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Southern Canadian Rockies Lynx Project

Lynx Ecology and Conservation Requirements in theSouthern Canadian Rocky Mountains

Annual Summary ReportYear 4

1999-2000

Principal ResearcherClayton D. Apps, RPBio1

Primary Field ResearchersRhonda OwcharBryce Bateman

Project PartnersAlan Dibb, Parks Canada

Gary Tipper, BC EnvironmentJohn Krebs, CBFWCP

FundingHabitat Conservation Trust

Columbia Basin Fish and Wildlife Compensation ProgramParks Canada

May 2000

This report contains preliminary results, analyses, and conclusions from a continuingstudy, and is not to be cited or distributed without the permission of the author.

1Aspen Wildlife [email protected]

Southern Canadian Rockies Lynx Project • 1999/2000 Summary Report • C. Apps • May 2000 1

Acknowledgments

Funding for the Southern Canadian Rockies Lynx Project 1999 - 2000 field season wasprovided by the Habitat Conservation Trust, the Columbia Basin Fish and WildlifeCompensation Program, and Parks Canada. In-kind contributions were also provided byParks Canada and the Columbia Basin Fish and Wildlife Compensation Program. Projectadministration was provided by Alan Dibb of Parks Canada, Gary Tipper of BCEnvironment, and John Krebs of the Columbia Basin Fish and Wildlife CompensationProgram. Liaison between the field program and operational management within ParksCanada was provided by Jim Mamalis, Joe Owchar, and Barb Bertch. Skiing Louiseprovided free lift access for telemetry work.

Many thanks to Bryce Bateman and Rhonda Owchar for their continued dedication, effort,and skill as primary field researchers this year. Field assistance this year was also capablyprovided by Wayne Holstrom, Dave Lewis, Steve Bertollo, and John Flaa. Laboratorywork was conducted by Todd Shury. Thanks also to the following individuals whocontributed to this work during previous years: Doerte Poszig, Peter Wandeler, SaundiStevens, Mike Vassal, Dave Quinn, Chris Darimont, Barb Bertch, Brian Baxter, JasonBryan, Mark Hebblewhite, John Weaver, Stuart Hawes, Diana Ghikas, Trevor Kinley, andNancy Newhouse.


Summary

The southern Canadian Rocky Mountains support a Canada lynx population near thespecies’ southwestern range extent, although inherent habitat potential varies greatly.Here and throughout the western mountains of North America, heightened concern forlynx conservation relates primarily to intensive forest management and human-causedhabitat fragmentation at several scales. Relative to northern populations, southwesternlynx populations may exhibit ecotypic differences that lessen their resilience to habitat andpopulation mismanagement. However, because associated hypotheses have not beenaddressed with field research, it is difficult to assess the influence of land management onthe health and viability of local populations or to account for lynx requirements at theappropriate planning levels. In response to this and the recognition of the lynx as a focalspecies in managing for biodiversity across spatial and temporal scales, the SouthernCanadian Rockies Lynx Project was initiated as a cooperative research effort between BCEnvironment and Parks Canada. Principle objectives are to 1) describe the broad habitatassociations and current/potential population distribution of lynx in southeastern BC; 2)describe lynx-habitat relationships specific to foraging, travel, and denning at stand andlandscape levels; 3) describe aspects of lynx ecology relating to space use, movements,diet, and demographics relative to habitat and prey variation; and 4) develop a scale-dependent model of lynx habitat suitability and associated management guidelines.

The regional focal area for this study includes the southern Canadian Rocky Mountains ofsoutheastern British Columbia and southwestern Alberta. Within this, the field study areaencompasses the Beaverfoot and upper Kootenay drainages on BC provincial land, and theVermilion, middle Bow, and Kicking Horse drainages within Kootenay, Banff, and Yohonational parks. Preliminary results and field evidence suggest the greater study area ofapproximately 3500 km2 may support 14 to 18 resident lynx during the increase to highphase of an eight to 11 year snowshoe hare population cycle.

The first three years of this study coincided with the increase to high phase of a snowshoehare cycle, facilitating comparisons with populations studied in boreal ecosystems. Basedon annual sampling during this period, hare densities varied greatly throughout the studyarea but have been low, ranging from 0.01 – 0.47 hares/ha1. Maximum hare densitieswere ≤ 7% of those reported in northern lynx study areas during previous populationpeaks, and were near the minimum hypothetical threshold thought to sustain resident lynx.The fourth year (1999/2000) coincided with the crash phase, documented in northern andsouthern regions of lynx and snowshoe hare range in western North America. Harepopulations may have declined during 1998/99, and probably did so during 1999/2000, butsampling was not complete at the time of this report. Field observations during 1999/2000were consistent with a marked decline, in accordance with evidence to date that southernhare populations cycle two to 25 fold in synchrony with boreal populations.

1 Among 3 overstory cover types and 3 landscape blocks. Sampling areas do not necessarily coincide withlandscapes that support resident lynx or habitats used by resident lynx.


During the first three years, lynx space use, movements, diet, and demographics resembledthat of northern populations during cyclic hare lows, consistent with the apparent lowdensities of their primary prey. Determined by snowtracking, winter diet (n = 172 killsover 4 years) was diverse and included hares (56% of kills), red squirrels (29%), northernflying squirrels (4%), martens (3%), grouse (2%), voles (2%), and mule deer (1%). Thelarge proportion of prey alternate to hares, including ungulate kills, is consistent withopportunistic food habits previously reported during hare lows. Kitten recruitment towinter was zero among litters of adult study animals (4 lynx-years), but one to threeuncollared family groups of two kittens each occurred to spring in the study area duringeach of the first three years. Although habitat quality may have varied among femalehome ranges, overall kitten production and survival was notably lower than what would beexpected during the high phase of the cycle. Among residents, multi-annual home ranges(90% adaptive kernel estimator; 4 years) averaged 221 and 222 km2 for females and malesrespectively. Home ranges were roughly five to ten times larger than those reported forresident lynx in previous studies, even during the low phase of the hare cycle. Minimumdaily movements (first 3 years) were 3.5 and 4.0 km/day for females and malesrespectively. This is higher than reported for most studies of comparable methods in theboreal forest, implying greater foraging effort, but similar to results of one study only afterhare densities declined below 1.0 hares/ha.

The dispersals of four juvenile lynx (2M, 2F) were documented during the first three years.Of the three with known fates, dispersals were short (17 – 44 km) and ended in starvation.Limited home range vacancy is expected during the high phase of the hare cycle, and ahigh dispersal rate and low survival of juveniles may be expected in an unexploitedpopulation with limited availability and distribution of potential habitat. Most adult lynxalso made extensive exploratory movements at various times of the year, consistent withthe patchy distribution of the resident population, but usually returned to core homeranges. However, one adult male dispersed from an established (>1 year) home rangeduring June 1998. This lynx was eventually trapped and killed north of Swan Hills,Alberta in January 2000, representing a minimum linear dispersal of 498 km. Longdistance dispersal by adult lynx has been reported from northern study areas, but onlyduring the snowshoe hare crash.

Several notable changes in lynx behaviour and vital rates occurred during winter1999/2000. A minimum of six kittens were born to five resident females, but nonesurvived past early winter. This is consistent with reports for other populations followedthrough the hare crash. Although annual survival among resident adults was high (0.90)during the first three years, six of eight resident study animals died during late winter2000, again similar to adult survival in other unexploited populations followed through thehare crash. Five mortalities were due to starvation and one was likely due to predation bya wolverine, although energetic stress may have rendered this lynx vulnerable to predation.

Lynx were snowtracked for 1208 km over the four winters. During the first three years,lynx made an average of one successful hare kill for each 11 km snowtracked. However,the hare kill rate decreased to one per 28 km during 1999/2000, suggesting a considerable


increase in foraging effort. Contrary to expectation, the proportion of alternate prey inlynx diet did not increase this year. The squirrel kill rate also decreased from an averageof one per 20 km during the first three years, to one per 94 km during year four. Redsquirrel populations are also known to fluctuate with conifer cone production, and oneIdaho study has documented a marked population decline of both squirrels and hares overthe past two years. In some southern lynx populations, resident animals may persist inlandscapes where, during the increase to high phase of the cycle, hare densities are abovebut still near the minimum threshold that can sustain them. However, during the declinephase, even a subtle drop in hare densities could have quick and dramatic effects on lynxpopulation vital rates relative to northern regions where hare populations drop from muchhigher peak densities. Although our preliminary results support this hypothesis,population effects may be further exacerbated by a coincidental decline in alternate preypopulations and increased competition in landscapes where human use may facilitate theyear-round persistence of canid competitors.

Results to date suggest that the lynx population in the southern Canadian RockyMountains is not subject to dramatic pulses of increased productivity reported for northernpopulations, and that alternate prey may be important in sustaining lynx throughout thehare cycle. Only some landscapes with suitable terrain and climatic conditions will supportresident lynx, resulting in a patchy distribution of the regional population. However,through the low phase of the hare cycle, lynx may persist in even fewer pockets with thehighest quality habitat. In addition, preliminary analyses showed that lynx crossed majorhighways within their home ranges less than random expectation, suggesting a negativeinfluence on movements. Crossing frequency appears to be at least a partial function oftraffic volume and highway allowance. Residents also exhibited topographic preferenceswithin home ranges, with seasonal differences that may relate to habitat partitioning inresponse to competing predators. Preliminary conservation recommendations areprovided.


Table of Contents

Acknowledgements…………………………………………………………………………………. 1

Summary……………………………………………………………………………………………... 2

1.0 Introduction……………………………………………………………………………………... 7

2.0 Study Area…….……………………………………………………………………….……….. 8

3.0 Regional Lynx Population Distribution and Habitat Associations….…………………….. 10

4.0 Snowshoe Hare Densities by Cover Type and Landscape……………………………….. 10

5.0 Study Animals………..………………………………………………………………………… 12

5.1 Capture……………………….…………….…………………………………………. 12

5.2 Detection of Unmarked Lynx…………….…………………………………………. 12

6.0 Radio-telemetry…….……………………………………………………………………..……. 14

6.1 Adult Survival………………………………………………………………………... 14

6.2 Reproductive Ecology and Kitten Survival.……………………………………… 16

6.3 Space Use and Movements of Resident Lynx….………………………………… 17

6.4 Adult Dispersal…….…………………………………………………………………. 22

6.5 Juvenile Dispersal and Survival…..………………………………………………... 22

6.6 Habitat Selection……………………………………………………………………... 24

6.7 Highway Influence………………………………………………………………….... 25

7.0 Snow-tracking…………………..………………………………………………….…………... 25

7.1 Winter Food Habits………....……………………………………………………….. 25

7.2 Habitat and Behaviour………………………………………………………………. 27

7.3 Positional Error of Movement Routes………..………………….………………… 27

8.0 Collaboration with other Research…………………………………………………………... 28

9.0 Communications and Consultations…………………………………………………………. 29

10.0 Discussion of Vital Rate Changes during 1999/2000……..…………………………….. 32

11.0 Local and Regional Conservation Recommendations……….………………………….. 32

Literature Cited………………………………………………………………………………………. 35


List of Figures

Figure 1. Southern Canadian Rockies Lynx Project focal region and study area within thesouthern Canadian Rocky Mountains………………………………………….……………... 9

Figure 2. Snowshoe hare densities (± 95% C.I.) among 3 landscapes and 3 successional stagesin the southern Canadian Rocky Mountains, 1996 – 1998………………………………….. 11

Figure 3. Snowshoe hare densities (± 95% C.I.) by landscape and year in the southern CanadianRocky Mountains, 1996 – 1999. Note: results are not adjusted to reflect the differentialarea of each successional stage in each landscape…...……………………………………. 11

Figure 4. Adaptive Kernel 90% multi-annual utilization distributions forresident lynx in thesouthern Canadian Rocky Mountains, 1996 – 2000…..…………….…………….………… 20

Figure 5. Minimum convex polygons (100%) of multi-annual radiolocations for resident lynx inthe southern Canadian Rocky Mountains, 1996 – 2000………..……….………………….. 21

Figure 6. Minimum linear dispersal movement of one adult male (M/03) from an established (>1yr) home range in the southern Canadian Rocky Mountains……………………………….. 23

Figure 7. Dispersal movements of 2 of 4 juvenile lynx monitored in the southern Canadian RockyMountains, 1996 – 1999………………………………………………………………………… 24

Figure 8. Lynx kills in the southern Canadian Rocky Mountains, 1996 – 2000, as determined bysnow-tracking………………………………………………….…………………………………. 26

Figure 9. Total bihourly activities (total events and activity by 1 min interval) and number of daysthat M/01 interfered with camera monitor while feeding on a mule deer kill over an 11day period.……………………………………………………………………………………….. 27

Figure 10. Positional error associated with Eagle Explorer non-differentially correctable GPS units,as determined from 129 locations obtained at fixed stations………………………………. 28

List of Tables

Table 1. Southern Canadian Rockies Lynx Project animal capture summary, 1996 – 2000……… 13

Table 2. Summary of telemetry and snow-tracking data/samples collected from radio-collaredlynx in the southern Canadian Rocky Mountains, 1996 – 2000………………….….……... 15

Table 3. Reproduction and kitten survival among resident lynx during 1999 – 2000 in thesouthern Canadian Rocky Mountains…………………………………………………………. 17

Table 4. Minimum convex polygon (MCP) and adaptive kernel (ADK) multi-annual home rangeestimates for lynx in the southern Canadian Rocky Mountains, 1996 – 2000………..…... 18

Table 5. Minimum daily movements (MDM) of lynx in the southern Canadian Rocky Mountains,from sequential radiolocations independent by 18-36 h, to March 2000………………….. 19


1.0 Introduction

The Canada lynx (Lynx canadensis) has been of regional conservation concern insoutheastern British Columbia since the early 1980's, when harvest trends indicated thatpopulations were not recovering to levels observed in past decades (A. Fontana, BCEnvironment, pers. comm.). Since then, populations have not rebounded and, as indicatedby a regional trapper survey (Apps 1996a), may have continued to decline in some areas.Similar concerns have resounded across other provincial and state jurisdictions at or nearthe southwestern extent of lynx range (Koehler and Aubry 1994), prompting the recentlisting of the lynx under the US Endangered Species Act (USFWS 2000). Near theirsouthwestern range extent, lynx populations are expected to occur at low densities withinpeninsular and often disjunct distributions. The recovery and continued viability of thesefringe populations requires functional habitat and genetic connectivity with morecontiguous northern populations, for which southeast BC and the southern CanadianRockies represent an important interface.

Intensive forest management is considered the greatest threat to the conservation ofsouthwestern lynx populations (Hatler 1988, Weaver 1993, Koehler and Aubry 1994). Itis expected that fragmentation associated with this and other human impacts will beamplified in mountainous landscapes where suitable habitat is naturally discontinuous. Forlynx, such effects are likely manifested at several scales. Individuals may respond towithin-stand conditions and to landscape composition and structure. At broader scales,the density and spatial distribution of populations may relate to habitat quality anddistribution across the regional landscape, with fragmentation influencing dispersal successand genetic connectivity among subpopulations.

The impetus for this research stems from conservation concern not only for lynx but forthe persistence and integrity of those natural communities with which they and other avianand terrestrial predators are associated. Because they are expected to respond to habitatconditions at multiple scales, lynx are an important focal species for assessing the efficacyof hierarchical biodiversity management strategies specific to montane and subalpineecosystems. However, relative to other carnivore species, we know little of lynx ecologyand associated ecotypic variation near their southwest range extent.

The primary objectives for the Southern Canadian Rockies Lynx Project are to:

1. describe the broad habitat associations and the current and potential populationdistribution of lynx in southeastern BC;

2. describe lynx-habitat relationships, specific to foraging, travel and denning, at standand landscape levels;

3. describe aspects of lynx ecology relating to space use, movements, diet anddemographics relative to habitat and prey variation; and

4. develop a scale-dependent model of lynx habitat suitability and associated managementguidelines.


The ultimate aim of this work is to facilitate the integration of probable stand andlandscape requirements for lynx with biodiversity management at the appropriate planninglevels.

This report is a summary of work carried out on the Southern Canadian Rockies LynxProject during year four of field study, May 1999 to May 2000, and of progress to date.The project working plan (Apps 1996b) provides additional information on researchbackground, design, and study area selection. Two publications have resulted from thisresearch to date (Apps 2000; Apps et al. in press) based on data collected to October,1998.

2.0 Study Area

The regional focal area for this research is the East Kootenay wildlife management sub-region of southeastern BC. The field study area falls largely within this region and thesouthern Canadian Rocky Mountains of British Columbia and Alberta (Figure 1). Itencompasses the Beaverfoot and upper Kootenay drainages on BC provincial land, and theVermilion, middle Bow and Kicking Horse valleys within Kootenay, Banff and Yohonational parks. Overall, the greater study area makes up roughly 3500 km2, althoughmuch of this is rock, ice, and other inherently unsuitable lynx habitat. Elevations rangefrom 1200 m to > 3000 m, and span the Interior Cedar-Hemlock, Montane Spruce,Englemann Spruce-Subalpine Fir, and Alpine Tundra biogeoclimatic zones (Meidinger andPojar 1991). The macroclimate is continental, with mean temperatures ranging from –18°C (January) to 23°C (July) and 42-63% of the 51-81 cm mean annual precipitationfalling as snow (Achuff et al. 1984). Natural and human conditions vary throughout thestudy area, and linear features include 2- to 4- lane paved highways with annual (1994)average daily traffic volumes of 1119 – 8322 vehicles per day (A. Clevenger, BanffNational Park, pers. comm.). Several other predators and potential competitors and adiversity of potential prey occur within the study area (Scotter and Ulrich 1995, Scotter etal. 1990).

Because the study area encompasses much variation in natural and man-caused habitatconditions, it is appropriate for addressing multi-scale questions of habitat suitability. It isalso associated with various types and levels of human development, providingopportunity to address questions of lynx response to specific disturbance. Logistically, thearea is appropriate for the intensive data collection that is required to address these andother field-level research objectives, and it is large enough that a minimum animal samplecan be achieved. Despite a limited hunting and trapping season in British Columbia, lynxare not known to have been harvested on the provincial section of the study area for atleast 15 years. Based on this, and the protection from harvest afforded to lynx within thenational parks, I consider the resident population to be unexploited.


Figure 1. Southern Canadian Rockies Lynx Project focal region (grey) and study area(red) within the southern Canadian Rocky Mountains.


3.0 Regional Lynx Population Distribution and Habitat Associations

Data collected through the East Kootenay Regional Trapper Survey have been compiledand results summarized (Apps 1996a). We are near completion of a regional 1:20,000habitat and human use database, against which a 13-year harvest database will be analyzedin conjunction with the trapper survey. This will discern habitat and human factors thatinfluence distribution, relative abundance, and vulnerability of lynx and bobcats to human-caused mortality.

4.0 Snowshoe Hare Densities by Cover Type and Landscape

Methods.—The range of snowshoe hare (Lepus americanus) densities occurring withinthe study area have been estimated using established 305 × 5.1 cm quadrats (Krebs et al.1987). Ten quadrats were spaced at 30.5 m intervals along each of 61 transects.Transects were placed randomly among early- (20 - 60 y), mid- (60 – 120 y), and late-(>120 y) successional stands, spaced by ≥ 500 m. Sampling was further stratified among 3landscapes within the study area, representing geographically distinct areas where Iconsider terrain conditions conducive to supporting lynx. Sampling was conducted in anadditional landscape (middle Bow Valley) during 1998 and 1999. Hare densities werecalculated with a modified version of the Krebs et al. (1987) regression equation, using theHARETURD program (27 October, 1998; C.J. Krebs, University of British Columbia,pers. comm.).

Track counts were conducted to provide an additional index of relative snowshoe hareabundance among landscapes. A technician slowly drove highways dissecting the studyarea 1 – 2 days after snowfall and recorded the distance of observable habitat/roadallowance edge and associated hare tracks per km.

Results--For early- mid- and late-successional stands respectively, mean hare densities bylandscape were 0.16/ha, 0.08/ha, and 0.01/ha in the upper Kootenay Valley, 0.25/ha,0.06/ha, and 0.10/ha in the Beaverfoot Valley, and 0.47/ha, 0.39/ha, and 0.32/ha in theVermilion Pass (Figure 2). Among landscapes, a slight but consistent decline wasdetected from 1998 to 1999 (Figure 3). However, variation among years has not beenstatistically tested at the time of this report. It should be noted that in all landscapes,many, if not most pellet plots fall in sites where there is no evidence that a lynx core homerange occurs. Thus, it is not necessarily appropriate to infer minimum hare densities thatwill support lynx from these results. Moreover, our sampling design is based on methodsdeveloped in a boreal ecosystem with much higher hare populations, and due to thenumber of “zero-counts” in our plots, it may be difficult to detect variation among years ifit is occurring.


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1997 1998 1999

Figure 2. Snowshoe hare densities (±± 95% C.I.) among 3 landscapes and 3successional stages in the southern Canadian Rocky Mountains, 1997/98.

Figure 3. Snowshoe hare densities (±± 95% C.I.) by landscape and year in the southernCanadian Rocky Mountains, 1996 – 1999. Note: results are not adjusted to reflect thedifferential area of each successional stage in each landscape.


5.0 Study Animals

5.1 Capture

Methods.— Details of animal capture and handling methods are described in the projectworking plan and in a separate capture and handling protocol that is available uponrequest. Over the previous four years, study animal capture work occurred during eitherearly winter (Oct. – Nov.) or late winter (Mar. – Apr.). To date, most animals werecaptured using live traps. These have been either #3 “soft-catch” foot-hold traps with #2springs in cubby sets, or 122 cm × 51 cm × 66 cm cage-traps. Trap stations were baitedwith visual attractants and scent lures. Efforts were focused in the following landscapeswhere lynx sign has been observed: the Beaverfoot Valley, upper Kootenay Valley,Vermilion Pass, and the middle Bow Valley.

This year we employed the use of a Walker hound and a Redbone hound /Blue-tick houndcross to track and tree potential study animals, and capture work was conducted onlyduring early winter. Between 16 October and 18 December, at least three field techniciansfocused track searches in target landscapes when and where snow conditions wereconducive. When tracks were detected, lynx were front-tracked to ensure that the trailwas fresh (i.e. < 6 hrs old) before the two dogs were deployed. Treed lynx wereimmobilized using either an extendable syringe-pole, a 1cc Pneu-dart immobilizationdart, and/or a hand-syringe. All handling methods were as detailed in the captureprotocol.

Results.— Hounds were deployed to track and tree lynx on ten occasions during earlywinter. Lynx were not treed on four occasions, most likely due to marginal snow andtrack conditions. On one occasion, a cougar was incidentally treed on the Lake Louise skihill. On one occasion, a lynx was treed but was too high to attempt immobilization. Lynxwere successfully treed and immobilized on five other occasions. This included onecapture in the Vermilion Pass, three in the middle Bow Valley, and one in the BeaverfootValley. To date, 17 lynx have been captured 19 times (Table 1). Capture efforts andresults during previous years are detailed in previous annual summary reports.

5.2 Detection of Unmarked Lynx

Methods.—The occurrence of uncollared lynx within the study area was noted by fieldpersonnel throughout the winter during lynx snow-tracking and track searches. Track oranimal observations reported by other individuals were also collected.

Results.— One uncollared family group with at least two kittens was detected on twodates during early winter in the vicinity of Symond and Whitetail creeks in the upperKootenay Valley. One adult was detected in the Beaverfoot Valley on three dates in earlywinter and on one date during late winter. At least one uncollared adult was detected oneither side of the Trans-Canada Highway in the middle Bow Valley during mid-winter andat least one detection occurred during late winter. One adult lynx was detected duringmid winter near Johnston Creek east of Castle Junction. One adult male was detected


Table 1. Southern Canadian Rockies Lynx Project animal capture summary, 1996 - 2000.

Animal Sex/ID CaptureCapture

DateGeneralLocation

AgeClassa

Weight(kg)

Length(cm)b

ToothWear

ConditionClass

M/01 Stanley 3 11/04/96 Vermilion Pass A 13.6 113.0 Min Good

M/02 Fraser 1 03/20/97 Beaverfoot A 12.5 111.0 Min Fair

M/03 Clawson 1 03/21/97 Beaverfoot A 13.6 112.0 Mod Good

M/04 Storm 1 11/14/97 Vermilion Pass K 5.5 71.0 No Fair

M/05 Hector 1 03/19/98 Middle Bow J/A 11.3 99.0 Min Fair

M/00 Unnamed 1 03/31/98 Middle Bow A 9.5 107.0 Maj Poor

M/06 Parker 1 03/19/99 Middle Bow J 9.9 96.0 Min Fair

M/07 Bosworth 1 11/28/99 Middle Bow A 15.0 110.0 Mod Good

F/01 Rudi 1 11/05/96 Vermilion Pass A 11.8 97.5 Mod Good

F/02 Tanu 1 10/28/97 Middle Bow A 10.9 102.0 Mod Good

F/03 Numa 1 03/22/98 Vermilion Pass J 9.5 97.0 No Good

F/04 Oesa 1 11/22/97 Middle Bow A 10.9 96.5 Mod Good

F/05 Shelagh 1 12/01/97 Beaverfoot K/J 5.5 No Fair

F/06 Margaret 1 11/25/99 Middle Bow A 10.0 94.0 Min Fair

F/07 Chickadee 1 11/26/99 Vermilion Pass A 11.3 100.5 Min Good

F/08 Helen 1 12/05/99 Middle Bow A 10.7 94.0 Mod Fair

F/09 Aquila 1 12/17/99 Beaverfoot J/A 10.4 98.5 Min Good

a A = adult (> 2 yrs), J = juvenile (1 – 2 yrs), K = kitten (< 1 yr)b dorsal: nose to tip of tail


traveling with F/06 on at least three dates during late winter and was directly observed onone other date. Landscapes where uncollared lynx were detected this year are consistentwith observations during previous years. However, in all parts of the study area, lynxwere not detected as frequently during late winter 2000 as in previous years.

6.0 Radiotelemetry

Methods.—Radiotelemetry was conducted following protocol outlined in the projectworking plan, with the vast majority of data collected using ground-based methods.Ground locations were estimated from 3 bearings taken within 1.5 hours, with originslocated using a GPS receiver. Although monitoring frequency has varied among animals,daily radiolocations have been obtained at least 5 days per week for most (Table 2). M/02has been located with lower frequency due to poorer ground accessibility. Intensivemonitoring has been conducted for several animals to address questions of movementpatterns at finer temporal scales, with locations obtained every 2 – 4 hours for up to 48hours. All radiotelemetry data are maintained in a digital database. Lynx carcasses and/orradiocollars were retrieved as soon as possible after mortality was detected, and probablecause of death was determined from necropsies and/or field evidence.

6.1 Adult Survival

Results.—Adult survival was high (0.90) during the first three years of study, with a singlemortality due to starvation and/or Helicobacter infection. During the fourth year of study,annual survival was low (0.25), with natural mortalities in six of eight resident adults in thestudy sample. This was coincident with a known population decline of snowshoe haresand possibly red squirrels (see Discussion). One mortality (M/01) occurred in January,four (F/04, F/06, F/08, F/09) occurred in Feburay, and one (M/07) occurred in March.The ultimate cause of five mortalities is thought to have been starvation. However,histology tests were not complete at the time of this report, and disease cannot be ruledout as an exacerbating factor. We are providing samples to one investigator studyingdifferences in the role of disease between peripheral and core lynx populations (M. Poss,University of Montana, pers. comm.). One lynx (F/04) appears to have been either killedor scavenged by a wolverine. All that could be recovered after her death was her head andneck, cached in the snow. If this animal was not killed but scavenged, the wolverine musthave found her within three days of her death. Cursory inspection of fat depositsassociated with remains that were recovered was not consistent with death fromstarvation. However, it is possible that this lynx was energetically stressed and thus moresusceptible to predation.


Table 2. Summary of telemetry and snow-tracking data/samples collected from radio-collared lynx in the southernCanadian Rocky Mountains to April, 2000.

Animal IDPeriod

Monitored

Totaltelemetrylocations

Dailytelemetrylocationsa

Snow-trackSequences

Snow-trackKm Kill Sites Scats

M/01 Stanley 11/96 – 01/00 1047 778 105 356 51 58

M/02 Fraser 03/97 – 04/00 292 263 17 45 7 10

M/03 Clawson 03/97 – 06/98 261 189 13 24 3 5

M/04 Storm 11/97 – 12/97 11 7 0 0 0 0

M/05 Hector 03/98 – 05/98 61 35 6 7 6 4

M/06 Parker 03/99 – 04/99 17 17 0 0 0 0

M/07 Bosworth 11/99 – 03/00 104 90 36 113 5 13

F/01 Rudi 11/96 – 09/97 257 200 11 28 5 9

F/02 Tanu 11/97 – 11/98 419 254 137 334 69(10)b 101(20)b

F/03 Numa 03/98 – 04/98 33 23 5 13 1 26

F/04 Oesa 04/08 – 02/00 370 334 26 102 10 10

F/05 Shelagh 05/98 – 10/98 38 37 0 0 0 0

F/06 Margaret 11/00 – 02/00 58 56 3 11 0 0

F/07 Chickadee 11/99 – 04/00 82 81 21 85 8 13

F/08 Helen 12/99 – 02/00 40 39 9 51 2 3

F/09 Aquila 12/99 – 02/00 7 7 2 4 1 1

UnknownVP 9 25 3 6

UnknownMBV 10 10 1 2

TOTALS 3097 2410 410 1208 172 261a independent by >24 hrsb the number recorded while F/02 was traveling with an unmarked male are in parentheses.VP = Uncollared lynx in the Vermilion PassMBV = Uncollared lynx in the Middle Bow Valley


6.2 Reproductive Ecology and Kitten Survival

Results.—Over the course of the project, two known natal den sites have been identifiedand their use monitored. The movements of a female (F/02) known to have bred weremonitored during the natal denning period May – July, 1998. Radiolocations of this lynxwere within 500 m of an activity center on 82% of days between 22 May and 14 July, andshe was observed with one kitten in August. The denning site was central within her homerange, and ground inspection indicated that it was a mesic site within an old (>250 yrs)stand dominated by Engelmann Spruce (Picea engelmannii). Extensive low-levelstructure was apparent, resulting from many downed and uprooted large diameter trees.The daily locations of another female (F/01) were within 250 m of an activity centerbetween 20 May and 1 June, 1997, after which she resumed movements and did not re-visit the presumed den site. From this, I inferred that she lost or abandoned her litter at orjust after parturition.

One adult female (F/04) was monitored during the 1999 denning period. She spent atleast three days near the eventual den site in mid-May, then began using it consistently asof 3 to 5 June. The site was located approximately 7 km from the Bow valley up theBaker Creek valley. After documenting consistent use of the site for 17 days, weconducted a visit on 22 June and recorded two kittens at approximately 15 days old (est.birth date 4 – 7 June). Five 48-hour intensive monitoring sessions were conducted duringJune and July, and two multi-day sessions of continuous activity monitoring wereconducted using a datalogging receiver. Two extensive forays away from the den sitewere recorded to 5 August, after which the den site was no longer consistently used andnormal movements appeared to resume. Early winter snowtracking confirmed the survivalof one kitten to 11 November.

During the first three years of study, no kittens are known to have survived to early winterwithin the study sample (4 lynx-years). However, uncollared family groups are known tohave occurred within the study area through to late winter of each year. One group withtwo kittens was detected during 1996 – 97, three groups with two kittens were detectedduring 1997-98, and two groups with at least two kittens were detected during 1998-99.During 1999-2000, no kitten survival was documented past December. Based on the densite visit, snowtracking, and placental scar counts, known litter sizes ranged from zero totwo among study animals, and no kittens survived past December (Table 3).


Table 3. Reproduction and kitten survival among radiocollared lynx during 1999 – 2000 inthe southern Canadian Rocky Mountains.

Lynx Kits Evidence

Latest survival

month

F/04 2 (known) Den site Nov

F/05 1 (known) Tracking and in utero a Dec

F/06 1 (min.) Tracking Dec

F/07 2 (max.) In utero a Nov

F/08 0 (known) In utero a --a Placental scars

6.3 Space Use and Movements of Resident Lynx

Methods.—From the study sample to date, eleven lynx (4M, 7F) were known to beresident adults, exhibiting distinct home range affinity. Several measures of movement andspace use were determined for these animals. I estimated home ranges of resident adultsseasonally and annually, with seasons defined by the typical 6-month snow and snow-freeperiods beginning 1 November and 1 May. Home ranges were estimated using the 90%minimum convex polygon (MCP) and the 95%, 75%, and 50% adaptive kernel utilizationdistributions (UD; Kie et al. 1996). Lynx movements indicated that home ranges could betraversed within 24 h; thus only radiolocations that were temporally independent by thisinterval were used (Swihart and Slade 1985). I defined minimum daily movements

(MDM) as the distance between sequential radiolocations separated by 18 – 36 h ( × = 24h), and I determined mean MDM for each resident annually and by season.

Results.-- From data pooled over the four years, the mean multi-annual 90% UD was 221km2 (n = 4, SD = 64) for resident males and 222 km2 (n = 7 SD = 209) for residentfemales (Table 4, Figure 4). As indicated by 100% MCP home ranges (Figure 5),extraterritorial forays were occasionally made by all lynx. Although such movementsoccurred throughout the year, they were more prevalent during the late February to earlyApril breeding period. Annual MDM averaged 4.0 km (n = 3, SD = 0.8) for males and 3.5km (n = 3, SD = 1.0) for females (Table 5). Analyses of season and sex differences inspace use and movements are detailed by Apps (2000) and Apps et al. (in press).


Table 4. Minimum convex polygon (MCP) and adaptive kernel (ADK) multi-annual homerange estimates for resident lynx in the southern Canadian Rocky Mountains, 1996 – 2000.

M C P A D KAnimal Sex/ID Monitored n 100% 90% 95% 90% 75% 50%M/01 Stanley 11/96 – 01/00 778 1003 259 477 270 117 31

M/02 Fraser 03/97 – 04/00 263 433 157 388 240 117 40

M/03 Clawson 03/97 – 06/98 189 346 154 330 246 49 21

M/04 Storm 11/97 – 12/97 -- -- -- -- -- -- --

M/05 Hector 03/98 – 05/98 -- -- -- -- -- -- --

M/06 Parker 03/99 – 04/99 -- -- -- -- -- -- --

M/07 Bosworth 11/99 – 03/00 90 232 70 272 127 54 25

F/01 Rudi 11/96 – 09/97 200 408 211 505 205 85 27

F/02 Tanu 11/97 – 11/98 255 133 64 94 62 25 10

F/03 Numa 03/98 – 04/98 -- -- -- -- -- -- --

F/04 Oesa 04/08 – 02/00 334 181 60 117 73 42 15

F/05 Shelagh 05/98 – 10/98 34 321 108 563 499 51 3

F/06 Margaret 11/00 – 02/00 56 138 64 195 130 72 29

F/07 Chickadee 11/99 – 04/00 76 84 25 67 48 26 13

F/08 Helen 12/99 – 02/00 40 319 273 611 535 272 55

F/09 Aquila 12/99 – 02/00 7 21 21 -- -- 69 9

Male Mean 504 160 367 221 84 29

Male SD 343 77 87 64 38 8

Female Mean 201 103 307 222 80 20

Female SD 134 91 241 209 80 17


Table 5. Minimum daily movements (MDM) of lynx in the southern Canadian RockyMountains, from sequential radiolocations independent by 18 - 36 hrs, to March, 2000.

Minimum Daily Movements (km)Animal ID Monitored n Range Mean S.D.M/01 Stanley 11/96 – 01/00 577 0.1 - 25.2 3.6 3.3

M/02 Fraser 03/97 – 04/00 292 0.1 - 23.2 3.9 4.2

M/03 Clawson 03/97 – 06/98 128 0.3 - 19.8 3.0 2.6

M/04 Bosworth 11/99 – 03/00 73 0.1 - 11.7 2.4 2.2

F/01 Rudir 11/96 – 09/97 151 0.1 - 12 2.8 2.5

F/02 Tanu 11/97 – 11/98 169 0.1 - 10.9 2.5 2.1

F/04 Oesa 04/08 – 02/00 158 0.2 - 13.7 3.2 2.5

F/05 Shelagh 05/98 – 10/98 12 0.4 - 13.2 3.3 3.7

F/06 Margaret 11/00 – 02/00 40 0.1 - 10.3 3.2 2.4

F/07 Chickadee 11/99 – 04/00 46 0.3 - 11.3 2.1 2.2

F/08 Helen 12/99 – 02/00 21 0.1 - 6.1 1.9 1.7

F/09 Aquila 12/99 – 02/00

Males 3.2 0.7

Females 2.7 0.6


Figure 4. Adaptive kernel 90% multi-annual utilization distributions for resident lynx inthe southern Canadian Rocky Mountains, 1996 - 2000.


Figure 5. Multi-annual 100% minimum convex polygons home ranges for resident lynx inthe southern Canadian Rocky Mountains, 1996 – 2000, illustrating the extent ofexploratory movements.


6.4 Adult Dispersal

Results.— Field evidence during 1998/99 suggested that one adult dispersal had occurred.In early June, 1998, M/03 was monitored while on a presumed exploratory movement.After radio-contact was lost, three aircraft searches were conducted over the next twomonths, one of which covered all drainages within a 35 km radius of his home range. TheWest Slopes Bear Study also continued to scan his frequency during their telemetryflights. Although hair-snag sampling within his home range was successful at several sitesin 1997, similar sampling was not successful in the fall of 1998. This, and the fact thatM/04 was outside his home range when contact was lost, is more suggestive of homerange abandonment than radiocollar failure, but we had no conclusive evidence that he hadin fact dispersed. On 25 January, 2000 M/03 was legally trapped and killed north of SwanHills, Alberta. This represents a minimum linear movement of 498 km from his KickingHorse Valley home range (Figure 6).

6.5 Juvenile Dispersal and Survival

Methods.—Lynx exhibiting consistent, linear movements were defined as transient ordispersing animals. Where possible, their movements were summarized with respect totiming of dispersal, direction, distance, MDM, and proximate outcome.

Results.— Five juvenile lynx have been radiocollared over the course of study. Four ofthese lynx were known to disperse from their natal home range. A juvenile male (M/05)made a 44 km southeast movement down the Bow Valley over 11 days, beginning inMarch, 1998. He then used one meadow complex intensively for the next 31 days beforehis death on 3 May. After she was radiocollared within her natal range in late March,1998, a juvenile female (F/03) re-joined her mother for three days, then movedindependently for three days before 28 March when she initiated a southeast linearmovement of 17 km over three days. She then used one area intensively for the next 15days before her death. The movements of both animals followed the major valley in whichthey occurred and paralleled but did not cross the Trans-Canada Highway (Figure 7). InOctober 1998, female (F/05) made two separate exploratory movements from her natalrange in the Beaverfoot Valley, of a minimum 74 km and 55 km over 24 and 11 daysrespectively. The second movement continued in a southward dispersal down the upperKootenay Valley, at which time radio-contact was lost and aircraft search through theKootenay drainage north of Canal Flats failed to locate her. A juvenile male (M/06) wasmonitored from March to May, 1999 while dispersing from a presumed natal range in themiddle Bow Valley. His minimum movements totaled 38 km over 20 days in asoutheastward dispersal before his death on 7 April. Deaths of the three dispersingjuveniles were due to starvation, while one lynx (M/04) appears to have been killed byanother unmarked lynx before he presumably became independent.


Figure 6. Minimum linear dispersal movement of one adult male (M/03) from anestablished (> 1 year) home range in the southern Canadian Rocky Mountains. Dispersalwas initiated in June 1998 and the lynx was trapped and killed in January, 2000.

BRITISHCOLUMBIA

ALBERTAEdmonton

Calgary

Swan Hills

Golden

Yoho N.P.

Lesser Slave Lk

50 km


Figure 7. Dispersal movements of two of four juvenile lynx monitored in the southernCanadian Rocky Mountains, 1996 – 1999.

6.6 Habitat Selection

Methods.—Lynx selection for attributes of forest overstory has not yet been analyzed.However, I did analyze selection by resident lynx for topographic attributes availablewithin respective home ranges, based on data collected to October, 1998. Analyses werestratified by season, and mean elevation use was also compared between seasons.Methods and results are detailed by Apps (2000).

Results.—Lynx either avoided or did not select highest elevations (> 1850 m), preferredor did not avoid mid elevations (1550 – 1850 m), and although lowest elevations (< 1550m) were not avoided during winter, two animals did avoid them during summer. All malesand one female used mean elevations 90 – 133 m higher during summer than during winter(P < 0.008). However, seasonal use of elevations did not differ for two females (P >0.240). Most lynx avoided or did not select steeper (> 40%) slopes, and preferred or didnot avoid moderate (20 – 40%) or gentle (< 20%) slopes. Mean slopes used were 4 – 9%


steeper during summer than during winter for two males and two females (P < 0.009),whereas slope use did not differ between seasons for one male and one female (P >0.344). Selection for aspect varied among animals. Among all six lynx, there were nosignificant seasonal differences in the use of northward aspects (P > 0.025), and althoughone male used eastward aspects greater during summer than during winter (P = 0.011),seasonal differences were not significant among the other five lynx (P > 0.055).

6.7 Highway Influence

Methods and Results.—Paved highways dissected the 95% UD home ranges of allresident lynx in a relatively linear manner. I therefore assumed that a crossing occurred ifsequential radiolocations were obtained on opposite sides of a highway. This facilitatedtests of the null hypothesis that lynx movements were random with respect to highways.Methods and results are detailed by Apps (2000). The analysis was not stratified bytemporal, seasonal, or spatial variation in traffic volume, or by road allowance width,lanes, or juxtaposed habitat. The analysis was intended as a first approximation within amore detailed investigation of the influence of point and linear human features on lynxmovements and habitat use within home ranges. Results of this preliminary analysisindicated that all six resident lynx monitored to October, 1998 crossed highways less thanrandom expectation within respective 95% UD home ranges (P < 0.001). It appears thatcrossing frequency is at least partially a function of traffic volume and highway allowance.

7.0 Snow-Tracking

Methods.—A detailed data collection protocol was developed for the snow-trackingcomponent of this study. When snow conditions were conducive, field technicians usedradiotelemetry to cut the track of individual study animals without disturbing them.Tracks were then either back- or front-tracked, depending on the animal’s proximity andprevious tracking sessions. The movement route was logged using a GPS receiver at anapproximate 50 m interval estimated using a pace-counter. Direct and adjacent overstorycover were recorded along the animal’s movement route. Stand understory, overstory,and terrain attributes were measured at random “route” plots systematically distributedalong the tracking route. Attributes were measured at successful or attempted kill sites,resting sites, and scat locations, and associated behavioural data were also recorded.Snow penetration by lynx was measured at all plots. GPS data were downloaded to adatabase and field data associated with specific locations were entered.

7.1 Winter Food Habits

Results.—Lynx were snow-tracked for 283 km this year, for a total of 1208 km logged forat least 14 different lynx over four winters of study. This year’s tracking resulted in 18kills documented and 39 scats collected, for four year totals of 172 and 261 respectively(Table 2). Kills to date have been of 96 snowshoe hares, 49 red squirrels (Tamiasciurushudsonicus), six flying squirrels (Glaucomys sabrinus), four spruce grouse (Dendragapuscanadensis), four microtine rodents, six pine marten (Martes americanus), four unknown


species, two mule deer (Odocoileus hemionus), and one northern flicker (Colaptesauratus) (Figure 8). In addition, two cases of mule deer scavenging have beendocumented.

Figure 8. Lynx kills (n = 172) in the southern Canadian Rocky Mountains, 1996 – 2000, asdetermined by snow-tracking. Proportions represent numeric occurrence for eachspecies and do not reflect biomass.

The mule deer kills were of a fawn and a doe and were made by M/01 while outside hiscore home range. The fawn was killed on 14-15 January, 1999, and the doe was killed on18 January, 1999. Both occurred in the vicinity of Hawk Creek in the Vermilion Valleyand were separated by a distance of ~10 m. The site was at 1580 m elevation, 240 mabove the valley bottom, and was mid-position on a 15° slope of northwest aspect. Thesnow base was 1.5 m and lynx snow penetration was 9 – 14 cm. Both kills were mostlyconsumed over a period of 28 days before M/01 returned to his core use area. Details ofthe habitat attributes, prey condition, and lynx behaviour associated with the kills and theirconsumption were recorded by field researcher Doerte Poszig. A remote camera andperiodic site inspections suggested that no other species interactions occurred during thetime that the lynx was at the site. Using the Trailmaster™ camera, Doerte documentedthe 24 hour activity of M/01 over 11 days at the kill site by analyzing the number andtiming of camera beam interferences. His activity at the kill appeared to peak around mid-day and mid-night (Figure 9).

hares56%red squirrels

29%

grouse2%

marten3%

microtines2%other

3%

flying squirrels4%

mule deer1%


Figure 9. Total bihourly activities (total events and activity by 1 min interval) and numberof days that M/01 interfered with camera monitor while feeding on a mule deer kill over an11 day period. Field work and data summary contributed by Doerte Poszig.

7.2 Habitat and Behaviour

Results.—Habitat attributes and behaviour classes associated with travel routes, attemptedand successful kills, resting beds, scat locations, and conspecific interactions have not beensummarized at the time of this report.

7.3 Positional Error of Movement Routes

Methods.— All field researchers used Eagle Explorer GPS units to log lynx movementroutes, data from which are not differentially correctable. Logged vectors will thereforebe associated with a positional error that must be accounted for during data analysis.Error was gauged during 1998-99 by obtaining UTM locations approximately every twodays while measuring snow conditions at fixed stations and at different times of the day.To test the influence of canopy closure, locations were obtained within a closed forest andin an opening at each station. Fixes were also obtained at high and low elevation snowstations. I tested for differences in positional error between sites using Student’s t tests.

Results.— GPS positional error did not differ between closed canopy and open sites (P =0.505) or between high and low elevation sites (P = 0.357). Mean positional error was 56m (n = 129, SD = 43.7). Ninety percent of GPS fixes deviated from the true location by<107 m (Figure 10).

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Figure 10. Positional error associated with Eagle Explorer non-differentially correctableGPS units, as determined from 129 locations obtained at fixed stations and independentby >24 hrs.

8.0 Collaboration with Other Research

Summary.—We have and will continue to collaborate with several other researchers andinstitutions over the course of this study. This has included work with J. Weaver of theWildlife Conservation Society on feasibility and sampling designs for lynx detection andpopulation estimation from hair-snag/DNA sampling (preliminary results are describedbelow). This technique also holds promise for addressing questions of regional populationabundance, distribution, connectivity, and questions of broad-scale habitat associations; itis thus relevant to the underlying objectives of our research. This year, DNA sampleswere provided to E. Rueness of the University of Oslo, Norway for a study oncircumpolar lynx genetics. Samples have also been provided to M. Schwartz of theUniversity of Montana for his work in comparing lynx genetics between core andperipheral populations. This work will compare relatedness and genetic distance amongour study animals to that of other populations. We have also agreed to provide hair andtissue samples to J. Roth of the University of Idaho for a diet study across lynx rangeusing stable isotope methodology. Finally, we are providing organ samples fromnecropsied animals to M. Poss of the University of Montana who is looking at viralgenetics as an indicator of movements and connectivity of host lynx populations fromCanada to the U.S.A.

Vermilion Pass Hair-Snag Sampling.—A sampling effort to test mark-recaptureassumptions was applied in the Vermilion Pass during July to September, 1997. Hair-snagstations (Weaver 1997) were distributed according to a sampling grid of 25 4 mi2 cells

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centered on the composite home range of two study animals in the Vermilion Pass. Adifferent spatial distribution and checking frequency of stations was applied during each oftwo 30 day sampling sessions. In conjunction, intensive telemetry was carried out onstudy animals to document the proportion of time they spent within the grid and withineach cell, and their probable movement routes in relation to hair-snag stations. The first30-day sample session was designed to compare two approaches in estimating the “edge”effect associated with sample grids. One used an “assessment” strip of cells around the3x3-cell core grid, while the other used an estimate of time telemetered animals spentinside and outside the core grid. Accordingly, there were 18 stations in the core grid, withan additional 32 stations in the assessment strip, for a total of 50 stations during this firstsession. Lynx hair was collected from four stations. The second 30-day sample sessionwas designed to evaluate “re-captures” subsequent to the first session on the core gridonly. Lynx hair was collected from two of 18 stations during this period. All DNAsamples collected from study animals and lynx harvested within the region this year havebeen sent to the New York genetics lab of the Wildlife Conservation Society for analysis.

9.0 Communications and Consultations

Scientific Communication.— Our plans, progress, and preliminary results have beenpresented during each of the past five years at the annual Western Forest CarnivoreCommittee meetings. This is a loose association of state, provincial and federal wildlifemanagers and researchers from western Canada and the northwestern United States withinterest in the conservation of lynx and other mid-sized carnivores. Preliminary results andconservation implications were also presented at a symposium on species and habitats atrisk during February, 1999 at Kamloops, BC. A presentation and field consultation wasprovided to the Washington Interagency Lynx Committee in August, 1998, at Penticton,BC. Consultation was provided to the Colorado Dept. of Wildlife for their lynxreintroduction program, to the USDA Forest Service for the development of their LynxConservation Strategy, the US Fish and Wildlife Service regarding lynx habitat definitionsas required under the Endangered Species Act, and the Washington Dept. of Fish andWildlife regarding their state lynx recovery plan. Advice based on preliminary results andother research findings was also provided to the following: BC Ministry of Forestsregarding lynx/bobcat habitat supply modeling; BC Ministry of Environment regardingpopulation management; four consulting companies regarding lynx habitat quality ratings,terrestrial ecosystem mapping, and linear disturbance effects; the Colorado Department ofTransportation regarding highway development within potential lynx habitat; IrisEnvironmental Systems regarding lynx and ski resort operations; Biosphere Institute of theBow Valley regarding lynx and bobcat ecology and conservation; and the NationalWildlife Federation regarding their North American wild cat conservation program. Twopapers were submitted and accepted for publication (Apps 2000; Apps et al. in press).

Public Communication.— Public communications have involved presentations in localcommunities including Golden, Invermere, Banff, Lake Louise, and throughout the EastKootenay via Shaw Cable TV. Interviews about the project have been aired on local and


regional radiostations. The project was featured on the CBC current events televisionprogram “Country Canada” and as an episode of a wildlife conservation program aired onDiscovery Channel, other educational networks, and archived with the National FilmBoard. Consultation has been provided to Parks Canada interpreters for the developmentof public education programs on lynx, wild cats, and associated conservation issues. Aproject overview document that outlines lynx research, ecology and conservation in thesouthern Canadian Rockies has been developed for public distribution.

10.0 Discussion of Vital Rate Changes during 1999/2000.

The first 3 years of this study were conducted during an assumed increase to high phase ofa snowshoe hare cycle (C. J. Krebs, University of British Columbia, pers. comm.).Despite this, hare densities in our study area have been comparable to those reported fornorthern lynx study areas during cyclic lows (Brand et al. 1976, Ward and Krebs 1985,Poole 1994, Slough and Mowat 1996). All evidence to date suggests that the harepopulation cycle is out of synch by no more than one year across most of North America(Hodges 2000a). Thus, our results are consistent with the suggestion that, althoughsouthern populations cycle in synchrony with boreal populations, hare densities in southernBritish Columbia and the northwestern United States do not approach the peaks reportedin the boreal forest and remain relatively low (Hodges 2000b).

During the first 3 years of study, lynx space use, movements, winter diet, anddemographics has resembled that of northern populations during cyclic hare lows,consistent with the apparently low hare densities in this study area (Apps 2000). The largeproportion of prey alternate to snowshoe hares occurring in the winter diet of studyanimals, including ungulate kills, is consistent with opportunistic food habits previouslyreported during hare lows (Koehler and Aubry 1994, Staples 1995, O’Donoghue et al.1988). Kitten production and survival was also significantly lower than would beexpected in northern populations during this phase of the hare cycle (Mowat et al. 2000).The apparent lack of recruitment to early winter among study animals is consistent withnorthern populations after a hare crash, when winter litter sizes were zero (Brand et al.1976, Poole 1994, Mowat et al. 1996). However, the occurrence of some kittens in thestudy area suggests that habitat quality varied among female home ranges. The highdispersal rate and low survival of juveniles would be expected in an unexploitedpopulation with limited availability and distribution of potential habitat. Annual homeranges were roughly five to ten times larger than previously reported (Koehler and Aubry1994, Poole 1994, Slough and Mowat 1996), with the exception of the year after a harecrash in the Yukon (Slough and Mowat 1996). Our extremely large 100% MCP homeranges reflect exploratory movements that were made throughout the year by all residentstudy animals. Although large 90% home ranges may reflect the low density and patchydistribution of prey, they likely also reflect the configuration of potential habitat asinfluenced by physiography in our study area. Minimum daily movements were mostlyhigher than in other studies of comparable methods (Ward and Krebs 1985, Poole 1994),


implying greater foraging effort. Ward and Krebs (1985), however, reported similar dailymovements as hares densities in their study area declined below 1.0 hares/ha.

Hare populations in North America are known to have peaked during the late 1990’s and,as has been expected, a decline was detected in several regions during 1999. In thesouthwest Yukon, where hare populations are closely tracked, the decline began duringwinter 1998/99 and populations have continued to slide during 1999/2000 (C. J. Krebs,University of British Columbia, pers. comm.). In the NWT, hare densities may have stillbeen peaking this winter, but this area has lagged behind the Yukon by one year inprevious cycles (K. Poole, Timberland Consultants Ltd., pers. comm.). Hare populationsapparently declined in Riding Mountain National Park, Manitoba, during 1999, and a crashwas expected during 2000 or 2001 (Nylen-Nemetchek 1999). A decline in hareabundance has also been detected from 1998 to 1999 in Washington State (C. Quade,Washington Dept. Nat. Res., pers. comm.), and in an Idaho population with a meandensity of 0.43 hares/ha, mark/recapture results have documented a precipitous declinebetween fall 1998 and 1999 (finite rate of change = 0.4687; A. Wirsing, University ofIdaho, unpubl. report). In our study area, pellet counts were consistent with a declinefrom 1998 to 1999 (though not statistically tested), and observations by primary fieldresearchers and one long term resident in the upper Kootenay Valley have been consistentwith a significant decline from recent years to winter 1999/2000. Collectively, preliminaryresults from our study area and other regions is consistent with evidence to date thatalthough southern hare populations do not achieve peak densities comparable to that ofthe boreal forest, densities still fluctuate two to 25 fold in synchrony with borealpopulations (Hodges 2000b). Evidence of cyclic fluctuation in hare populations is alsoconsistent with a regional trapper survey conducted for southeastern British Columbia(Apps 1996a).

The zero kitten survival past early winter that we documented among five resident femalesis similar to other lynx populations followed through the hare crash (Mowat et al. 1996).Although adult survival has been high during our first three years of study, survival waslow during year four due to natural mortality. High levels of natural mortality areexpected in unexploited lynx populations during the first year of very low hare numbersafter the crash. For example, similar to our results, annual survival rates in two largelyuntrapped northern populations were high (>0.89) prior to the crash, but low (0.09 –0.40) during the first year of very low hare densities following the crash (Poole 1994,Slough and Mowat 1996, O’Donoghue et al. 1997). In the south-central Yukon, only twopreviously resident lynx remained on the study area after the hare decline, compared toover 130 animals during the peak (transients, residents, and all age classes; Slough andMowat 1996). Hare populations always decline through winter months, but survival issignificantly lower during the decline phase of the cycle (Hodges 2000a). Survival ofresident red squirrels in Idaho was also 10 times lower during winter than during summerover the past two years (T. Steury, University of Idaho, unpubl. report.). As expectedthen, our adult mortalities this year occurred during late-winter, consistent with the timingof starvation in other studies during the hare decline phase (Mowat et al. 2000).Moreover, periods of deep, unconsolidated snow during February may have exacerbated


the energetic stress to lynx due to prey population declines. As would be expected duringor shortly after the hare decline, snowtracking data suggest that the snowshoe hare killrate this year was considerably lower than that of previous years. While lynx weresnowtracked for roughly 11 km on average between successful hare kills during the firstthree years, an average of 28 km were tracked between hare kills during winter1999/2000. This would indicate of a considerable increase in foraging effort and energeticoutput per success.

In the boreal forest, the numeric response in lynx populations typically lags the onset ofthe hare crash by one year (Mowat et al. 2000). In these northern regions, it is expectedthat hare densities can decline considerably from their peak before eliciting a significantdemographic response in the lynx population, but as they fall below a hypothetical haredensity threshold, dramatic effects are seen in lynx population vital rates (Ibid.). Becausepeak hare densities in this and perhaps other southern lynx populations are likely muchcloser to this threshold, it can be expected that effects to lynx reproduction and survivalwill be manifested much closer to the onset of the hare decline. As a result, considerablenatural mortality or dispersal among residents may be clustered within the first year of thedecline.

In contrast to northern populations, our results suggest that alternate prey likely play animportant role in the southern Canadian Rockies even during the hare population peak.Red squirrels, the most important alternate prey, are also subject to populationfluctuations with conifer cone productivity (Klenner and Krebs 1991). Red squirrelpopulations in the larger region may have shown a marked decline in the past two years,presumably due to two consecutive years of cone crop failure (T. Steury, University ofIdaho, unpubl. report; T. Thier, Montana Fish Wildlife and Parks, pers. comm.).Consistent with this, we documented a notable change in squirrel kill success rate this yearrelative to previous years. Over 283 km of snowtracking conducted during winter1999/2000, three squirrel kills were documented (1 per 98 km) as compared to 46 in thefirst three years (1 per 20 km). Similarly, spruce grouse populations also cycle (Boutin etal. 1995) and have been scarce in the past two years relative to the first two years of study.

11.0 Local and Regional Conservation Recommendations

The following management recommendations are based on our preliminary results inconjunction with other completed and ongoing research on lynx community ecology.Forest management guidelines are currently being developed. These will specifically relateto provincial ecosystem and forest classification mapping standards, nomenclature, andoperational forest development planning under the Forest Practices Code.

• Close the lynx harvest season in southeastern British Columbia and southwesternAlberta. Although this is standard practice throughout lynx range during the lowphase of the hare cycle, it is assumed that human mortality is compensatory to naturalmortality during the first two years of the crash (Mowat et al. 2000). However, due to


much lower peak hare densities, natural lynx mortality may be much more precipitousin the southern Canadian Rockies, and population harvest may only be compensatoryduring the first year of the hare decline. After this, lynx that survive the crash will stillbe vulnerable, and human mortality may be additive to natural mortality, threateningthe recovery of the regional population.

• Consider the distribution of current and potential regional lynx habitat and ensurethat protective measures are in place in the most productive habitat nodes. Regionallynx conservation depends on the existence of unharvested refugia of the mostproductive habitat. Implications of localized overharvest may be severe in regionswhere the most productive habitat nodes are patchy and populations occur at lowdensities. Regional modeling to identify these landscapes is currently being conductedas part of this project.

• Ensure that mitigative crossing structures associated with major human features suchas the twinned Trans-Canada Highway (TCH), are of maximum width (i.e., > 500 m)and are associated with a continuation of suitable habitat with maximum cover, andnatural terrain. It is not known whether lynx will use existing TCH crossingstructures (A. Clevenger, Banff National Park, pers. comm.). Because they areparticularly sensitive to interference competition with other terrestrial predators, lynxmay not use travel routes where movement options are constrained and where thelikelihood of interspecific interaction is increased. Extensive understory cover and/orescape habitat such as “jackstrawed” blowdown and tall, large diameter trees at ordirectly proximal to crossing structures may encourage their use. Although majorhuman and natural features may normally represent home range boundaries, lynx mayattempt to cross the TCH during the breeding period and when prey densities declineand resident animals become partially nomadic. Broad-scale movement options tofacilitate successful dispersal may be especially important for population recovery asprey densities increase. Our data can be used to identify actual and model probablepreferred crossing zones.

• Conduct comprehensive vegetation, terrain, and human use inventories withinnational parks that are that are consistent with provincial standards and initiatives.This should begin with 1:20,000 forest overstory and land cover mapping. It isessential that these data are available for regional habitat supply modeling and analysesand to direct habitat management within and adjacent to national parks.

• Avoid promoting increased and unnatural competition with other terrestrialpredators. In the southern Canadian Rockies, subalpine landscapes may support lynxlargely because winter snow conditions normally preclude canid competitors. Snowcompaction due to extensive and formalized road, snowmobile, ski, and snowshoenetworks may facilitate year-round residency by competitors, particularly coyotes.This may influence lynx through interference competition and displacement fromeffective habitat, but will also have a major impact on prey densities, intensifying harepopulation declines through winter months.


• Avoid creating fire guards of simplified forest structure around developments. Suchbuffers will largely negate habitat value for many small mammal and avian species,rendering them ineffective as foraging habitat for lynx and other predators.Consequences to lynx depend on the current and potential ability of the largerlandscape to support resident animals.

• Maintain future habitat management options that employ prescribed burning. Overtime, a prescribed burning program is necessary to achieve an area and distribution ofseral classes that reflect a natural disturbance regime, especially within protected areas.However, even in landscapes where prescribed burning is not currently beingconsidered, it is important to maintain future options to use fire as a habitatmanagement tool. This should be an explicit consideration in all environmental impactassessments for recreational and resort development.

• Minimize high impact human development and recreational activities within andproximal to potential lynx denning habitat. Lynx are expected to be sensitive todisturbance by humans and other predators in their selection of natal den sites. Habitatmodeling conducted through this project can identify potential denning areas, whichwill assist in resource planning and access management. Although inherently lessobtrusive than motorized traffic, human foot or bicycle traffic may still render potentialdenning habitat ineffective, especially if accompanied by off-leash domestic dogs.


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