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The influence of wolves on the ecology of mountain caribou Allison, Bradley Armstrong 1998

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THE INFLUENCE OF WOLVES ON THE ECOLOGY OF MOUNTAIN CARIBOU  by BRADLEY ARMSTRONG ALLISON H . B . O . R . , Lakehead University, 1991 B . S c , Lakehead University, 1991  A THESIS SUBMITTED IN P A R T I A L F U L F I L L M E N T OF THE REQUIREMENTS FOR T H E D E G R E E OF M A S T E R OF SCIENCE in THE F A C U L T Y OF G R A D U A T E STUDIES (Department o f A n i m a l Science)  We accept this thesis as conforming tp the required standard  THE UNIVERSITY OF BRITISH C O L U M B I A M a y 1998 © Bradley Armstrong Allison, 1998  In  presenting this  degree at the  thesis  in  partial  fulfilment  of  the  requirements  University  of  British  Columbia,  I agree that the Library shall make it  freely available for reference and study. I further agree that copying  of  department  this thesis for scholarly or  by  his  or  her  for  It  is  understood  head of my  that  publication of this thesis for financial gain shall not be allowed without permission.  Department The University of British Columbia Vancouver, Canada  DE-6 (2/88)  advanced  permission for extensive  purposes may be granted by the  representatives.  an  copying  or  my written  ABSTRACT The w o l f (Canis lupus)/mountam caribou (Rangifer tarandus caribou) spatial separation model (Seip 1992a) was examined in highland and mountainous areas o f east central and southeastern British Columbia to determine the influence o f wolves on mountain caribou ecology. Three key elements investigated were: the importance o f w o l f predation as a mountain caribou mortality factor, the elevational relationships o f mountain caribou, wolves, and moose (Alces alces), and the seasonal dietary importance o f moose to wolves. Mountain caribou mortality data from the Columbia Mountains and Quesnel Lake supported the hypothesis that w o l f predation is greater i n highland than i n mountainous areas. W o l f predation was the main mortality factor o f caribou i n the highlands around Quesnel Lake, but was a minor factor i n the other three study areas. W o l f predation at Quesnel Lake occurred primarily during summer/fall at low elevations. Mountain caribou, w o l f and moose radio-telemetry data suggested that wolves i n both highland and mountainous areas are more closely associated elevationally with moose than caribou throughout the year. Caribou i n highland areas appeared as adept as those i n rugged mountains at spacing elevationally away from wolves during all seasons. Significant elevational overlap between wolves and caribou occurred only i n the rugged Columbia Mountains, and then, only during summer/fall. W o l f scats from the Columbia Mountains indicated that mountain caribou were o f lesser dietary importance than moose to wolves throughout the year. Moose, particularly i n winter, were the main diet item o f wolves. Beaver was an important dietary item o f wolves during summer/fall. Elevational separation appears inadequate i n explaining the variation i n w o l f predation on mountain caribou i n the highland and mountainous study areas. Differences i n w o l f densities and  the relative densities o f moose and caribou may be the main factors influencing the susceptibility o f mountain caribou to w o l f predation. It is recommended that studies be conducted i n both topography types across a wide range o f caribou and moose densities to better explain the influence o f wolves on mountain caribou ecology. In addition, it w i l l be necessary to compare the geographic distribution o f mountain caribou and wolves to assess i f geographic rather than elevational separation occurs. Determining how timber harvesting impacts the numerical and spatial responses o f wolves, mountain caribou and moose is also recommended.  iii  TABLE OF CONTENTS Abstract  ii  Table of Contents  iv  List of Tables  I  vi  List of Figures  vii  Acknowledgements  viii  Chapter 1. General Introduction  1  Study Areas  2  Chapter 2. Wolves: Their role in mountain caribou mortality Introduction  6  Methods  7  Results  10  Discussion  14  Chapter 3. Elevational relationships of mountain caribou, wolves and moose Introduction  18  Methods  19  Results  20  Discussion  24  Chapter 4. The seasonal dietary importance of moose to wolves in the Columbia Mountains Introduction  27  Methods  27  Results  28  Discussion  30  iv  Chapter 5. General conclusion and management recommendations Literature Cited  LIST OF TABLES Table 1.  Table 2.  Table 3.  Table 4.  Table 5.  Level o f mountain caribou monitoring and annual mortality rate ( % ) ± (sd) within study areas.  11  Causes o f radio-collared mountain caribou mortalities within study areas.  12  Seasonal causes o f radio-collared mountain caribou mortalities across all study areas (Columbia Mountains 1981-85, 1992-96; Wells Gray Provincial Park 1986-89; Quesnel Lake 1984-89; Quesnel Highlands 1993-96). Length o f season (days) i n brackets.  12  Type and frequency o f mortalities occurring above or below the median seasonal elevation o f radio-collared mountain caribou (Columbia Mountains 1981-85, 1992-96; Wells Gray Provincial Park 1986-89; Quesnel Lake 1984-89; Quesnel Highlands 1993-96).  13  Mountain caribou density estimates based on mean annual distribution o f populations.  14  Table 6.  M e a n w o l f density estimates within pack territories.  14  Table 7.  Wolf/mountain caribou and wolf/moose seasonal intermedian distances (m) within study areas.  24  Frequency o f occurrence and percent biomass o f w o l f diet items by season i n the Columbia Mountains.  29  Table 8.  vi  LIST OF FIGURES Figure 1.  Figure 2.  Figure 3.  Figure 4.  Figure 5.  Location o f study areas: Columbia Mountains, Wells Gray Provincial Park, Quesnel Lake and Quesnel Highlands, British Columbia.  3  Columbia Mountains radio-collared mountain caribou, w o l f and moose median monthly elevations and interquartile ranges (1992-96 caribou and wolf; 1984-85 moose).  21  Wells Gray Provincial Park radio-collared mountain caribou and w o l f median monthly elevations and interquartile ranges (1986-89).  21  Quesnel Lake radio-collared mountain caribou, w o l f and moose median monthly elevations and interquartile ranges (1984-89).  22  Quesnel Highlands radio-collared mountain caribou and w o l f median monthly elevations and interquartile ranges (1993-96).  22  vii  ACKNOWLEDGEMENTS " W e are made to exaggerate the importance o f what work we do; and yet how much is not done by us!" (Thoreau 1854) F r o m the outset o f m y search for knowledge and understanding o f wolves, mountain caribou and science i n general, many individuals graciously helped me move toward attaining this goal. A s always, I recognize m y path was directed by G o d and that He sustained me faithfully throughout this adventure. I thank my wife, Sue, for providing patient support and steadfast encouragement throughout all the various phases o f this thesis, in particular when I was absent from home and out tracking wolves i n the Columbia Mountains. I greatly appreciate my family members and friends for cheering me onward and providing necessary moments o f levity. B . N . M c L e l l a n , m y research supervisor guided me throughout this research, and most notably provided sage advice, fund-raising support, good coffee and occasionally a floor to crash on. D . Shackleton, m y academic supervisor, tried diligently to keep me focused whenever I was within arms length o f "the hut". M y other committee members: D . Seip, M . Pitt, and T. Sullivan, assisted freely, whenever asked. R . Ramcharita deserves a special note o f thanks for volunteering countless hours i n m y initial efforts to radio-collar wolves. In addition to almost always keeping the snowmobile tanked up, he shared many ideas which undoubtedly increased the quality o f this research. J. Flaa assisted i n the field, and together with M . Super and D . M a i r spent many hours airborne, locating radiocollared caribou and wolves in the Columbia Mountains. J. Townley helped i n backtracking wolves during the winter, while the assistance o f D . Lewis, J. Krebs, K . Atkinson and A . L a y also eased m y work i n the field. G . G a l z i , S. W i l s o n and S. Leung offered advice and assistance with various database queries. I. Clemens assisted with computer programming and data analysis. K . A l l i s o n created a map o f the study areas. Individuals that generously provided data for analysis were: B . N . M c L e l l a n and J. Flaa, K . Simpson and G . Woods (Columbia Mountains caribou data); D . Seip (Quesnel Lake caribou/wolf/moose data; Wells Gray caribou/wolf data); J. Y o u n g and P. Dielman (Quesnel Highlands caribou/wolf data); K . Simpson (Columbia Mountains moose data); and J. Woods (Banff National Park ungulate weight data). Financial support for this study was provided by the B . C . Ministry o f Forests, Mount Revelstoke and Glacier National Parks, Columbia Basin Fish and Wildlife Compensation Program, Canadian Wildlife Service - University Research Support Fund, The Friends o f Mount Revelstoke and Glacier, Revelstoke Community Forest Corporation, Canadian Wildlife Federation - Orville Erickson Memorial Scholarship, University o f British Columbia - James Robert Thompson Fellowship, E . A l l i s o n , E . and W . Bauman and P. Spaetzel. Although they w i l l never read this, I am grateful for the wolves, caribou and moose that through direct and indirect observation, allowed me a brief glimpse into their lives and provided many enjoyable moments afield. Ultimately, I realize that together with the help o f many fine individuals, m y burdens were lightened, progress was made, and I was truly allowed to enjoy "one o f the best times o f m y life" ( B . N . M c L e l l a n , personal communication).  CHAPTER 1 GENERAL INTRODUCTION Mountain caribou  (Rangifer tarandus caribou) are on the blue or vulnerable list o f species i n  British Columbia (Harper et al. 1994). Their low numbers (Simpson et al. 1997), low rate o f reproduction (Bergerud 1978) and reliance on old growth forests for food and shelter which conflicts with timber harvesting needs (Simpson et al. 1996) collectively resulted in this designation. The distribution and abundance o f mountain caribou began to decrease across southern British Columbia i n the early 1900s (Munro 1947). Explanations for the decline have included habitat degradation, overhunting and predation (Bergerud 1974; Bloomfield 1980; Edwards 1954; Munro 1947), but w o l f  (Canis lupus) predation has recently been suggested as the most  significant limiting factor across southern British Columbia (Bergerud 1988a, 1996; Seip 1991,  ( 1992a; Seip and Cichowski 1996). Bergerud and Elliot (1986) and Seip (1992a) suggested that w o l f populations within mountain caribou range are sustained primarily by moose  (Alces alces),  but in areas o f gentle topography, wolves were also a significant predator o f mountain caribou. Ultimately, it was concluded that mountain caribou i n more rolling, highland areas experience more w o l f predation than caribou in rugged mountainous areas due to their inadequate degree o f spatial separation with wolves and moose (Seip 1992a). To determine the influence o f wolves on the ecology o f mountain caribou, I investigated three elements o f the spatial separation model (Seip 1992a): the importance o f w o l f predation as a mountain caribou mortality factor, the elevational relationships o f caribou, wolves and moose, and the seasonal dietary importance o f moose to wolves. Chapter 1 includes this introduction to the research and a description o f the four study areas. In Chapter 2,1 use mountain caribou mortality data collected during five previous studies ( B . N . 1  M c L e l l a n and J. Flaa, unpublished data; Seip 1990, 1992b; Simpson and Woods 1987; J . A . Y o u n g and P. Dielman, unpublished data) to compare the rates, causes, seasons and elevations o f mountain caribou mortality among four study areas, and i n particular, the impact o f w o l f predation on animals in highland and mountainous areas. In Chapter 3,1 use radio-telemetry data ( B . N . M c L e l l a n , J. Flaa, and B . A l l i s o n , unpublished data; Seip 1990, 1992b; K . Simpson, unpublished data; J . A . Young, unpublished data) from the four study areas to examine the elevational relationships o f mountain caribou, wolves and moose i n both highland and mountainous areas. In Chapter 4,1 estimate the seasonal dietary importance o f moose to wolves in the Columbia Mountains using w o l f scats I collected from that area. The final chapter summarizes m y findings and provides some management recommendations.  STUDY AREAS The four study areas are located in east central and southeastern British Columbia. These areas are: the Columbia Mountains north o f Revelstoke (51°30'N, 118°15'W), Wells Gray Provincial Park (52°15'N, 120°W), the Quesnel Lake area (52°45'N, 120°45'W) and the Quesnel Highlands (52°45'N, 121°W) near Horsefly (Fig. 1). The Quesnel Highlands study area encompasses the Quesnel Lake study area, but since the data examined come from two different time periods, and the areas are o f considerably different size, they are referred to as separate studies. Although there are some moderate slopes and plateaus, the topography o f the Columbia Mountains and the majority o f Wells Gray Provincial Park is typical o f rugged terrain. Valleys are narrow and steep with mountains rising to massive rock outcrops and glaciers. Elevations range from 575 to 3200 m in the Columbia Mountains and 630 to 2850 m i n Wells Gray Provincial Park. A n estimated 375 caribou inhabit the Columbia Mountains study area ( M c L e l l a n  2  F i g . 1. Location o f study areas: Columbia Mountains, Wells Gray Provincial Park, Quesnel Lake, and Quesnel Highlands, British Columbia.  3  et al. 1994b), while 275 and 302 were estimated within and adjacent to Wells Gray Provincial Park i n 1987 and 1989 respectively (Seip 1990). The Quesnel Lake and Quesnel Highlands study areas contain some rugged mountains but generally consist o f highlands with rounded, vegetated mountains and moderate slopes (Holland 1976). Large rock outcrops are rare and glaciers are absent. Elevations range from 728 m to almost 2600 m. The Quesnel Lake study area was estimated to have 220 mountain caribou in 1986, but dropped to 94 i n 1989 and 95 i n 1997 (Seip 1992b; J . A . Y o u n g , unpublished data). In 1997, 358 mountain caribou (J.A. Young, unpublished data) were estimated to reside i n the Quesnel Highlands area. A portion o f the Quesnel Highlands study area is rugged with mountains reaching 2800 m. The majority o f the valley bottoms within the four study areas are i n the Interior-CedarHemlock (ICH) biogeoclimatic zone which occurs up to 1250 - 1350 m. Forests i n the I C H zone are dominated by closed canopies o f western red cedar {Thuja plicata) and western hemlock {Tsuga heterophylla)  (Ketcheson et al. 1991). L o w elevations i n the northern and western  portions o f the Quesnel Highlands contained the Sub-Boreal Spruce (SBS) zone and are dominated by hybrid white spruce (Picea engelmannii x glauca) and subalpine fir {Abies lasiocarpa) (Meidinger et al. 1991). M i d slopes in the four study areas are occupied by the Engelmann Spruce-Subalpine F i r (ESSF) biogeoclimatic zone, which extends to 1800 - 2000 m. Lower portions o f the E S S F are dominated by closed canopy forests o f hybrid white spruce and subalpine fir (Coupe et al. 1991) while subalpine fir increases i n abundance at higher elevations and near timberline, grows i n small clumps, forming the more open subalpine parkland (Coupe et al. 1991). The Alpine Tundra ( A T ) zone occurs at the highest elevations i n the four study areas. Snowpacks i n the mountainous portions o f the study area are high and average 1.3 m i n valley bottoms and 2.8 m at snow stations between 1620 and 1920 m ( B . C . Ministry o f Environment  4  1985). Snowfall accumulations i n the Quesnel Highlands are lower than those o f the mountains and average 1.0 m i n valley bottoms and 1.9 m at stations between 1570 and 1900 m during the late winter ( B . C . Ministry o f Environment 1985). Ungulates in all four study areas include: woodland caribou, moose, mule deer (Odocoileus hemionus), white-tailed deer (O. virginianus), and mountain goats (Oreamnos Wolves, grizzly bears (Ursus arctos), black bears (U. americanus),  americanus).  and wolverines (Gulo gulo)  are the main predators within the study areas.  5  CHAPTER 2 WOLVES: THEIR ROLE IN MOUNTAIN CARIBOU MORTALITY INTRODUCTION Since the latter part o f the nineteenth century, woodland caribou populations have declined across a large portion o f their historic North American range (Anderson 1938). In southern British Columbia, the mountain ecotype began decreasing around 1900 and disappeared from some areas (Munro 1947). Currently there are about 2500 mountain caribou and almost all are found i n British Columbia (Simpson et al. 1997). Mountain caribou feed largely on arboreal lichen during the winter (Antifeau 1987, Edwards and Ritcey 1960, Rominger and Oldemeyer 1990, Servheen and L y o n 1989; Simpson et al. 1997; Terry 1994), and because these lichens are most abundant on old trees, caribou management conflicts directly with forest harvesting. Maintaining present mountain caribou populations and their requisite habitats is a major conservation challenge i n British Columbia (Simpson et al. 1997, Stevenson and Hatler 1985). Several hypotheses have been proposed to explain the decline o f mountain caribou. Overharvesting by native (Munro 1947) and recreational hunters due to liberal hunting regulations, combined with improved access, was suspected (Bergerud 1974; Bloomfield 1980; Seip and Cichowski 1996). Habitat degradation due to fires (Edwards 1954) and timber harvesting was thought to reduce caribou winter food, decrease shelter and disrupt travel corridors (Bloomfield 1980). In portions o f southern British Columbia, cougar (Puma  concolor)  predation was believed to have caused the extirpation o f caribou i n one area (Munro 1947), while predation by an increasing w o l f population during the 1930s and 1940s was suggested i n other areas (Bergerud 1974; Munro 1947). Although more pluralistic hypotheses have been proposed to explain the widespread decline in mountain caribou (Bloomfield 1980; Stevenson and Hatler  6  1985), w o l f predation is currently emphasized as the primary factor involved (Bergerud 1974, 1988a, 1996; Seip 1991, 1992a; Seip and Cichowski 1996). Bergerud and Elliot (1986) reported that in areas where wolves are abundant, caribou populations decline, and Seip (1992a) proposed that mountain caribou i n highland areas with less ability to space away from wolves and moose, experience more w o l f predation particularly i n summer, than caribou i n rugged mountains. The objectives o f this chapter are to: 1) compare rates, causes, seasons and elevations o f mountain caribou mortality among four study areas, 2) compare w o l f and caribou densities among those study areas, and 3) test whether w o l f predation on caribou is greater i n highland areas than i n mountainous areas.  METHODS Capture and monitoring The mountain caribou mortality data from the four study areas (Columbia Mountains; Wells Gray Provincial Park; Quesnel Lake; Quesnel Highlands) were collected during five previous studies ( B . N . M c L e l l a n and J. Flaa, unpublished data; Seip 1990,1992b; Simpson and Woods 1987; J . A . Y o u n g and P. Dielman, unpublished data). Most mountain caribou i n the four study areas were captured by net-gunning from a helicopter i n subalpine habitats during late winter ( M c L e l l a n et al. 1994a; Seip 1990, 1992b; Y o u n g et al. 1996). Other caribou were captured by darting with immobilization drugs during the 1981-85 Columbia Mountains study (Simpson and Woods 1987) and at the start o f the 1984-89 Quesnel Lake study (Seip 1992b). A l l caribou were fitted with mortality-motion sensitive radio collars. Radio-collared animals were relocated from aircraft two to four times per month and location and elevation were recorded for each relocation.  7  Mortality factors and seasons When a signal from a motion-sensitive radio collar indicated that a caribou was dead, the site was investigated and i f possible, the caribou was necropsied to determine cause o f death. Mortality causes were first classified as predation, not predation, and unknown. Cases o f predation were then divided into wolf, bear, wolverine or unknown. Non-predation deaths were divided into accident, avalanche, malnutrition or unknown. Criteria used to attribute predation mortalities to wolf, bear or wolverine were: presence o f predator, tracks, scats, or hair at the mortality site; extensive blood on carcass, ground or snow; distribution o f the carcass, and site conditions ( B . N . M c L e l l a n and J. Flaa, unpublished data; D . R . Seip, unpublished data; J . A . Y o u n g , unpublished data). Malnutrition deaths in all four study areas were determined from bone marrow fat content ( B . N . M c L e l l a n and J. Flaa, unpublished data; D . R . Seip, unpublished data; K . Simpson, personal communication; J . A . Young, unpublished data), with renal and rump fat measures also used i n the Columbia Mountains. Mortalities were classified as unknown when insufficient carcass remnants were available at the mortality site to assign a mortality cause. Mortalities were also classed as not predation but unknown when necropsies on relatively intact carcasses failed to reveal a mortality agent. A l l caribou mortalities were assumed to be classified to the correct mortality factor. Date o f mortality was assumed to fall halfway between the date the caribou was last relocated alive and the date when the mortality signal was first heard (Mayfield 1975). I classified season o f mortality as spring ( M a y 1 - M a y 28), summer/fall (May 29 - N o v 7), early winter (Nov 8 Dec 7) or late winter (Dec 8 - A p r 30). These seasons denote shifts i n elevation and habitat use by mountain caribou and correspond to the conventional seasons o f other mountain caribou studies (Antifeau 1987; Simpson and Woods 1987; Terry et al. 1996). Seasonal delineation for  8  each study area was determined by plotting mean caribou elevation by Julian week ( M c L e l l a n et al. 1994a). Boundaries between seasons were then identified when the weekly change between mean caribou elevation was greatest, indicating a major elevational shift. Seasonal boundaries were then averaged among study areas to attain the four standardized seasons.  Mountain caribou and wolf density estimates The 9 5 % adaptive kernel method (Seaman and Powell 1991) was used to estimate the annual distribution o f each caribou population and w o l f pack from the relocations o f radio-collared animals. Density estimates for each population were obtained by dividing the mean annual distribution by the estimated number o f individuals in each caribou population or w o l f pack. The mean o f all w o l f pack density estimates i n each study area was used for inter-study comparisons. Estimates o f caribou population size in each study area were obtained from censuses conducted during late winter as described by M c L e l l a n et al. (1994b), Seip (1990, 1992b) and J . A . Y o u n g (personal communication). Estimates o f winter w o l f pack sizes (Fuller and Snow 1988) were obtained during relocation flights, while the number o f packs i n each study area were estimated from incidental observations and w o l f radio-telemetry data ( B . N . M c L e l l a n , J. Flaa, and B . A l l i s o n , unpublished data; Seip 1990, 1992b; J . A . Young, unpublished data).  Statistical analyses Annual mortality rates were estimated by dividing the total number o f caribou mortalities by the total number o f caribou years (one caribou monitored for one year is a caribou-year), based on methods described by White and Garrott (1990) for survival data. Multiple male and female caribou were radio-collared i n the Columbia Mountains so tests for differences i n male and  9  female caribou mortality rates were made. I f no difference between rates was found, the mortality data o f both sexes were pooled. Chi-square tests o f independence (Siegel and Castellan 1988) were used to test for differences i n mortality rates among genders and areas. I f different rates were found among areas, the contingency table was partitioned into subtables following the methods o f Siegel and Castellan (1988) to determine the cause o f the difference. Fisher's extact test (Siegel and Castellan 1988) was used to test whether w o l f predation on caribou is higher in highland areas than in mountainous areas using mortality from study areas with adequate sample sizes. Levels o f significance for all tests were set atp < 0.05.  RESULTS Mortality rates In total, 155 mountain caribou (132 adult females; 11 adult males; 9 juvenile females; 3 juvenile males) from the four study areas were radio-collared and monitored for a total o f 410 caribouyears between 1981 and 1996 (Table 1). Different annual mortality rates were found among the four study areas  = 15.56, d f = 3,p = 0.002). Partitioning the contingency table showed no  difference between rates i n Wells Gray Provincial Park, the Columbia Mountains and the Quesnel Highlands  = 0.18, df = 1, p = 0.75), but the mortality rate o f caribou at Quesnel  Lake was higher than the other study areas (%  - 14.56, d f = 1, p < 0.001).  10  Table 1. L e v e l o f mountain caribou monitoring and annual mortality rate ( % ) ± (sd) within study areas. Study Area  No. of caribouyears of monitoring  No. (age - sex) of radio-collared animals  Annual mortality rate  Ad F  AdM  JuvF  Juv M  Total  Columbia Mountains 1981-85 & 1992-96 Wells Gray Prov. Park 1986-89 Quesnel Lake 1984-89 Quesnel Highlands 1993-96  162  41  9  7  3  60  123 + 2 6 "  75  27  1  2  -  30  8.0 ± 3 . 1  81  31  1  -  -  32  28.4 ± 5 . 0  92  33  -  -  -  33  13.0 ± 3 . 5  " Combined mortality rate of male and female caribou, as their mortality rates were not significantly different (  2 x  =0.02, df= l,/? = 0.90).  Causes, seasons and elevations of mountain caribou mortalities Mountain caribou i n the four study areas died from a variety o f factors. The proportion o f w o l f to non-wolf caribou deaths differed between the highland area, Quesnel Lake and the Columbia Mountains (Fisher's exact test, p = 0.0002). W o l f predation was the main mortality factor at Quesnel Lake, accounting for 10 (56%) o f the known deaths, but was a minor factor i n the other three study areas (Table 2). Bear predation was the most prominent mortality factor i n Wells Gray Park, causing three (75%) o f the known deaths. Avalanches were the greatest single factor contributing to caribou deaths in the Columbia Mountains, causing six (35%) known mortalities. Mountain caribou i n the Quesnel Highlands, though, died from a variety o f predation (43%) and non-predation (57%) factors. The highest seasonal mortality rate occurred during spring, but most mountain caribou died during the summer/fall or late winter (Table 3). Thirteen (56.5%) caribou deaths at Quesnel Lake occurred during the summer/fall, eight o f which were due to w o l f predation. The summer/fall accounted for five (83.3%) caribou deaths i n Wells Gray Park, with bear predation causing three  11  T a b l e 2. C a u s e s o f radio-collared m o u n t a i n c a r i b o u mortalities w i t h i n study areas. Study Area  Predation  Non-predation  Unk.  Total mortalities  Wolf  Bear  Wolverine  Malnutrition  Avalanche  Accident  Unk.  -  2  3  1  6  3  2  3  20  1  3  -  -  -  2  6  10  3  2  1  2  -  5  23  1  1  1  2  -  5  12  4  4  8  7  2  15  61  Columbia Mountains 1981-85 & 1992-96 Wells Gray Prov. Park 1986-89 Quesnel Lake 1984-89 Quesnel Highlands 1993-96 Total  2 13  8  o f these. T e n ( 5 0 % ) deaths o c c u r r e d d u r i n g late w i n t e r i n the C o l u m b i a M o u n t a i n s w i t h avalanches c a u s i n g f i v e o f these and w o l v e r i n e p r e d a t i o n three. I n the Q u e s n e l H i g h l a n d s , f o u r ( 3 3 . 3 % ) deaths o c c u r r e d i n b o t h summer/fall and late w i n t e r , but n o clear seasonal m o r t a l i t y factor w a s o b s e r v e d . D e a t h s r e s u l t i n g f r o m accidents and m a l n u t r i t i o n d i d not e x h i b i t strong seasonal affinities i n a n y study area.  T a b l e 3. S e a s o n a l causes o f radio-collared m o u n t a i n c a r i b o u m o r t a l i t i e s across a l l study areas ( C o l u m b i a M o u n t a i n s 1981-85, 1992-96; W e l l s G r a y P r o v i n c i a l P a r k 1986-89; Q u e s n e l L a k e 1984-89; Q u e s n e l H i g h l a n d s 1993-96). L e n g t h o f season (days) i n brackets.  Predation  Season Spring (28) Summer/Fall (163) Early Winter (30) Late Winter (144)  Wolf  Bear  10  2 6  -  -  3  Non-predation  Wolverine  Malnutrition  Avalanche  Accident  4  1 1 1 1  1 1 6  2 2 1 2  Unk.  Total  3 7 2 3  8 29 5 19  Unk.  2 -  I n study areas w h e r e w o l f and bear p r e d a t i o n o c c u r r e d , m o s t c a r i b o u w e r e k i l l e d b y these predators at l o w elevations ( T a b l e 4). W o l v e r i n e p r e d a t i o n , h o w e v e r , o c c u r r e d at b o t h h i g h a n d l o w elevations. C a r i b o u w e r e most l i k e l y to b e k i l l e d b y avalanches w h i l e at h i g h altitudes, b u t  12  mortalities caused by malnutrition and accidents occurred with the greatest frequency at low elevations.  Table 4. Type and frequency o f mortalities occurring above or below the median seasonal elevation o f radio-collared mountain caribou across all study areas (Columbia Mountains 198185, 1992-96; Wells Gray Provincial Park 1986-89; Quesnel Lake 1984-89; Quesnel Highlands 1993-96). Elevation  Predation  Non-predation  Wolf  Bear  Wolverine  Above Median  2  1  2  Below Median  11  7  2  Malnutrition  4  Unk.  Total  1  4  18  1  7  38  Avalanche  Accident  Unk.  6  2  1  5  Mountain caribou and wolf densities A m o n g the four study areas, the highest estimated caribou density (11.4 caribou/100 km ) 2  occurred i n the highlands around Quesnel Lake in 1986 (Table 5), but i n 1989, the estimated density fell to 4.9 caribou/100 k m . This was within the range o f caribou densities (3.4 to 5.3 2  caribou/100 k m ) observed in the other highland (Quesnel Highlands) and mountainous 2  (Columbia Mountains; Wells Gray Provincial Park) study areas. The two highland study areas (Quesnel Lake; Quesnel Highlands) had the highest estimated w o l f densities and observed winter pack sizes, but w o l f density was not estimated for Wells Gray Park as the number o f wolves/pack was unknown (Table 6). Wells Gray Provincial Park, the Columbia Mountains and the Quesnel Lake study areas each had two known w o l f packs, while the Quesnel Highlands study area had four.  13  Table 5. Mountain caribou density estimates based on mean annual distribution o f populations. Study  Columbia Mountains (1992-96) Wells Gray Prov. Park (1986-89) Quesnel Lake (1984-89) Quesnel Highlands (1993-96)  Year of population estimate  Number of caribou (95% C.I.)  Mean annual distribution (km )  1994  375 (337 -413)  7018  5.3  1987 1989 1986 1989 1997  275 302 220 94 358  8206  3.4 3.7 11.4 4.9 3.8  (220 - 297) (258 - 346) (197 - 243) (86 - 102) (294 - 422)  Density (caribou/100 km ) 2  2  It  1931 It  9398  Table 6. M e a n w o l f density estimates within pack territories. Study  Columbia Mountains (1992-96) Wells Gray Prov. Park (1986-89) Quesnel Lake (1984-89) Quesnel Highlands (1993-96)  Number of known wolf packs/study area  Number of wolves/radiocollared pack (winter)  Mean density (wolves/100 km of pack territory)  2  3-4  0.61  2  Unknown  Unknown  2  6-8  0.84  4  5-8  0.81  2  DISCUSSION Variation in wolf predation among highland and mountainous study areas Mortality data from the four study areas showed that mountain caribou at Quesnel Lake had a much higher mortality rate than i n the other three study areas, as the caribou density dropped from 11.4 to 4.9 caribou/100 k m between 1986 and 1989. The results support the hypothesis that w o l f predation is greater in highland areas than i n mountainous areas as suggested by Seip (1992a). W o l f predation was the main cause o f caribou deaths at Quesnel Lake (Seip 1992a, 1992b) but heavy w o l f predation was not reported in any other study area. N o n - w o l f mortality  14  factors were more common in rugged mountainous terrain. It appeared that bears were the major predator o f caribou i n Wells Gray Park (Seip 1990). Avalanches were the single most important mortality factor o f caribou i n the Columbia Mountains, but both wolverine and bear predation were also notable ( B . N . M c L e l l a n and J. Flaa, unpublished data; Simpson and Woods 1987). Another study, i n the southern Selkirk Mountains, found cougar predation was the cause o f most caribou mortalities (Compton et al. 1995).  Seip (1992a) attributed differences in w o l f predation on mountain caribou i n highland and mountainous areas to differences in the degree o f elevational and geographic separation between caribou and wolves. U p o n examination, elevational separation (Seip 1992a) does not appear to adequately explain the large variation i n w o l f predation among the four study areas. Except for summer/fall i n the Columbia Mountains, caribou in both highland and mountainous areas were found to maintain elevational and hence geographic separation from wolves (see chapter 3). The separation o f mountain caribou and wolves i n geographically different areas during summer/fall in the Columbia Mountains although not examined i n this thesis, may as reported elsewhere (Bergerud 1985; Bergerud et al. 1990; Cumming et al. 1996; Seip 1990, 1992a) explain w h y w o l f predation on caribou was not observed in this study area.  Two other factors may account for the variation i n w o l f predation on mountain caribou among the highland and mountainous study areas. Firstly, w o l f density differences (Bergerud and Elliot 1986) may explain why few caribou were killed by wolves i n the mountainous study areas (Columbia Mountains; Wells Gray Provincial Park), while caribou at Quesnel Lake frequently died from w o l f predation. Bergerud and Elliot (1986) reported that caribou populations decline in areas where w o l f densities exceed 0.65/100 k m . W o l f density within pack territory i n the 2  15  Columbia Mountains (0.61/100 k m ) did not reach this critical level and it appears unlikely that 2  wolves i n Wells Gray Provincial Park exceeded it either. Wolves in the highlands around Quesnel Lake, however, with more wolves per pack, and smaller territories than wolves i n the Columbia Mountains, had densities (0.84/100 k m ) above the critical threshold. This may have 2  lead to more w o l f predation on Quesnel Lake caribou. I f w o l f abundance is ultimately determined by the availability o f prey biomass (Boertje and Stephenson 1992; Dale et al. 1994; Hayes 1995; Messier 1985, 1994; Packard and M e c h 1980), and alternate ungulate prey remain sparse i n the wet, rugged mountains o f mountain caribou range (Terry et al. 1996), caribou in those areas should continue to be less susceptible to w o l f predation.  Secondly, the relative densities o f moose and caribou (Cumming et al. 1996; Seip and Cichowski 1996) may also be a factor influencing the susceptibility o f mountain caribou to w o l f predation. Optimal foraging theory (Krebs and M c C l e e r y 1984) predicts that wolves should hunt in the most optimal areas available to them, selecting the most energetically profitable prey (MacArthur and Pianka 1966). Assuming wolves i n the four study areas hunted optimally, one can infer from the indirect evidence (wolf scat contents and/or elevations used by wolves) that at the prey densities wolves encountered, moose were considered the more profitable prey. I f the density o f moose decreased and/or caribou density increased significantly, wolves would be expected to switch predation efforts (functional response) from moose to caribou (Farnell et al. 1996). This behaviour would continue until once again it became more profitable to prey on moose (Messier 1995). Perhaps this explains why caribou numbers plunged at Quesnel Lake between 1986 and 1989 (Seip 1992a), but have since stabilized (J.A. Y o u n g , personal communication), even though w o l f density remained high. The relatively high caribou density (11.4/100 k m ) i n 1986 may have been sufficient to alter wolves from preying largely on moose  16  to also prey on caribou as this density is above the equilibrium density o f 3.0-4.0 mountain caribou/100 k m  2  as suggested by Seip and Cichowski (1996).  Seasons and elevations of mountain caribou mortality factors It has been reported that mountain caribou are most vulnerable to wolf, bear, and cougar predation during summer (Compton et al. 1995; Seip 1990; Seip 1992b) especially at lower elevations (Seip 1992a). Mortality data from the Quesnel Highlands and Columbia Mountains studies support these findings. Unlike other carnivores, wolverines killed radio-collared caribou only during the late winter at both valley bottom and subalpine elevations ( B . N . M c L e l l a n and J. Flaa, unpublished data; J . A . Young, unpublished data). Mountain caribou may be most susceptible to predation by this relatively small carnivore in late winter because the caribou have depleted most o f their fat reserves and may be i n poor condition. Malnutrition deaths did not exhibit this biological pattern in the four areas examined, but this is likely due to a relatively small number o f mountain caribou mortalities.  17  CHAPTER 3 ELEVATIONAL RELATIONSHIPS O F MOUNTAIN CARIBOU, W O L V E S A N D MOOSE INTRODUCTION  Wolves that prey primarily on moose, but sometimes switch to woodland caribou, have been implicated as a major cause o f both past and present declines i n mountain caribou populations (Bergerud 1974, 1988a, 1996; Seip 1991, 1992b, Seip and Cichowski 1996). It was suggested that woodland caribou could only exist on ranges shared with high densities o f moose or deer and wolves i f a habitat feature allowed caribou to avoid wolves and their alternate ungulate prey (Bergerud 1974). Elevation appears to provide caribou with a unique means to space away from wolves and their alternate ungulate prey. In northern B . C . , the northern ecotype o f woodland caribou used rugged mountainous terrain and intra-specific dispersion to provide spatial separation from wolves and moose (Bergerud et al. 1984; Bergerud and Page 1987). In the Quesnel Lake area o f east central B . C . , wolves were sustained primarily by moose (Seip 1992b), but during the summer, their distribution overlapped with mountain caribou and they appeared to k i l l enough caribou to cause a rapid decline (Seip 1992a). Meanwhile, caribou that were spatially separated from moose and wolves in the nearby rugged mountains o f Wells Gray Provincial Park experienced little w o l f predation (Seip 1990). Seip (1992a) proposed that caribou living i n highland habitats were more available and thus more prone to w o l f predation than caribou in more mountainous terrain where they were separated from wolves. The spatial separation model (Seip 1992a) o f which elevation is a component, may apply throughout mountain caribou range. The objective o f this chapter is to test the following hypotheses: 1) wolves show more elevational overlap with moose than with mountain caribou, and 2) mountain caribou in highland 18  areas show less elevational separation from wolves than those living in rugged mountains.  METHODS Capture and elevational monitoring of mountain caribou, wolves and moose Radio telemetry data on mountain caribou and wolves from two study areas (Columbia Mountains 1992-96; Quesnel Highlands 1993-96), and moose radio-telemetry data from one study area (Columbia Mountains 1984-85), were used to evaluate the two hypotheses. Seip (1992a) did not statistically test the mountain caribou and w o l f radio-telemetry data (Wells Gray Provincial Park 1986-89; Quesnel Lake 1984-89) nor the moose radio-telemetry data (Quesnel Lake 1984-89) so these data were also examined. Captured animals were fitted with radio collars and relocated from aircraft two to four times each month. Ground telemetry was also used to locate wolves i n winter during the 1992-96 Columbia Mountains study. The location and elevation were recorded for each relocation.  Statistical Analyses Elevation o f radio locations were often asymmetrically distributed, thus two-tailed robust rankorder tests (Siegel and Castellan 1988) were used to test whether wolves and caribou, and wolves and moose used different elevations during each season i n the highland and mountainous study areas. Frequency o f use analyses were not used due to inadequate sample sizes i n each study area during some seasons. Seasonal intermedian distances o f wolves and caribou, and wolves and moose i n highland and mountainous areas were also compared as another measure o f the hypotheses. Seasons were based on elevational movements o f caribou ( M c L e l l a n et al. 1994a) and were classified as spring, summer/fall, early winter and late winter (see chapter 2). The level o f significance for all statistical tests was set atp < 0.05.  19  RESULTS Seasonal use of elevations by mountain caribou, wolves and moose In spring, caribou i n the highland (Quesnel Lake; Quesnel Highlands) and mountainous (Columbia Mountains; Wells Gray Provincial Park) study areas used lower elevations than during late winter, however, they remained separate from wolves which occupied still lower elevations (allp < 0.001) (Figures 2,3,4 and 5). In contrast, no difference was observed in the A  elevations used by wolves and moose in the Columbia Mountains (U [19,10] = 1.69, p = 0.091) A  or in the highlands around Quesnel Lake (17 [6,14] = 0.608, p = 0.543) during spring (Figs. 2 and 4). During summer/fall, wolves i n all study areas increased their use o f higher elevations (Figs. 2,3,4 and 5), particularly i n the rugged Columbia Mountains where elevations used b y wolves and caribou did not differ (U [38,1314] = 1.46,p = 0.143), nor did those used b y wolves and A  moose (U [38,230] = 0.396,p = 0.692) (Fig. 2). Wolves and caribou during summer/fall i n Wells Gray Provincial Park, and i n both highland areas (Quesnel Lake; Quesnel Highlands) remained elevationally separate (all p < 0.001), although wolves at Quesnel Lake showed some use o f high elevations during June (Fig. 4). In the Quesnel Lake area, moose and wolves used different elevations i n the summer/fall (U [92,64] = -2.34,p = 0.019) with moose using higher elevations (Fig. 4). During early winter, wolves and caribou in highland and mountainous areas again used lower elevations, but still little altitudinal overlap occurred (Figs. 2,3,4 and 5) (p < 0.01 i n all areas). Alternately, wolves and moose were found at similar low elevations i n both the Columbia A  A  Mountains (U [15,6] = 0.134,;? = 0.893) (Fig. 2) and the Quesnel Lake study areas (U [15,15]  20  •—Caribou —•—Wolf —*—Moose • • 4  238 13 19  224 17 34  351 13 17  211 6 1  412 21 10  316 10 21  195 7 60  261 8 64  165 6 41  219 3 39  271 9 11  322 18 0  2100 i  Late Winter  I Spring I  | V | Ean  Summer/Fall  w  i  n  t  e  r  Month  Fig. 2. Columbia Mountains radio-collared mountain caribou, wolf and moose median monthly elevations and interquartile ranges (1992-96 caribou and wolf; 1984-85 moose).  -•-Caribou -•—Wolf •  113  Jan  85  Feb  127  Mar Late  Winter  122  Apr  123  May  122  Jun  141  Jul  | Spring |  159  Aug Summer/Fall  105  Sep  110  125  Oct  63  Nov |  Dec  Early w  i  n  t  e  | r  Month  Fig. 3. Wells Gray Provincial Park radio-collared mountain caribou and wolf median monthly elevations and interquartile ranges (1986-89).  Caribou 110  78  197  •  14  12  7  4  18  37  H  46  190  193  7  Wolf —*— Moose 221  191  150  120  112  110  13  27  18  16  16  13  9  20  11  10  7  31  Aug  Sep  Oct  Nov  23  14  15  Apr  May  Jun  108  2100 1900 1700 1500 g »-—• >  m  LU  1300 1100 900 700 500  Jan  Feb  Mar  Jul  Spring  Late Winter  |  Summer/Fall  Month  Early Winter  Dec |  Fig. 4. Quesnel Lake radio-collared mountain caribou, wolf and moose median monthly elevations and interquartile ranges (1984-89). Caribou •  83  48  123  31  48  •  12  14  19  7  12  Feb  Mar Late Winter  May  Wolf  37  48  Jul  25  36  40  142  184  6  12  7  25  22  Aug Summer/Fall  Spring  Month  Sep  Nov  Dec  Early Winter  Fig. 5. Quesnel Highlands radio-collared mountain caribou and wolf median monthly elevations and interquartile ranges (1993-96). 22  = 1.14,/? = 0.255) (Fig. 4) during the early winter. During late winter, the elevations used by wolves i n the highlands and mountains were generally lowest (Figs. 2,3,4 and 5) with wolves separate from caribou i n each study area (all/? < 0.001). Wolves and moose in both the Columbia Mountains and the Quesnel Lake area used lower elevations than caribou during the late winter (Fig. 2 and 4), yet the elevations used by wolves and moose i n the Columbia Mountains did not differ (U [59,71] = 0.113,/? = 0.91), A  whereas those o f wolves and moose at Quesnel Lake did (U [47,147] = 2.72, /? = 0.007).  Seasonal intermedian distances of mountain caribou, wolves and moose In all seasons, in both the rugged Columbia Mountains and the highlands around Quesnel Lake, the median elevation o f wolves was more closely associated with the median elevation o f moose than that o f caribou (Table 7). Seasonal intermedian distances o f wolves and moose ranged from 46 to 305 m i n the Columbia Mountains, while those o f wolves and caribou ranged between 198 and 975 m . A t Quesnel Lake, the wolf/moose seasonal intermedian distance ranged between 35 and 213 m , compared to a range o f 347 to 747 m for wolves and caribou. Caribou and wolves i n highland areas did not always have seasonal intermedian distances lower than those o f caribou and wolves in the mountainous study areas. Only during spring and late winter was the Quesnel Lake (highland area) wolf/caribou intermedian distance less than that o f wolves and caribou in Wells Gray Provincial Park (Table 7). A similar comparison between wolves and caribou i n the Quesnel Highlands and the Columbia Mountains showed that only i n late winter did wolves and caribou i n the Quesnel Highlands have a seasonal intermedian distance less than that o f wolves and caribou in the rugged Columbia Mountains (Table 7).  23  Table 7. Wolf/mountain caribou and wolf/moose seasonal intermedian distances (m) within study areas. Season  Mountains Columbia Mountains  Spring Summer/Fall Early Winter Late Winter  Highlands Wells Gray Prov. Park  Quesnel Lake  Quesnel Highlands  wolf-moose  wolf-caribou  wolf-caribou  wolf-moose  wolf-caribou  wolf-caribou  305 76 46 61  579 198 366 975  600 400 200 850  35 213 68 137  347 510 492 747  778 746 427 736  DISCUSSION The hypothesis that wolves show more elevational overlap with moose than with mountain caribou was supported i n both highland and rugged mountains. W o l f scat analyses from these two areas further support these conclusions as the main diet item o f wolves throughout the year in both the Quesnel Lake area (Seip 1992b) and the Columbia Mountains (see chapter 4) was moose. Similar elevational relationships between wolves and moose were reported i n northern British Columbia during the summer (Bergerud et al. 1984; Bergerud and Page 1987) and throughout the year in Wells Gray Provincial Park and the Quesnel Lake area (Seip 1992a). The hypothesis that mountain caribou i n highland areas show less elevational separation from wolves than those living i n rugged mountains was not clearly supported. Caribou and wolves in the highlands (Quesnel Lake; Quesnel Highlands) were as likely to use different elevations during each season as were caribou and wolves i n rugged mountainous areas (Columbia Mountains; Wells Gray Provincial Park). Even during summer/fall, when wolves were most elevationally mobile and likely to prey upon mountain caribou (Seip 1992a), caribou i n the rolling highland areas appeared to space further away from wolves than did caribou i n the more mountainous areas. Despite the advantage o f greater elevational range to space away from wolves, mountain caribou i n the Columbia Mountains were the only population to exhibit  24  significant elevational overlap with wolves, and then, only during summer/fall. The Columbia Mountains caribou population has not declined and no w o l f predation o f radio-collared caribou has been observed to date ( B . N . M c L e l l a n , personal communication). This may indicate that caribou and wolves occupied geographically different areas during summer/fall, similar to caribou and wolves i n Wells Gray Park (Seip 1990, 1992a). Wolves were reported as the main cause o f caribou deaths i n the highlands near Quesnel Lake (Seip 1992a); however, the elevations used by the two species differed during all four seasons. A brief overlap i n elevational use by wolves and caribou may have occurred during the calving period (Bergerud et al. 1984; Bergerud and Elliot 1986; Bergerud and Page 1987) and influenced the predation rate. This does not seem likely though as few adults at Quesnel Lake were killed by wolves during this period (Seip 1992b) and w o l f predation on calves was reported to be low in June (Seip 1992a). It appears that without significant elevational overlap, wolves caused a dramatic decline i n the caribou population at Quesnel Lake as 80% (8/10) o f the w o l f predation occurred at elevations below the median elevations o f radio-collared caribou (see chapter 2). M y results suggest that elevational separation is inadequate i n explaining the large variation i n mountain caribou deaths attributed to w o l f predation i n the highland and mountainous study areas examined (see chapter 2). Differences in w o l f densities (Bergerud and Elliot 1986) and the relative densities o f moose and caribou (Cumming et al. 1996; Seip and Cichowski 1996) may be the primary factors influencing the susceptibility o f mountain caribou to w o l f predation within highland and mountainous areas i n any given season (see chapter 2). It would be useful i f future studies were conducted in both highland and mountainous areas to examine how different mountain caribou, moose and w o l f densities influence the elevational relationships o f these species. Additionally, where elevational separation between caribou and wolves does not occur, the geographic locations o f caribou and wolves should be compared to determine i f any spatial  25  separation exists. These studies should also be o f longer duration than those conducted thus far i n order to ascertain the seasonal and annual variability within mountain caribou, w o l f and moose spatial interactions.  26  CHAPTER 4 T H E SEASONAL DIETARY IMPORTANCE OF MOOSE T O WOLVES IN T H E COLUMBIA MOUNTAINS INTRODUCTION Early this century, the distribution and abundance o f moose increased rapidly i n parts o f British Columbia (Edwards 1954; Hatter 1949; Spalding 1990). This was possibly due to the clearing and burning o f forests for land settlement and railway development (Hatter 1949). Bergerud (1974) and Bergerud and Elliot (1986) suggested that the increased availability o f moose stimulated an increase i n wolves that also preyed upon and reduced or eliminated caribou populations lacking an adequate antipredator strategy. W i t h an ample alternate ungulate prey source available, a decrease i n caribou would not necessitate reduced w o l f numbers (Bergerud 1988b; Seip 1992b). Wolves i n the Quesnel Lake area were reported to be sustained primarily by moose (Seip 1992b); however, studies o f the importance o f moose i n w o l f diet i n other areas o f mountain caribou range are scarce. The objective o f this chapter is to test the hypothesis that wolves i n mountain caribou range are sustained primarily by moose throughout the year.  METHODS The seasonal importance o f moose for two packs o f radio-collared wolves i n the Columbia Mountains was estimated from 79 w o l f scats collected between December 1993 and November 1995. Scats were classified as either summer ( M a y 1 - N o v 7) or winter (Nov 8 - A p r 30). In summer, scats were collected at den and rendezvous sites, and also monthly along an extensive network o f logging roads. During winter, scats were collected at kill/scavenge sites, along roads, and while backtracking wolves i n snow. Logging roads i n the study area were checked just after snowmelt i n the spring for winter-deposited scats (Forbes and Theberge 1992). Since coyotes 27  (Canis latrans) inhabited this study area, only scats > 30 m m i n diameter (Weaver and Fritts 1979) or smaller scats associated with a w o l f den, rendezvous site or w o l f tracks were collected (Fuller and Keith 1980). Hair from scats was identified to species using the methods o f Adorjan and Kolenosky (1969), Kennedy and Carbyn (1981), and Spilborghs (1996). White-tailed and mule deer hair could not be separated. Frequently, multiple scats collected at the same k i l l or rendezvous site contain hair from the same k i l l and are not independent samples (Huggard 1991). In an attempt to minimize this bias, the effective sample size for these sites was taken as the number o f prey species represented i n the scats. The regression equation proposed by Weaver (1993) was used to estimate the relative weight o f each prey species consumed by wolves, based on the frequency o f occurrence o f each prey species i n scats. Paired t-tests ( S Y S T A T 1992) were used to determine whether moose was the primary diet item o f wolves i n each season. In testing the w o l f diet hypothesis statistically, "primary diet item" was defined as being >50% o f the biomass consumed by wolves. Levels o f significance were set &tp < 0.05.  RESULTS Frequency of occurrence of wolf diet items Moose occurred approximately twice as often i n winter scats (50%) than i n summer scats (22.0%), whereas the frequency o f caribou remains i n w o l f scats differed little between summer and winter seasons, with each containing 14.0% and 18.2% respectively (Table 8). Deer remains were found only i n winter w o l f scats and then, occurred infrequently (6.8%). Beaver, a common food item o f wolves i n the Columbia Mountains i n the summer, comprised 42.0% o f the total diet. Although several other species were found i n both summer and winter w o l f scats, each occurred infrequently (Table 8).  28  Table 8. Frequency o f occurrence and percent biomass o f w o l f diet items by season i n the Columbia Mountains. Summer  Wolf diet items  Winter  Frequency of occurrence  % Biomass  Frequency of occurrence  % Biomass  7 7 4  18.9 42.0 5.7  8 22  12.7 77.5  Caribou Moose (adult) Moose (calf) Deer Beaver Other*  -  -  -  -  3  3.4  21 11  21.8 11.6  -  -  11  6.4  No. of scats No. of items  44 50  35 44  Note: Percent biomass is based on the regression equation: y = 0.439 + 0.008x, where x is animal mass (Weaver 1993). Animal masses used were as follows: caribou, 124.7 kg (based on a ratio of 22 bulls : 78 cows : 21 calves (McLellan et al 1994b) and D.C. Thomas, unpublished data); adult moose, 343.4 kg (J. Woods, Banff NP unpublished data); moose calf in July, 40.0 kg (Banfield 1974); adult deer (mule and white-tailed combined), 74.2 kg (J. Woods, Banff NP unpublished data); beaver, 14.0 kg (Leege & Williams 1967). *Includes wolf, hoary marmot, red-backed vole, red squirrel, grouse, snowshoe hare, abattoir scraps, and deer mouse.  Percent biomass of wolf diet items During winter, wolves i n the Columbia Mountains were sustained primarily by moose (t-test = 4.12, d f = 34, p < 0.001), with moose clearly the most important species (77.5%; Table 8). A l l five w o l f kills found while backtracking wolves during the winter o f 1994-95 were moose, confirming this result. Caribou, the second most prominent diet item, contributed 12.7% o f the biomass consumed by wolves. Deer, the only other ungulate found i n winter scats, constituted a mere 3.4% o f w o l f diet during this season. Wolves i n the Columbia Mountains were not sustained primarily by moose i n the summer (ttest = -0.25, d f = 43, p = 0.403), yet moose was still the most important species i n the summer diet. Three prey items accounted for 88.4% o f the biomass consumed. In order o f importance these were: moose (47.7%; 42% adult, 5.7% calves), beaver (21.8%) and caribou (18.9%).  29  DISCUSSION Bergerud (1974) and Bergerud and Elliot (1986) suggested that in areas with both moose and caribou, wolves would prey primarily on moose. The estimate o f winter w o l f diet items in the Columbia Mountains supported this hypothesis, but the estimate o f summer w o l f diet items did not. Still, moose was the main diet item o f Columbia Mountains wolves throughout the year and as reported at Quesnel Lake (Seip 1992b), moose were clearly o f greater dietary importance to wolves than caribou. Unfortunately, moose census data corrected for sightability were not available for the Columbia Mountains, so testing whether wolves used moose and caribou equal to their availability was not possible. Uncorrected moose census data did indicate that moose were common i n the Columbia Mountains during the study (J. Krebs, personal communication). Wolves i n the Columbia Mountains relied to a greater extent on beaver during the summer, and less on moose than wolves i n the Quesnel Lake area (Seip 1992b). Differences i n the relative availability o f beaver (Theberge et al 1978) in the two areas may explain this variation, as wolves often prey heavily on beaver when available (Peterson 1977; Potvin et al. 1992; Voigt et al. 1976). It is also possible that the high occurrence o f beaver in the sample o f summer w o l f scats is an artifact o f collecting a disproportionate number (95%) o f w o l f scats at lower elevations during this season. During summer, radio-collared wolves i n the Columbia Mountains were located 2 5 % o f the time i n roadless, high-elevation areas, and often these areas were not visited as the probability o f finding w o l f scats at or near the bi-monthly relocation site o f a radio-collared w o l f was low. Assuming wolves consumed diet items at elevations proportionally equal to those o f their relocations, and that scats would be left i n proximity to k i l l or scavenge sites (Huggard 1991), the summer w o l f scat sample may have been biased towards lower elevation prey species (i.e., beaver), and against higher elevation prey species (i.e., caribou and hoary marmots). Therefore, •  30  caution should be used when interpreting the estimates o f w o l f summer diet i n the Columbia Mountains. This estimate o f w o l f diet in the Columbia Mountains and the Quesnel Lake w o l f diet estimate (Seip 1992b) support the view that wolves inhabiting mountain caribou range are largely sustained by moose. Both studies, however, lack sufficient data to test for annual and pack level variations i n w o l f diet across a range o f moose and caribou densities. Future mountain caribou studies i n which wolves are also radio-collared, should seek to address this inadequacy.  31  CHAPTER 5 GENERAL CONCLUSION AND MANAGEMENT RECOMMENDATIONS Wolves were suggested to be the primary factor limiting mountain caribou although they were sustained largely by moose (Bergerud and Elliot 1986; Seip 1992a). Mountain caribou i n areas o f highland topography were suggested to be most susceptible to w o l f predation as, unlike those i n more rugged mountains, they had less ability to space away from wolves and moose (Seip 1992a). The importance o f moose in the seasonal diet o f wolves was one element o f the spatial separation model (Seip 1992a) investigated. W o l f scat contents from the Columbia Mountains supported the hypothesis that wolves i n mountain caribou range are sustained primarily by moose. This was particularly evident during winter when few alternate prey species were available to wolves. Similar to wolves at Quesnel Lake (Seip 1992b), mountain caribou were a minor part o f Columbia Mountains w o l f diet. Since moose appear to be the main prey o f wolves in southern B . C . and w o l f abundance may be strongly associated with the density o f moose, attempts should be made to maintain moose densities at their present levels, particularly i n highland areas. The importance o f w o l f predation as a mountain caribou mortality factor was another element o f Seip's (1992a) model that was examined. The caribou mortality data indicated that w o l f predation on mountain caribou was higher at Quesnel Lake than i n the Columbia Mountains, supporting the hypothesis that w o l f predation on mountain caribou is greater i n highland areas than i n mountainous areas. There was no evidence though indicating that caribou i n highland areas continually experience high levels o f w o l f predation. Possibly wolves prey on mountain caribou i n a density-dependent manner (Seip and Cichowski 1996). Mountain caribou i n rugged mountains experience limited w o l f predation while avalanches, bear, wolverine, and cougar 32  predation appear as more common mortality factors. Acquiring a more comprehensive understanding o f the importance o f w o l f predation on mountain caribou populations in both highland and mountainous areas w i l l require more caribou mortality data than are presently available. Radio-collaring more caribou i n and o f itself, w i l l not necessarily increase the sample size o f caribou deaths with known causes. Increasing the frequency o f successive relocation flights (Heisey and Fuller 1985) and reducing response times to caribou mortality sites should help in differentiating among possible mortality factors, lessen the number o f caribou deaths o f unknown cause and increase sample sizes required to further test the w o l f predation hypothesis. The remaining element o f the spatial separation model (Seip 1992a) investigated was the degree o f elevational separation between wolves and moose, and wolves and mountain caribou in highland and mountainous areas. The radio-telemetry data supported the hypothesis that wolves show more elevational overlap with moose than with mountain caribou. N o support, however, was demonstrated for the hypothesis that mountain caribou i n highland areas show less elevational separation from wolves than did caribou i n rugged mountains. Differences i n w o l f density (Bergerud and Elliot 1986) and the relative densities o f moose and caribou (Cumming et al. 1996; Seip and Cichowski 1996) may explain the large variation i n the number o f mountain caribou mortalities attributed to w o l f predation i n the highland and mountainous study areas examined to date. Although the wolf/caribou radio-telemetry data used in this thesis incorporates all the current spatial data simultaneously collected on these species, it explains the elevational relationships o f wolves and caribou within a relatively short period o f time. To determine the annual and seasonal variation i n the elevational interactions o f mountain caribou, wolves, and moose it is recommended that future studies addressing this issue be o f longer duration. Further, to fully test  33  the spatial separation model (Seip 1992a), the elevational and geographic distributions o f mountain caribou, wolves and moose should both be examined. Additionally, the degree to which habitat alteration (timber harvesting and fires) influences wolf, mountain caribou, and moose densities and ultimately, their interactions is still largely unknown. Research directed towards understanding how wolves and their prey respond numerically and spatially to different landscape changes i n both highland and rugged mountainous areas should be a primary component o f future wolf/caribou/moose radio-telemetry studies.  34  LITERATURE CITED Adorjan, A . S . , and Kolenosky, G . B . 1969. A manual for the identification o f hairs o f selected Ontario mammals. Ont. Dept. o f Lands and Forests. Toronto, Ont. Anderson, R . M . 1938. The present status and distribution o f the big game mammals o f Canada. Transactions o f the 3rd North American W i l d l . 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