP5 0.01 3.0 B8 0.1 7 0.25 -0.08 6.0 09 0.00 0.00 0.00 1.O Y1 0.18 0.18 0.00 2.0 Y2 029 026 0.04 4.0 Y3 0.1 5 O24 4.09 7.0 Y6 0.1 4 0.30 -0.16 9.0 Absolute sum: Negative ranks 45 Positive rmks IO Significant as one-tailed test

Summer South vs North Animal South North O Rank BI 0.05 0.05 0.00 1.5 83 0.1 9 0.07 0.12 8.0 84 0.1 O 0.00 0.10 7.0 B5 026 0.03 0.23 10 88 0.1 7 0.39 -0.22 9.0 B9 0.00 0.07 -0.07 6.0 Y1 0.18 0.1 8 0.00 1.5 Y2 0.29 024 0.05 4.0 Y3 0.1 5 0.09 0.05 5.0 Y6 0.1 4 0.1 8 -0.04 3.0 Absolute sum: Negative ranks 36.5 Positive ranks 15.0 Not significant: T=lS>t=9,p>O.O5 Appendix H continued

Wnter South vs North

Animal South North Rank BI 0.29 0.03 9.0 03 0.33 0.00 10 64 0.1 4 0.00 4.0 B5 025 0.06 5.0 68 0.25 0.05 8.0 B9 0.00 0.00 1.O Y1 0.18 021 20 Y2 026 0.21 3.0 Y3 0.24 0.04 6.0 Y6 0.30 0.10 7.0 Absolute sum: Negative ranks 2.0 Positive fan ks 53 Significant as Two-tailed test T=2 c t = 9, p4.005