Rocky Mountain Bighorn Sheep ( Ovis Canadensis Canadensis (Shaw, 1804)) Where 4 Large

Canadian Journal of Zoology

Bighorn sheep winter habitat selection and seasonal movements in an area of active coal mining

Journal: Canadian Journal of Zoology

Manuscript ID cjz-2016-0069.R1

Manuscript Type: Article

Date Submitted by the Author: 29-Jun-2016

Complete List of Authors: Poole, Kim; Aurora Wildlife Research Serrouya, Robert; University of Alberta, Department of Biological Sciences Teske, Irene; British Columbia Ministry of Forests, Lands and Natural Resource OperationsDraft Podrasky, Kevin; Teck Coal Limited

bighorn sheep, British Columbia, coal mining, habitat selection, Ovis Keyword: canadensis, reclamation, winter range

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13 July 2016

Kim G. Poole Aurora Wildlife Research 1918 Shannon Point Road, Nelson, BC, V1L 6K1, Canada Tele. (250) 825-4063 e-mail: [email protected]

Bighorn sheep winter habitat selection and seasonal movements in an area of active coal mining Kim G. Poole, Rob Serrouya, Irene E. Teske, and Kevin Podrasky

K.G. Poole , Aurora Wildlife Research, 1918 Shannon Point Road, Nelson, BC V1L 6K1, Canada [email protected] R. Serrouya , Department of Biological Sciences,Draft University of Alberta, Edmonton, AB T6G 2E9, Canada [email protected] I. Teske , Ministry of Forests, Lands and Natural Resource Operations, Fish, Wildlife and Habitat Section, 205 Industrial Road G., Cranbrook, BC V1C 7G5, Canada [email protected] K. Podrasky , Teck Coal Limited, P.O. Box 2003, Sparwood, BC V0B 2G0, Canada [email protected]

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Bighorn sheep winter habitat selection and seasonal movements in an area of active coal mining

K.G. Poole, R. Serrouya, I.E. Teske, and K. Podrasky

Abstract: Winter is an important period for most mountain ungulates due to limited availability

of preferred forage and costs associated with travel through deep snow. We examined winter habitat

selection by Rocky Mountain bighorn sheep ( Ovis canadensis canadensis (Shaw, 1804)) where 4 large,

open-pit coal mines are in operation. Sheep in this area generally winter at high elevation on

windswept, south-facing native grasslands. We used GPS collars and Resource Selection Function

analysis to examine movements and habitat selection. Most (79%) of the sheep were migratory and

fidelity to winter ranges was high (88%). Sheep showed low use (~10–20%) of mine areas between

November and April, followed by increased use peaking at 60–65% in September-October. Wintering sheep were positively associated with highDraft elevations, closeness to escape terrain, and warmer aspects. High-elevation, native grasslands were the highest ranked cover class. Most sheep that used

mine areas during winter used reclaimed habitats, primarily reclaimed spoils and pits. Primary winter

ranges comprised 4.3% of merged sheep range, emphasizing the limited amount of occupied winter

ranges within the landscape. Disturbance to native winter range resulting from development should be

minimized or be conducted in a manner that effectively manages and/or mitigates the impacts.

Key words: bighorn sheep, British Columbia, coal mining, habitat selection, Ovis canadensis ,

reclamation, winter range

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Introduction

Most mountain ungulates exhibit complex spatial and temporal use patterns, in large part

because of the effects of winter and seasonality on forage production, availability and quality. Snow

and weather influence mountain ungulates by increasing costs of thermoregulation, energetic costs of

locomotion and by the burial of preferred forage (Burles and Hoefs 1984; Parker et al. 1984; Daily and

Hobbs 1989). Ungulates that typically live at high elevations cope with winter and deep snow cover by

using a variety of strategies. Some ungulates alter vegetation use (alpine chamois ( Rupicapra rupicapra

(L., 1758)) and Himalayan tahr ( Hemitragus jemlahicus (Smith, 1826)): Forsyth 2000), including

movement to areas of heavier cover (mountain caribou ( Rangifer tarandus (Gmelin, 1788)): Apps et al.

2001) or movement to open, high-elevation habitats in shallow snow zones (bighorn sheep (Ovis canadensis (Shaw, 1804)): Demarchi et al. Draft2000; mountain goats ( Oreamnos americanus (Blainville, 1816)): Poole et al. 2009; Abruzzo chamois ( R. pyrenaica ornate (Bonaparte, 1845)): Lovari and

Cosentino 1986). When snow depths increase, some may move to lower elevations (chamois: Lovari

and Cosentino 1986) or to higher elevations later in the winter to areas of deeper snow for access to

arboreal forage (mountain caribou: Apps et al. 2001). Lastly, some species move to escape terrain and

moderate slopes during winter (Asiatic ibex ( Capra ibex sibrica (Pallas, 1776)) and blue sheep ( Pseudois

nayaur (Hodgson, 1833)): Namgail 2006).

Habitat selection by ungulates is thought to be a trade-off between nutritional and anti-

predation constraints (Festa-Bianchet 1988 a; Hebblewhite and Merrill 2011). In western North

America, seasonal habitat selection by bighorn sheep varies among populations likely in response to

both predation risk and snow depth (Shackleton et al. 1999; Demarchi et al. 2000; Krausman and

Bowyer 2003). Proximity to escape terrain is an important aspect of most sheep habitat selection (e.g.,

Tilton and Willard 1982; DeCesare and Pletscher 2006; Bleich et al. 2009).

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The east side of the Elk Valley in southeastern British Columbia (BC) is home to a population of

Rocky Mountain bighorn sheep ( O. c. canadensis ) of provincial significance; the Ewin Ridge sheep range

has been considered the most important bighorn sheep winter range in BC (Demarchi 1968). Sheep

typically winter at lower elevations to avoid deep snows that accumulate in most parts of their range

(e.g., Tilton and Willard 1982), however, sheep in the Elk Valley are unique as they generally winter at

high elevation on windswept, south-facing native grasslands. Lower elevation habitat in this area is

primarily forested with heavy snow accumulation.

The Elk Valley supports important industrial activities, primarily coal mining and forestry. Four

large, open-pit coal mines occur within the east side of the Elk Valley (Fig. 1), and sheep use areas of

some mine sites to varying extent year-round (K. Podrasky, Teck Coal, unpubl. data). Coal mines can create bighorn sheep habitat where none Draftexisted previously, as has occurred at mines in the Alberta foothills (MacCallum 1991; MacCallum and Geist 1992). At these Alberta mines, bighorn sheep from

the adjacent occupied alpine habitat responded to the mine reclamation by occupying the new

habitats, expanding their range, and exhibiting rapid population growth. Bighorn sheep also can use

urban environments, which can provide higher forage quality (Rubin et al. 2002; Dibb et al. 2008).

While human disturbance (e.g., noise, activity) from mines do not appear to deter use by sheep

per se (Oehler et al. 2005; Jansen et al. 2006, 2009; Bleich et al. 2009), expansion of coal mining may

cause direct loss of winter range. Winter range is important habitat for bighorn sheep (Demarchi et al.

2000), and has been suggested as the single most important factor limiting sheep populations in the

Elk Valley area (Schuerholz 1984). The late winter/early spring (green-up) period is particularly

important, as delayed green-up may cause starvation in sheep populations (Burles and Hoefs 1984).

Identifying high value habitats can help assess and manage the potential impacts of industrial activities.

There is concern about management of mountain sheep and the potential impacts that habitat

alteration might have on those ungulates (Bleich et al. 2009). Incomplete knowledge about bighorn

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sheep ecology, including important winter use areas and habitat selection, can be an obstacle to

coordinating sheep management activities. We hypothesized that ( i) winter habitat selection would

differ between sheep wintering on native grasslands and those using active coal mine areas because of

limitations to how mine areas can duplicate establishment of native ranges, and ( ii ) greater winter

severity would result in smaller seasonal range size and decreased sheep survival. The information

collected in this study could be applied to other regions where mountain ungulate range and resource

development overlap.

Materials and methods

Study area

The 830 km 2 study area is located in the Rocky Mountains of southeastern BC, and focused on bighorn sheep populations wintering east Draftof the Elk River and stretching up to the Continental Divide (Fig. 1). Biogeoclimatic zones within the study area grade from Montane Spruce dry cool (MSdk) in

valley bottoms, Engelmann Spruce–Subalpine Fir dry cool (ESSFdk, including subalpine parkland) at

higher elevations, to small amounts of Alpine Tundra (AT) on ridge and mountain tops (Braumandl and

Curran 1992). Krummholz subalpine fir ( Abies lasiocarpa (Hook.) Nutt.) with scattered whitebark pine

(Pinus albicaulis (Engelm.) Rydb.) dominate the treeline, whereas closed to open stands of Engelmann

spruce (Picea engelmannii Parry), subalpine fir, lodgepole pine ( Pinus contorta Dougl. ex Loud.), and

aspen ( Populus tremuloides Michx) are found at lower elevations. Cliffs of various sizes, and open scree

slopes and meadows are located on most ridgelines and mountain tops. Sheep winter ranges are

characterized by herb-rich, tall grass meadows that occur at higher elevation (most between 2000–

2300 m elevation, but ranging from 1800–2500 m), and which retain low snow depths because of their

south and southwest aspects and strong westerly winds (Smyth 2014). Average mean daily

temperatures at Sparwood, British Columbia, located at 1140 m elevation in the Elk River valley near

the southern end of the study area, were –6.8 °C and 15.4 °C for January and July, respectively, and

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6 precipitation averaged 603 mm annually (Environment Canada 2011). Sparwood receives 248 cm of snowfall annually.

Snow pillow data (snow water equivalent) were obtained for Morrissey Ridge, located at 1966 m, 30 km southwest of the study area (http://bcrfc.env.gov.bc.ca/data/asp/realtime/index.htm). Snow depths varied among winters during the study, with winter 2009–10 characterized by 10–35% lower snow depths than average, and winter 2010–11 with 10–80% higher snow depths coupled with a late melt (as a result of a cold and wet spring). Winter 2010–11 had the highest snow depths since 2001–

02, and was 94% more severe (Baccante and Woods 2008) than winter 2009–10.

Potential predators of bighorn sheep in the study area include grizzly bear ( Ursus arctos (L.,

1758)), black bear ( U. americanus (Pallas, 1780)), cougar ( Puma concolor (L., 1771)), wolverine ( Gulo gulo (L., 1758)), wolf ( Canis lupus (L., 1758)),Draft coyote ( C. latrans (Say, 1823)), and golden eagle ( Aquila chrysaetos (L., 1758)). The area also supports a high density and diversity of other ungulates, including

large numbers of elk ( Cervus elaphus (L., 1758)) and mule deer ( Odocoileus hemionus (Rafinesque,

1817)), scattered mountain goat ( Oreamnos americanus ) populations, and low numbers of moose

(Alces alces (L., 1758)) and white-tailed deer ( O. virginianus (Zimmermann, 1780)). Mule deer and elk in

particular were observed on the same winter ranges used by sheep.

The primary resource development activities within the study area are open pit coal mining

and forestry. Approximately 17% (137 km 2) of the study area was composed of mine-related infrastructure, including pits, spoils, reclaimed areas, and some logged areas. These 4 mining operations have impacted terrain from the valley bottoms to the mountain tops (Fig. 1). Back country roads to varying degrees of functionality occur within the study area, but vehicle traffic is generally light. Forestry activities are limited to lower to mid-elevations off main access roads.

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Bighorn sheep capture and collaring

We deployed GPS collars on bighorn sheep within the study area in late February 2009,

spreading capture effort through each winter range and between sexes, with a greater emphasis on

ranges with higher sheep numbers. Capture and handling protocol was conducted under BC

Environment permit CB09-51173. Sheep were captured primarily by helicopter netgunning (Barrett et

al. 1982), and a radiocollar was attached (GPS collar model G2110B, Advanced Telemetry Systems,

Isanta, Minnesota, USA) set to a 10-hour fix rate. Timed collar drop-off was set for mid-May 2011,

covering 2 full winter seasons. We estimated age using annuli on the horns and tooth eruption.

We located collared sheep monthly using a fixed-wing aircraft to establish locations and to

estimate survival. When collared sheep were not detected we occasionally searched up to 75 km from the study area, and this searching was repeatedDraft at the end of the study. Collars detected on mortality mode were investigated as soon as possible, and the collar data downloaded. Collars were then

refurbished and redeployed at the earliest opportunity using ground and helicopter-based techniques.

Ground-based capture involved darting with immobilizing drugs and processing similarly as for

helicopter netting. We used 1.6–1.8 cc of a premixed mixture of detomidine hydrochloride

(Dormosedan) and tiletamine hydrochloride and zolazepam (Telazol) (5 ml of detomidine at 10 mg/ml

concentration in each vial of Telazol), with 6 cc of atipamezole hydrochloride (Antisedan) provided for

reversal. Prior to analysis we removed locations collected within 1 week of capture to reduce

movements associated with capture-related disturbance and for the animals to acclimate to the

transmitter (White and Garrott 1990; Morellet et al. 2009). We visually removed <10 locations

considered errant outliers. Mean GPS location error was assumed to be 11 m, with 50% and 95% of

locations within 6 and 31 m accuracy, respectively (D’Eon et al. 2002). Because of high fix success and

location quality (see Results), we did not correct for habitat-induced GPS bias (Hebblewhite et al.

2007).

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Fate and survival rates

To examine survival rates of sheep we used the staggered entry (monthly interval) Kaplan-

Meier survival estimator (Pollock et al. 1989), calculated in program Ecological Methodology version

6.1. (Kenney and Krebs 2002). Sheep that experienced suspected capture-related mortality were removed from the survival analysis. Since we had no indication that predation significantly affected collar transmission and long-distance search flights around the study area did not detect dispersal, and it appeared that rams were hard enough on the collars to break the VHF crystals (see Results), we assumed sheep were alive until censored when contact was lost. Survival rates were calculated separately for ewes and rams, and by year (year beginning mid-May, just prior to lambing), and were presented as finite survival rates for the period of collaring and converted to annual survival rates. We tested for differences between sex and yearDraft using log-rank tests. Seasonal ranges

We defined 4 broad seasons based on population average movement rates and changes in elevation and spatial distribution (unpubl. data), and assigned winter (1 December 2009 or 15

December 2010 to 30 April; generally low movement rates and stable use of elevation); spring (1 May

to 22 June; includes the bulk of lambing; use of low elevation [green-up]; increasing movement rates);

summer (23 June to 24 August; highest elevation use and movement rates); and fall (25 August to 30

November 2009 or 14 December 2010; includes the rut in late November; variable use of elevation,

declining movement rates [except for rams during the rut]). Where dates of individual arrival onto or

departure from winter range did not fit these patterns, we modified season dates to accommodate

these individual spatial shifts. Slightly later onset of winter in 2010–11 appeared to result in delayed

movement onto winter ranges compared with December 2009.

To determine individual home ranges we calculated 95% fixed kernels (Worton 1989; Seaman

and Powell 1996) using the Home Range Extension (Rodgers and Carr 1998) for ArcView, with unit

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variance standardization, a user-defined smoothing factor of 0.70, and raster resolution set to 120. We

defined a fixed smoothing parameter because least-square cross-validation tends to produce under-

smoothed data (Kie et al. 2010). We used a 90% fixed kernel to determine individual seasonal ranges.

We these levels of kernels as they best captured the extent and spatial pattern of annual and seasonal

sheep range use. We compared range sizes using t-tests on log-transformed data to correct for

heteroscedasticity to examine space use patterns between sexes and seasons in years of differing

severity.

Movements and winter range fidelity

We assumed migration occurred if winter and summer ranges did not overlap (Brown 1992;

Nicholson et al. 1997; Mysterud 1999) or if winter range encompassed <10% of summer range; we termed these sheep ‘migratory’, and termedDraft sheep that did not show such patterns as ‘non-migratory’. Migration distance was defined as the horizontal distance between seasonal range centers of activity

(centroids; Hayne 1949; Mysterud 1999), with the centroid calculated as the mean UTM coordinates of

the locations for a sheep in each season using the Animal Movement Extension for ArcView (Hooge

and Eichenlaub 2000). Migration distance between seasonal centroids was calculated between

summer and winter for both years. Comparison of distances between summer and winter centroids

was conducted for both years and between sexes using 2-tailed t-tests on log-transformed data.

We examined winter range fidelity by determining whether sheep remained on the same

winter range during winters 2008–09 (as indicated by late February and March 2009), 2009–10, and

2010–11. We included sheep that were monitored but the collars were not recovered. We also

summarized distances between consecutive winter use centroids.

We determined the proportion of sheep locations on mine areas, which included all mine

infrastructure, active mining areas, and reclaimed property; data were summed weekly and plotted by

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sex to examine seasonal use of mine areas. Individual variation also was examined by summing

locations for each collared sheep.

Resource selection

Habitat data and use

To examine habitat use and selection we obtained digital data for habitat variables likely of

importance to bighorn sheep during winter based on the literature (e.g., Festa-Bianchet 1988 a;

Sweanor et al. 1996; MacCallum 1991; Dicus 2002; DeCesare and Pletscher 2006): 1 m resolution LiDAR data flown in summer 2011 (D. Vasiga, Teck Coal, unpubl. data); land cover classification (Earth

Observation for Sustainable Development of Forests – EOSD; Wulder et al. 2008); and mine areas (D.

Vasiga, Teck Coal, unpubl. data). The LiDAR data were degraded to 20 m resolution from which we derived terrain variables of interest (TableDraft 1). All databases were rasterized to ArcGIS 9.3 (Environmental Systems Research Institute, Redlands California, USA) grid format at 20 m resolution.

We calculated terrain ruggedness (RUGGED) with the vector ruggedness measure (Sappington et al. 2005), using an ArcGIS script to calculate the 3-dimensional dispersion of vectors within the landscape at a 7 x 7 grid scale (script available online from the Environmental Systems Research

Institute ArcScripts website: www.esri.com/arcscripts). We used digital terrain modelling to calculate solar duration (SOLAR), a measure of solar radiation, which is the number of hours a pixel sees the sun

per day based on latitude (sun angle) and the shading effects of nearby topography (Kumar et al.

1997). We calculated means for solar duration for 11 January to 31 March to match work completed

elsewhere in the area (e.g., Poole et al. 2009). Although definitions of escape terrain for bighorn sheep

vary (e.g., Tilton and Willard 1982; Sweanor et al. 1996; McKinney at al. 2003; DeCesare and Pletscher

2006); MacCallum (1991) observed that bighorn sheep in the Alberta Rocky Mountains used steep

terrain between 35 ° and 45 ° for escape, travel, bedding, and lambing. We subjectively used ≥37 ° (75%)

slope to delineate escape terrain, very steep areas that are often cliffs that predators do not use (Arjo

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and Pletscher 2004). From each grid cell we calculated the distance to the nearest escape terrain

(ESCPDST).

EOSD data are derived from year 2000 classified Landsat-7 Enhanced Thematic Mapper Plus

(ETM+) images with a pixel resolution of 25 m (Wulder et al. 2008). The original EOSD land cover data

within BC were modified to remove shadows and small raster patches. Additional corrections and

additions were made through visual inspection of ortho-imagery taken in summer 2010 and GIS layer

files to update the mine areas. The original land cover classes in the study area were collapsed into 9

mutually exclusive classes (Table 1) to remove small classes and combine similar classes. Original EOSD

data were used for locations in Alberta (2.6% of the sheep locations).

We used the digital data to characterize sheep habitat use within each full winter of monitoring within corresponding 90% fixedDraft kernel winter ranges and 95% fixed kernel home ranges. Within each individual seasonal and home range we placed 1000 random points, and calculated

descriptive characteristics for each terrain and cover class variable. We determined the mean and 90 th

percentile range (i.e., lower 5% and upper 95% percentile values) for each terrain and cover class

variable for each sheep based on these point data, and then calculated the mean of individual means

and lower and upper limits to provide a sample mean and lower and upper limit for each variable. We

used the 90 th percentile as a measure of general use to avoid outliers and provide a more meaningful

measure of the range of habitat use. We compared terrain and cover class variables among winter use

and random home ranges using an analysis of variance (ANOVA) on log-transformed data to correct for

heteroscedasticity; seasonal means were compared using Duncan’s multiple-range test.

We also calculated mean and 90 th percentile range values for terrain variables between sheep

wintering on mine areas and native ranges, and compared means using t-tests on log-transformed

data. Within mine areas, we summarized cover type used by sheep: reclaimed land, spoil (waste rock

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dump), pit, highwall (the face of exposed overburden and coal), footwall (the interface between the

base of a coal seam and the surrounding rock formation) and roads.

Habitat selection

We defined selection as the process in which an animal chooses a resource, and used the

terms “selected” (use greater than availability) and “avoided” (use less than availability) to describe

selection for or against a resource, respectively (Manly et al. 2002:3). We considered that selection of

home range within their geographic range (2 nd order selection; Johnson 1980) would not be enlightening because these sheep select home ranges in areas with moderate to high elevation and generally open, rugged terrain within the landscape, and most valley bottoms and expanses of forest on non-steep slopes are not truly “available” to bighorn sheep because of behavioural constraints (heavily forested and/or distant from escapeDraft terrain)(summarized in Shackleton et al. 1999; Krausman and Bowyer 2003). We examined winter habitat selection at 2 scales: 1) the home range scale (winter habitat use compared to random points within the home range), and 2) the winter range scale (winter habitat use compared with random points within the winter range) (3 rd order selection; Johnson 1980).

Selection for resources at the winter range scale is important for predicting and understanding

seasonal animal occurrences, and it is also at this scale (3 rd order; stand-level) that land management

decisions are frequently made (Nielsen et al. 2002). Both resource use and availability were identified

by individual sheep (type III study design; Manly et al. 2002). Comparisons between winters were

conducted to examine the effects of winter severity on sheep habitat selection. Selection at the winter range scale was examined by sex (Bleich et al. 1997).

We considered the individual collared bighorn sheep as the sample unit (Johnson 1980; White and Garrott 1990; Aebischer et al. 1993). No 2 collared sheep shared the same herd year-round, therefore minimizing pseudoreplication (Millspaugh et al. 1998). Sample sizes of locations ranged from

275–325 per winter for most animals (range 59–358).

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We developed an a priori set of candidate models based on the literature and field

observations to explain winter habitat selection at the 2 scales by bighorn sheep. We developed 3

model themes consistent with selection for topography and security (elevation, slope, terrain

ruggedness, distance to escape terrain, and mine areas); Tilton and Willard 1982; MacCallum 1991;

McKinney at al. 2003; Sappington et al. 2005), habitat and forage availability (cover type, especially

open and grassland habitats, and solar duration; Festa-Bianchet 1988 a), and thermoregulation (solar

duration, conifer cover; Cook et al. 1998).We also examined several models that combined variables

from themes. A quadratic term for elevation was considered to model selection for intermediate

elevation (Gross et al. 2002). We included interaction terms with year (winter) with several variables

because of likely differences in selection between years due to observed differences in snow depth. We evaluated multicollinearity among variablesDraft (lmer package in R; R Development Core Team 2008) with sheep as a random effect, which produces an output of correlations among fixed effects. We

based the final coefficients on bootstrapped values (n = 1000) of coefficients from individual sheep

(see below). Between highly correlated pairs of variables (r > |0.7|; Tabachnick and Fidell 1996) we

chose the one with the greatest biological meaning according to the literature.

We examined sheep habitat selection using multivariate logistic regression. Rather than using

mixed effects models where individual animals are specified as a random effect (Gillies et al. 2006) we

created a separate resource selection function for each individual (Fieberg et al. 2010). With mixed

effects models the large number of locations per individual can produce results where every variable is

significant, revealing little about the strength among variables (preliminary analyses). To infer selection

we used the RSF equation w(x) = exp(β1x1 + β2x2 … βnxn), where β 1 . . . β n are coefficients and w(x)

represents the relative probability of occurrence. RSF values range from 1 to infinity (Johnson et al.

2006).

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When models included the set of cover classes, we set the most abundant cover class (conifer- leading stands) as the reference category. Following suggestions by Anderson and Burnham (2002), we assessed the strength of competing models using Akaike’s Information Criteria (AIC) values (Anderson et al. 2000). We calculated standardized coefficients (Menard 2004) that reflect the relative magnitude of each variable for each sheep. The modelling process was repeated for each sheep, but to obtain inference for the population we bootstrapped parameter estimates, weighted by the inverse of their unconditional (cross model weighted) standard errors, and presented mean values with 95% CIs using the percentile method. Inconclusive statistical inference is expected from covariates with confidence intervals that overlap 0. At the home range scale, to evaluate the predictive accuracy of models we calculated the receiver operating characteristic (ROC) curve (Boyce et al. 2002) for each sheep and determined the mean ROC and correspondingDraft 95% CIs. Models with a ROC value >0.9 were considered to reliably discriminate used from random points. Although ROC values can be biased due to the use- available design used here (Boyce et al. 2002), these biases would not compromise comparisons among models. Furthermore, for the winter range scale we combined 2 independent sources of data to validate models. We used annual mine total count aerial inventories within the BC portion of the study area between 2006 and 2011 (a recent period of complete coverage and uniform survey methods; L. Amos, Teck Coal, unpubl. data), covering 352 groups and totalling 2588 individuals; and

Forests, Lands and Natural Resource Operations (FLNRO) surveys of the Elk Valley East population conducted in 2008, 2010, and 2011, totalling 227 groups and 1320 sheep (I. Teske, BC FLNR, unpubl. data). Model validation was done by calculating the area-adjusted frequency of 10 equal RSF bins and plotting the predicted RSF values vs. observed data corresponding to the same bins (Boyce et al. 2002).

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Results

Captures

We captured and fitted GPS collars on 39 bighorn sheep (19 ewes and 20 rams) in February

2009, and subsequently re-deployed 11 collars (on 3 ewes and 8 rams) between May 2009 and October

2010. Re-deployment efforts were directed at rams as we became aware that more ram collars were

dropping off or lost. Five rams could not be located by the end of the project and collar failure was

suspected. Three collars that released remotely in active mining areas were deemed too dangerous to

collect and were never recovered. We ultimately obtained location data from 41 sheep (19 ewes, 22

rams) which provided an average of 19.1 months of location data (± 8.5; range 0.4–26.6 months). The

final data set contained 53 682 locations after censored locations were removed. Location success was 95.1%, while 85.0% of all locations were 3-dimensioDraftnal fixes. No collared sheep were detected outside of the study area or further than 1–2 km into Alberta, based on long-distance scanning flights. Based

on blood progesterone, 18 of 19 ewes (95%) 3+ years of age captured in February 2009 were pregnant

(unpublished data).

Fate and survival

Two mortalities occurred within 2 days of capture and handling, and were censored from

further analysis. Eleven sheep died during the study, 7 ewes and 4 rams. Six mortalities (5 ewes, 1 ram)

were attributed to unknown natural causes, 1 ram to grizzly bear mortality, 1 ewe to wolf mortality,

and 2 rams to starvation. Only 1 known human-related mortality occurred, related to a vehicle strike

on a mine access road. Six of the mortalities occurred on mine areas (1 human-related, 5 natural

causes). All but 1 mortality occurred between December and April.

We used 476 ewe-months and 443 ram-months of data for survival analysis. Average annual

survival rates were 0.83 (0.72–0.92) and 0.87 (0.77–0.97) for ewes and rams, respectively, and did not

differ between sexes ( X2 = 0.43, 1 df, P > 0.05). Three mortalities occurred between mid-May 2009 and

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mid-May 2010 (survival rate 0.93 ± 0.85–0.99) and 7 mortalities occurred between mid-May 2010 and

mid-May 2011 (0.78 ± 0.66–0.90), but the difference in annual survival rates was not significant ( X2 =

2.3, 1 df, P > 0.05).

Spatial ecology

Size of summer ranges did not differ between sexes for summers 2009 or 2010, or for winters

2009–10 and 2010–11 (90% ranges, t < 1.37, 27–28 df, P > 0.18). Sizes of summer ranges did not differ

between years (90% ranges, t = 0.09, 58 df, P = 0.93), but winter ranges were roughly one third the size

during the high snow winter 2010–11 compared with the previous year (90% ranges, t = 2.59, 57 df, P =

0.01; Table 2).

Most (79%) of the sheep monitored for a summer to winter session were migratory (both years considered; n = 53 sheep-years). A lower proportionDraft of ewes were migratory (67%) when compared to rams (96%). All of the non-migratory sheep resided on or near the northern 2 adjacent mines ( n = 10 sheep-years). Non-migratory sheep (10.5 ± 5.69 km 2; n = 6) had smaller individual ranges than migratory sheep (45.8 ± 38.68 km 2; n = 24; t = 2.05, 28 df, P < 0.001). For sheep that were migratory,

distance between summer and winter centres of activity (centroids) did not differ between years for

either ewes (4.1 ± 3.28 km vs. 4.9 ± 2.35 km; t = 0.81, 18 df, P = 0.43) or rams (7.6 ± 2.94 km vs. 7.1 ±

3.16 km; t = 0.25, 19 df, P = 0.80). Mean distance between seasonal centroids was greater for rams (7.7

± 2.70 km) compared with ewes (5.8 ± 2.47 km; t = 2.25, 39 df, P = 0.03). Fidelity to individual winter

ranges among years was 88% (ewes 88%; rams 88%) on a sequential-year basis (n = 60). Of the 7 cases

that did not show fidelity, only 4 (7% of overall pairings) did not involve adjacent winter ranges.

The proportion of collared sheep on mine areas ranged from about 10–20% between

November-December and April to peak at about 60–65% in September-early October (Fig. 2). Seasonal

patterns of use were generally consistent between sexes and years. During the 3 rd week of May 2009

and 2010 (roughly the peak of lambing), ewe locations were positioned on mine areas 50% of the time,

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suggesting that some ewes were lambing on mine areas. Differences among individuals and mine areas

were apparent. Three sheep (2 ewes, 1 ram) spent 82–94% of their time on one mine area, in essence

almost year-round residency, while 3 other collared sheep (2 ewes, 1 ram) made almost no use of mine

areas, occupying ranges east of the mines.

Resource use and selection

Winter habitat use

Compared with home ranges, winter sheep locations were located at higher elevations, on

warmer and steeper slopes, in more rugged terrain, and closer to escape terrain (Fig. 3, Table S1).

Among main cover classes, winter sheep locations were more often found in high-elevation grasslands

(winter 2009–10 only) and barren land, and less often found in coniferous forests, rock-rubble, and mine areas compared with home ranges (Fig.Draft 4, Table S1). Most (90%) winter sheep locations occurred within 90–95 m of escape terrain, as compared within 280 m within the home range. Overall sheep use

of resources did not differ markedly between winters. There were no differences in mean topographic

values between ewes and rams within either winter (t-test, t < 1.95, P > 0.06), and limited differences

among land cover variables (Table S1).

Compared with native areas, sheep wintering on mine areas were at lower elevation, on less

steep slopes and less rugged slopes, on cooler aspects, and further from escape terrain (Table 3).

Sheep that used mine areas in winter primarily used reclaimed habitats and spoils, and used of pits and

highwalls less.

Winter habitat selection

Slope and distance to escape terrain were correlated ( rs = –0.81), and we selected distance to

escape terrain as the most causative variable for modelling ( cf Sweanor et al. 1996, Dicus 2002,

DeCesare and Pletscher 2006). Barren land was correlated with several land cover variables (rs = 0.75–

0.79). Since barren land and high-elevation grasslands often occurred simultaneously on several of the

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sheep winter ranges, we combined these 2 variables for modelling. Ewes and rams demonstrated

similar selection of variables, with the exception of terrain ruggedness, where rams showed negative

to neutral selection and ewes showed positive selection, and selection was reversed for logged

habitats (Figs. 3, 4). However, the importance of these variables was weak, and thus would have little

influence on modelled output. We therefore combined sexes for analyses.

At the home range scale, variables representing topographic/security and habitat/forage

themes were most selected during winter (Table 4). The final model had excellent predictive accuracy

(ROC median = 0.94, 95% CI 0.93–0.95). Elevation, distance to escape terrain, solar duration and high- elevation grasslands/barren lands were the strongest variables, where the majority of sheep were showing consistent positive (elevation, solar, and grasslands) or negative selection (distance to escape terrain). The negative quadratic term for elevationDraft indicated selection for mid-elevations within the home range. Selection for terrain ruggedness was positive but showed inconsistent selection among sheep. Among land cover variables, high-elevation grasslands/barren lands were highest ranked and had the greatest positive selection by sheep. Low-mid-elevation grasslands also were selected by most sheep. Mine areas were equally selected and avoided by individual sheep. Except for selection for closer to escape terrain during the high snow year, all terms with year interactions had confidence intervals that overlapped 0, and all had low relative importance. Therefore, to simplify interpretation

(Murtaugh 2007); we removed year interaction terms.

At the winter range scale, sheep selected higher elevations and warmer aspects (higher solar duration), and selected areas closer to escape terrain (Table 5). The negative quadratic term for elevation indicated selection for mid-elevations within the winter range with a peak selection at about

2150 m. Sheep showed highest selection for areas in and immediately adjacent to escape terrain, about half as much selection by about 75 m distance and by 200 m only 10% of the highest selection.

Terrain ruggedness did not factor as a strong and consistent variable, with equal selection and

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avoidance among individuals. There was little importance of year in habitat selection at the winter

range scale; we therefore removed year variables from further analyses. All cover classes showed

positive selection (relative to conifer). High-elevation grasslands and rock-rubble (where available)

were selected by all sheep. The mine areas cover class was selected by three-quarters of the sheep.

The linear and quadratic terms for elevation were the most influential of these variables, with distance

to escape terrain, solar radiation, high-elevation grasslands, and rock-rubble next in importance. The

winter range model was very robust (rs = 1.00).

Between home range and winter range scales, selection of the stronger and highest ranked

model variables remained consistent, with slightly less pronounced selection for topographic variables

at the winter range scale. At both scales selection for high-elevation grasslands/barren lands and rock- rubble cover classes were consistent. SelectionDraft for mine areas was ranked higher at the winter range scale compared with the home range scale.

Based on winter collar distribution and survey data (FLNO unpubl. data), we identified 29 km 2

of habitat used by sheep during winter among 15 areas, approximately 23 km 2 of which 13 ranges

could be considered main winter range. These main ranges comprise 2.7% of the study area, or 4.3% of

the merged annual sheep ranges, emphasizing the limited amount of used winter ranges within the

landscape.

Discussion

Faced by increasing snow depths and declining availability of forage, mountain ungulates have

a number of options to survive winter, including movement to lower elevation forested habitats,

increased use of steep terrain which tends to shed snow, or selection of high-elevation windswept

ridges (Lovari and Cosentino 1986; Forsyth 2000; Apps et al. 2001; Poole et al. 2009). Snow restricts

movements of mountain ungulates (Parker et al. 1984; Daily and Hobbs 1989), and mountain sheep

generally avoid snow depths >25–30 cm (Stelfox 1976; Sweanor et al. 1996). Our study indicated that

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20 bighorn sheep vary their use of native and mine-altered habitats during the year, but wintered primarily on high-elevation windswept, south-facing native grasslands. Elk Valley sheep used mine areas to a greater extent during summer and fall and to a lesser degree during winter. Although fidelity to winter ranges was high, individual variability in winter habitat selection was evident. Elevation, distance to escape terrain, solar duration, and high-elevation grasslands were main drivers of winter habitat selection.

Winter is an important season for sheep survival, as indicated by the reduced winter range size and higher numbers of mortalities during the deeper snow and more severe winter of 2010–11.

Greater cumulative snow depth and longer snow season may cause sheep to seek specific habitat types having less snow cover (Richard et al. 2014). Mortality rates were 3 times higher during the more severe winter (2010–11) compared with theDraft winter of low snow accumulation (2009–10), although precision was poor due to low sample size. Stelfox (1976) concluded that severe winters were accompanied by greater winter weight loss and increased lamb mortality. Burles and Hoefs (1984) observed a decline in a Dall sheep ( O. dalli) population in the Yukon following a severe winter characterized by above average snowfall and below average temperatures and wind speeds; older sheep (≥7 years of age) and lambs were most affected. White et al. (2008) documented a population decline of bighorn sheep in Yellowstone following a severe winter. However, Jorgenson et al. (1997) found that winter severity had no effect on ewe or ram survival in Alberta populations. The apparent impact of deeper snow on survival of Elk Valley sheep may be related to the high-elevation wintering strategy, leaving sheep with few options as snow depths increase and access to forage decreases

(Stelfox 1976). Thus the impact of winter severity on sheep populations may vary among wintering ecotypes.

Although snow depth affects survival of this high-elevation wintering population, other climatic factors may also influence survival. The effects of summer climate on forage quantity and quality have

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also been highlighted in several studies, for example excessive heat or cool wet summers can

negatively affect survival while high spring or summer precipitation can have a positive affect (Stelfox

1976; Portier et al. 1998; Parker et al. 2009; White et al. 2011; Johnson et al. 2013). Portier et al. (1998)

found that winter bighorn sheep lamb survival was not affected by winter weather, but that wet

springs had a positive effect on neonatal and winter survival of lambs, and warm springs increased

lamb survival the following winter.

Resource selection

Resource selection by sheep varied slightly by scale, with stronger selection and avoidance

observed at the home range scale. Terrain variables of elevation, distance to escape terrain, and solar

duration, and high-elevation grasslands/barren lands were the most important variables at both scales. Previous studies have found slope and distanceDraft to escape terrain as important variables in sheep habitat selection, along with aspect or solar radiation index and low snow cover (Stelfox 1976; Sweanor

et al. 1996; Dicus 2002; DeCesare and Pletscher 2006; Bleich et al. 2009). Selection of high winter solar

duration and open grasslands in our study coincided with areas where snow cover would be less.

Overall, we did not see differences in selection between sexes. Security may be more

important to ewes with lambs than to rams, because rams can better defend themselves against

predators, and ewes may make trade-offs of reduced forage benefits to minimize predation risk to

their offspring (Festa-Bianchet 1988 a; Bleich et al. 1997; Shackleton et al. 1999). Although we did not

detect differences in selection for distance to escape terrain between sexes during winter, these

differences may be more pronounced during spring and summer.

Our 2-stage modelling approach considered individual variation in selection of parameters. The

results appear to be superior to a mixed modelling approach that considers the individual as a random

term (Gilles et al. 2006), because it explicitly highlights the range of responses among sheep.

Bootstrapping parameter estimates with less weight given to individuals with high variance allowed

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more realistic models less influenced by outlier data. Given differences in use of mine areas by sheep, the individual modelling approach acknowledges different types of selection within the study area.

Differences in selection among herds and areas are also likely. DeCesare and Pletscher (2006) found that resource selection models for ewes across sites gave mixed results, such that models developed for 1 herd were not well validated to 2 other herds. These authors found nearness to escape terrain and grasslands were the only consistent variables in sheep winter resource selection among the 3 herds. Selection for cover types varied among herds, although high-visibility habitats were generally preferred.

Our modelling appeared to effectively capture used sheep winter ranges because models performed well when validated with independent data. However, areas lightly or not used by collared sheep were also captured in the models. ItDraft may be that these areas were unused but suitable winter ranges, but it is more likely that our modelling did not capture some aspect of sheep winter range selection important to sheep. Elevation and solar radiation can be considered surrogate variables to model snow but may not translate to the fine scale or accurately reflect true snow depths. Wind direction and strength would also influence snow cover and depth. Predation risk, distance among seasonal ranges, or other factors could be influencing selection of winter ranges (Festa-Bianchet

1988 b; Shackleton et al. 1999).

Use of mine areas

The Elk Valley sheep population used mine areas 10–20% of the time during winter. However, as predicted the topographic characteristics of sheep wintering areas on mines differed markedly from native wintering areas, occurring on average 300 m lower in elevation, on gentler slopes further from escape terrain. Bighorn sheep winter range on mines consisted of reclaimed native and non-native grass mixtures and spoils while pits and highwalls were used less. Reclaimed habitats tend to produce greater amounts of forage compared with native grasslands (Smyth 2014). Use of active and reclaimed

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mines by bighorn sheep has been previously observed (Jansen et al. 2007, 2009). MacCallum and Geist

(1992) documented use of a reclaimed open-pit coal mine on the east slopes of the Rocky Mountains

in Alberta where reclaimed grasslands provided over twice the productivity of native ranges and likely

contributed to large body mass of ewes and high lamb survival.

Bighorn sheep numbers in the Elk Valley East population have increased since the late 1980s (I.

Teske, BC FLNR, unpubl. data) concurrent with active coal mining and reclamation within or near their

main habitats. High pregnancy rates suggest the population currently is likely not at carrying capacity.

Jansen et al. (2006, 2007) found that desert bighorn sheep behaviour and activity at an active copper

mine was generally similar to that recorded while outside the mine perimeter. These authors

suggested modern mining might be predictable enough to allow sheep to habituate to human activities. At most of the Elk Valley mines sheepDraft are regularly observed along active haul roads and around office complexes (K. Podrasky, Teck Coal, pers. comm.). Oehler et al. (2005) observed no

differences in size of annual home ranges, composition of diet, and lamb:ewe ratios between ewes

inhabiting mined and non-mined areas. Similarly, Bleich et al. (2009) observed sheep being associated

with areas closer to limestone mines than were random points. Desert bighorn sheep used an active

mine more than during mine closure (Jansen et al. 2009). However, relative predation risk on mine

areas compared with native habitats is unknown. Despite high visibility in generally open habitat on

mines likely enabling early predator detection (Cristescu et al. 2015), wolves have consumed Dall sheep

and grizzly bears have killed bighorn sheep on reclaimed mine areas (Elliott 1984; Cristescu et al. 2015).

Since reducing predation risk influences habitat selection (Festa-Bianchet 1988 a), predation may

influence habitat selection at the mine versus native habitat scale. Benefits to sheep populations from

occupancy of mine areas depends on demographic responses (Oehler et al. 2005), which can be

difficult to quantify (Bleich et al. 2009).

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Conclusions

Bighorn sheep require a balance of habitats supplying adequate year-round forage and security

from predation. Important components of winter habitat include juxtaposition of escape terrain and

adequate and accessible forage, and warm aspects (Tilton and Willard 1982; Jansen et al. 2006). There

are limited possibilities to use management to enhance winter habitat, which for the most part are

static and in specific locations within the environment. Avoidance of conifers and use of high visibility

habitats suggest that forest encroachment would be detrimental to sheep (Wakelyn 1987), and thus

could be controlled in areas where this is occurring using slashing or burning (Smith et al. 1999). Mines can reproduce or mimic many of the static features of sheep range in general (MacCallum and Geist

1992; Bleich et al. 1997; Jansen et al. 2006), but may be unable to reproduce the low snow depths that are important for high quality winter rangeDraft (Stelfox 1976). When developing suitable sheep winter range on mine areas, escape terrain in close proximity

(within 100 m) of high quality forage should be considered (MacCallum and Geist 1992). Using GIS to model prevailing wind direction in association with solar radiation could assist in pinpointing areas where snow depths would remain low (<30 cm) throughout the winter season. Reclamation should include maintenance of accessible or stepped highwalls (Jansen et al. 2006). Optimal design of features to enhance forage and security could reduce predation risk within these habitats. Security terrain is especially important for ewes and lambs, which may preferentially reduce predation risk at the expense of forage quality (Bleich et al. 1997). Reduced survival or reproduction in sheep could occur during more severe winters if sheep are attracted to areas that contain adequate physical structures but that otherwise lack some of the conditions or resources normally associated with those indicators under less than pristine conditions (i.e., they are ecological traps; Bleich et al. 2009).

High fidelity to winter ranges and the apparent influence of winter severity on survival suggest that disturbance to native winter range resulting from development should be minimized or be

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conducted in a manner that effectively manages and/or mitigates the impacts. Options for addressing

loss of high quality winter range through mitigation may be limited. The degree to which disturbance

to or removal of portions of existing winter range will affect either the carrying capacity of the range or

population numbers is difficult to quantify, as many factors influence overall population health and

survival. However, it is likely that large scale removal of high-elevation winter ranges in the absence of

effective mitigation measures would result in population declines.

Acknowledgements

This study was a joint effort developed through Teck Coal and BC FLNRO. Teck Coal, BC FLNRO,

and the Habitat Conservation Trust Foundation (HCTF) provided primary funding for this project, with

additional funding from The Wild Sheep Society and the East Kootenay Wildlife Association (BC Wildlife Federation). Teck Coal and BC FLNRO providedDraft much in-kind support. Bighorn Helicopters expertly conducted aerial captures. BearAir carried out telemetry flights, D. Lewis and L. Ingham assisted with

ground captures and collar collection, and H. Schwantje provided veterinarian and capture support.

We thank the following for assistance with fieldwork or logistics: L. Amos, T. Barr, T. Caldwell, N.

Caulkett, D. Charest, T. Chala, A. Fahlman, W. Franklin, K. Honeyman, N. Manklow, S. Medcalf, V.

Naude, L. Ozeki, K. Penney, B. Phillips, G. Sword, T. Szkorupa, S. Thiel, J. Thorner, G. Wilson, D. Vasiga,

and J. Volp, as well as various Teck staff who provided sightings and support. W. Burt skillfully

conducted the GIS analyses and extractions, and map production. The manuscript benefitted from

helpful comments by G. Mowat, B. MacCallum, and 2 anonymous reviewers.

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List of Figures

Fig. 1. The Elk Valley bighorn sheep study area in southeastern British Columbia, 2009 to 2011. Mine

areas refer to the cumulative disturbance footprint up to 2011.

Fig. 2. Proportion (%) of use on mine areas by collared bighorn sheep ewes and rams in the Elk Valley,

British Columbia, March 2009 – May 2011.

Fig. 3. Distribution of topographic variables for the random home range (RHR), random winter range

locations for winter 2009–10 (RWIN1) and 2010–11 (RWIN2), and sheep use on winter ranges for

winter 2009–10 (SWIN1) and 2010–11 (SWIN2), by female (F) and male (M) bighorn sheep in the Elk

Valley, British Columbia, 2009–11 (open symbols; box and whisker plots with outliers [solid symbols]). Y-axis values: elevation (m); slope (%); terrainDraft ruggedness (units); distance to escape terrain (m); solar (units. See Table 1 for description of acronyms.

Fig. 4. Distribution of cover class variables for the random home range (RHR), random winter range

locations for winter 2009–10 (RWIN1) and 2010–11 (RWIN2), and sheep use on winter ranges for

winter 2009–10 (SWIN1) and 2010–11 (SWIN2), by female (F) and male (M) bighorn sheep in the Elk

Valley, British Columbia, 2009–2011 (open symbols; box and whisker plots with outliers [solid

symbols]). Y-axis values are cover classes (scaled from 0–1). See Table 1 for description of acronyms.

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Draft

Fig. 1. The Elk Valley bighorn sheep study area in southeastern British Columbia, 2009 to 2011. Mine areas refer to the cumulative disturbance footprint up to 2011.

194x248mm (300 x 300 DPI)

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Ewes 50 Rams 40 Draft

Proportion (%) oflocations on mine property 10

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2000 ELEV 1800 1600 1400

80 SLOPE

40 0.07 Draft

0.06 RUGGED 0.05 0.04 Value 0.03 0.02

100 ESCPDST

600 SOLAR 500

400

300 https://mc06.manuscriptcentral.com/cjz-pubs RHR RWIN1 RWIN2 SWIN1 SWIN2 RHR RWIN1 RWIN2 SWIN1 SWIN2 SEASON Page 39 of 46 Canadian Journal of Zoology F M 0.8 0.6 CONIF 0.4 0.2 0.0 0.6

0.4 CUT 0.2 0.0

0.3 SHRUB 0.2 0.1 0.0 GLHIGH 0.4 Draft 0.2

0.0 GLLOW

Value 0.3 0.2 0.1 0.0

0.75 BARREN 0.50 0.25 0.00

0.4 ROCRUB 0.3 0.2 0.1 0.0 1.00

0.75 MINE 0.50 0.25 0.00 https://mc06.manuscriptcentral.com/cjz-pubs RHR RWIN1 RWIN2 SWIN1 SWIN2 RHR RWIN1 RWIN2 SWIN1 SWIN2 SEASON Canadian Journal of Zoology Page 40 of 46

Table 1. Description of variables examined in modelling habitat selection by bighorn sheep in the Elk

Valley, southeastern British Columbia, 2009 to 2011.

Category Acronym Variables

YEAR Represents less severe (0) and more severe (1) winters

Terrain variable ELEV Elevation (m)

ELEV2 Squared term for elevation

SLOPE Slope (%)

RUGGED Index of ruggedness (Sappington et al. 2005) ESCPDST DistanceDraft to nearest escape terrain (m), defined as slope ≥75% SOL AR Solar duration value (hrs)

Cover class * GLHIGH High -elevation grasslands , forbs, >33% herb

GLLOW Low -mid -elevation g rass lands, forbs , >33% herb

BARREN Barren, non -vegetated areas

ROCRUB Bedrock, rubble, talus

SHRUB >20% ground cover, with >33% shrub

CUT Cutblocks , low to mid -elevation

MINE Disturbed mine areas

OTHER Includes deciduous -leading stands and wetlands

CONIF Conifer -leading stands

* Cover class modified from EOSD coverage (Wulder et al. 2008); see text.

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Table 2. Seasonal range sizes (90% kernel;km 2) for bighorn sheep (summer 2009, 2010 and winter

2009–10, 2010–11), Elk Valley, southeastern British Columbia.

Both sexes* Females Males

Season n x SE x SE x SE

Summer 2009 30 33.0 5.34 32.3 5.63 33.8 9.72

Summer 2010 30 37.0 7.88 26.8 4.23 48.7 15.93

Winter 2009–10 30 9.5 2.56 10.3 3.10 8.5 4.31

Winter 2010–11 29 3.2 0.58 3.9 0.09 2.3 0.54

* Range sizes did not differ between sexes for any season ( P > 0.18) Draft

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Table 3. Mean habitat variable values for wintering bighorn sheep (2009–10 and 2010–11) in the Elk

Valley, southeastern British Columbia. Mining ( n = 23 sheep-winters) vs. native grassland areas ( n =

60 sheep-winters).

Mine Native 90 th 90 th

x † SE percentile x 2 SE percentile Variable * range ‡ range ‡

Topographic Elevation (m)* 1839 50.7 1340–2096 2156 24.3 1779–2355 Slope (%)* 48 2.4 31–65 72 1.1 61–88 Terrain ruggedness* 0.035 0.0032 0.020–0.058 0.046 0.0017 0.027–0.070 Dist. to escape terrain (m)* 51 4.3 18–79 23 1.4 8–44 Solar duration (hrs)* 563 25.2 Draft 256–663 608 8.8 500–675 Mine cover classes (%) Reclaimed lands 35 6.0 0–84 Spoil 29 5.5 0–74 Pit 8 2.4 0–31 Highwall 8 2.7 0–32 Footwall 1 0.3 0–3 Roads 1 0.6 0–4 Undifferentiated 18 3.5 0–47 * Significant differences ( t-test, P < 0.05) between mine vs. native indicated with asterisks.

† Mean values calculated as mean of means from individual sheep locations.

‡ Mean upper and lower limits of 90th percentile ranges are from individual sheep.

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Table 4. Variables predicting winter habitat selection at the home range scale by bighorn sheep

based on GPS collar data, Elk Valley, southeastern British Columbia. Presented are standardized

coefficients (Std. Coefficient) and 95% confidence intervals, and the proportion of sheep that showed

positive selection for that parameter (sheep showing positive selection/sample size). Winter 2009–

10 is coded as 0 and 2010–11 is coded as 1, thus negative interaction terms with year signify a

reduced selection for that variable during the deep snow winter of 2010–11. Numbers with the

strongest selection or avoidance are bolded.

Std. Coefficient % sheep

Variable * β Lower 95% CI Upper 95% CI selected ( n)

YEAR –0.81 –2.67 0.76 42 (13/31) Terrain Draft ELEV 18.4 8.1 30.7 84 (26/31)

ELEV ×YR 3.73 –12.19 21.24 52 (16/31)

ELEV2 –18.1 –29.6 –8.0 19 (6/31)

ELEV2 ×YR –4.95 –22.33 11.96 48 (15/31)

RUGGED 0.33 0.12 0.56 68 (21/31)

ESCPDST –2.59 –3.23 –2.14 0 (0/31)

ESCAPE ×YR –0.51 –1.00 –0.03 39 (12/31)

SOLAR 2.11 1.74 2.51 97 (30/31)

SOLAR ×YR –0.10 –1.06 0.98 48 (15/31)

Cover class †

GLHIGH ‡ 1.93 1.59 2.26 100 (31/31)

GLLOW 0.61 0.44 0.80 83 (24/29)

ROCRUB 0.99 0.61 1.32 48 (12/25)

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SHRUB 0.51 0.33 0.71 73 (22/30)

CUT 0.53 0.17 0.94 30 (8/27)

MINE 0.58 0.05 1.11 45 (14/31)

OTHER 0.24 0.15 0.33 48 (12/25)

CONIF 0 0 (0/31)

* See Table 1 for description of acronyms.

† Conifer is the reference value for all other cover classes, thus coefficient values and whether sheep selected the cover class were relative to selection of conifer.

‡ GLHIGH includes BARREN.

Draft

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Table 5. Variables predicting winter habitat selection at the winter range scale by bighorn sheep

based on GPS collar data, Elk Valley, southeastern British Columbia. Presented are parameter

estimates (Coefficient), standardized coefficients (Std. Coefficient) and 95% confidence intervals,

relative importance (RI), and the proportion of sheep that showed positive selection for that

parameter. Winter 2009–10 is coded as 0 and 2010–11 is coded as 1, thus negative interaction terms

with year signify a reduced selection for that variable during the deep snow winter of 2010–11.

Numbers with the strongest or significant selection or avoidance are bolded.

Std. Coefficient % sheep

Variable * β Lower 95% CI Upper 95% CI selected ( n)

YEAR 0.17 –1.87 1.67 52 (13/25) Terrain Draft ELEV 12.2 6.7 17.9 85 (28/33)

ELEV ×YR –4.66 –14.18 7.26 36 (9/25)

ELEV2 –11.7 –17.3 –6.1 18 (6/33)

ELEV2 ×YR 4.90 –6.15 14.19 64 (16/25)

RUGGED 0.04 –0.15 0.24 52 (17/33)

ESCPDST –1.61 –2.02 –1.23 9 (3/33)

ESCAPE ×YR –0.30 –0.67 0.02 40 (10/25)

SOLAR 1.84 1.52 2.20 97 (32/33)

SOLAR ×YR 0.12 –0.76 1.08 48 (12/25)

Cover class †

GLHIGH ‡ 1.57 1.34 1.81 100 (33/33)

GLLOW 0.68 0.51 0.87 81 (26/32)

ROCRUB 1.03 0.70 1.37 100 (14/14)

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SHRUB 0.52 0.36 0.69 75 (24/32)

CUT 0.75 0.28 1.26 33 (7/21)

MINE 0.77 0.51 1.07 78 (21/27)

OTHER 0.25 0.16 0.31 42 (13/31)

CONIF 0 0 (0/33)

* See Table 1 for description of acronyms.

† Conifer is the reference value for all other cover classes, thus coefficient values and whether sheep selected the cover class were relative to selection of conifer.

‡ GLHIGH includes BARREN.

Draft

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