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The Vancouver Island wolf (Canis lupus crassodon) : an initial study of food habits and social organization Scott, Barbara Mary Victoria 1979

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THE VANCOUVER ISLAND WOLF  (Canis lupus crassodon),  [ INITIAL STUDY OF FOOD HABITS AND SOCIAL ORGANIZATION  by  BARBARA MARY VICTORIA SCOTT B.Sc., The University of British Columbia, 1976  A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Department of Animal Science  We accept this, thesi .v, as conforming t^K thfi//ip^nuire^L/^tandarc^  THE UNIVERSITY OF BRITISH COLUMBIA September 1979 ©  Barbara Mary Victoria Scott, 1979  In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives.  It is understood that copying or publication  of this thesis for financial gain shall not be allowed without my written permission.  Department of  VUUMftAi  )C  The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5  ABSTRACT  The food habits and social organization of Vancouver Island wolves  (Canis lupus crassodon Hall) inhabi-  ting an area on Northeastern Vancouver Island were from January, 1977 through January, 1979.  studied  During this  period, two adjacent packs were studied in detail between March and November, 1978. Three members of the Upper Adam pack, an adult male, adult female and yearling female, were fitted with r a d i o - t r a n s m i t t e r s tracked for five months  captured,  and subsequently  (April to August, 1978).  radioThe same  procedures were carried out on the adult male and female of the Lower Adam pack, who were and five months respectively August). months  radio-tracked for eight  (April to November; April to  A lone male was captured and tracked for eight  (November, 1978 to June, 1979). Black-tailed deer  (Odocoileus hemionus  colum-  bianus) were the major prey in the diet of both packs, while Roosevelt elk (Cervus elaphus roosevelti) and beaver  (Castor canadensis leucodontus) were utilized as  secondary food sources.  Seasonal variation in the diet  of the wolves was apparent, with deer fawns and elk calves constituting the bulk of the summer (June 1 to August 31) diet. winter months.  Beaver were taken primarily during the i  Individual packs showed different  foraging patterns.  The lower pack depended less upon adult elk during the winter and more upon ungulate young during the summer, while the Upper pack utilized relatively more adult elk during the winter with more emphasis on adult deer as opposed to ungulate young during the summer. The Upper pack consisted of ten individuals, two adult males, one adult female, one yearling femaling, two unknown, and four pups. bited a 64 km 2  The radio-collared members  home range.  inha-  Wolf density within their  home range was one per 6.4 k m 2 . The Lower pack consisted of five  individuals  including one adult male, one adult female, and three pups which ranged throughout a 75 km 2  area.  Density  within this area was one wolf per 15 k m 2 . Den sites were situated within pristine  coniferous  timber where hollow logs, tree bases and root systems were utilized for denning purposes.  Both packs occupied  from late April until mid-July.  densites  During the post-denning  period rendezvous sites were frequented, situated in open meadows bordering on timber stands or river side areas. Both types of sites were generally typified by a water source nearby, structural suitability giving a view of the surrounding area, activity and resting areas, and several well used trails.  i  i  Seasonal shifts in the use of core areas of home ranges were apparent for both packs, and den and rendezous sites were spatio-temporally distributed at significant distances from adjacent pack sites.  iii  TABLE OF CONTENTS  Page 1  ABSTRACT  V1  LIST OF TABLES LIST OF FIGURES  vii  ACKNOWLEDGEMENTS  XII  CHAPTER 1  INTRODUCTION  1  CHAPTER 2  THE STUDY AREA  4  CHAPTER 3  FOOD HABITS  10  3.1  Introduction  10  3.2  Methods  14  3.3  Results  16  3.4  Discussion  24  SOCIAL ORGANIZATION  30  4.1  Introduction  30  4.2  Methods  32  CHAPTER 4  4.3  4.2.1  Capture Procedures  32  4.2.2  Radio-Location Methods  34  4.2.3  Den and Rendezvous Sites  36  Results 4.3.1  *  37  Weights, Measurements and Metabolic Profiles  iv  37  Page  4.4 CHAPTER 5  4.3.2  Pack Sizes and Composition  4.3.3  Home Range Sizes and Dis-  37  ., • tributions  4S  4.3.4  Wolf Densities  72  4.3.5  Daily and Seasonal Movements  4.3.6  Movements of a Lone Wolf  83  4.3.7  Den Site Characteristics  87  4.3.8  Rendezvous Site Characteristics.  92  ...  72  94  Discussion FINAL DISCUSSION  102 109  LITERATURE CITED  v  LIST OF TABLES Page  Table 1  Major mammal species  inhabiting  the study area Table 2  Metereological data from two stations in the area surrounding the study site  Table 3  The diet of Vancouver Island wolves (Canis lupus crassodon) determined by analysis of scats from two packs collected between February  and 17  November, 1979 Table 4  Diets of the Upper and Lower Adam wolf packs, expressed  in terms of the  relative weights of each prey in the scats Table 5  species  (after Floyd et al., 1978). 23  Summary of results from previous  studies  of the social organization of wolves  in  North America Table 6  Weights  31  (kg) and measurements  nine Vancouver Island wolves  (mm) of (Canis  lupus erassodon) captured between April vi  10 and November 1, 1978, near Kelsey Bay, British Columbia Table 7  Haematology of some Vancouver  38 Island  wolves Table 8  39  Serum chemistries of Vancouver  Island  wolves Table 9  40  Serum proteins and hormones of Vancouver Island wolves  Table 10  41  Individual and pack home range sizes (km 2 ) for the whole study period combined for two packs of Vancouver  Island  wolves Table 11  52  Individual and pack home range sizes (km 2 ) shown for denning  (April 11 to  July 11) and post-denning periods of two packs of Vancouver Island wolves ..  vii  53  LIST OF  FIGURES Page  Figure 1  The study site  Figure 2  Seasonal changes in the diets of wolves obtained from combined samples from both packs, estimated by relative weight  Figure 3  (A) and relative occurrence  (B).  18  Diets of wolves determined from scats collected at den (D-S) and rendezvous 21  (R-S) sites Figure 4  Observation-area curves for Lower Adam adult male's  (86) and adult  female's  (90) 100 and 90 per cent home ranges . Figure 5  Observation-area curves for Upper Adam adult male's  (87), adult female's  and yearling female's  (97)  (4) 100 and 90 49  per cent home ranges Figure 6  47  100, 90 and 50 per cent home ranges of the adult male  (86) and the adult  female  (4) from the Lower Adam pack for the whole study period  (86: April 11 -  November 20; 90: April 14-August 29, 1978) 54 viii  Page  Figure 7  100, 90, and 50 per cent home ranges of the Lower Adam adult male during the denning periods  (D:  (D) and post-denning  (PD)  April - July; PD:  August - November, 1978) Figure 8  56  100, 90 and 50 per cent home  ranges  of the Lower Adam female during the denning  (D) and post-denning  periods.  (PD)  (D: April to July; PD:  August, 1978) Figure 9  Home ranges  58  (100, 90, and 50 per cent)  of Lower Adam  (LA) and Upper Adam  packs for whole study period  (UA)  super-  imposed on topographic map Figure 10  61  100, 90, and 50 per cent home ranges of the adult male  (87) , adult  (90) , and yearling female  (4), of the  Upper Adam pack for the whole period  female  study  (87: April 10-September 10;  97: April 30 - September 3; 4: May 4- September 3, 1978) ix  63  Page  Figure 11  100, 90 and 50 per cent home ranges of the Upper Adam adult male the denning periods  (D:  (87) for  (D) and post-denning  (PD)  April - July; PD:  August  - September, 1978) Figure 12  65  100, 90 and 50 per cent home for the Upper Adam female the denning periods  ranges  (97) during  (D) and post-denning  (D: April-July; PD:  (PD)  August -  September, 1978) Figure 13  67  100, 90 and 50 per cent home ranges of the Upper Adam yearling female for the denning periods  (4)  (D) and post-denning  (D: April - July; PD:  August -  September, 1978) Figure 14  69  100, 90 and 50 per cent home ranges of the Lower Adam pack  (LA) and the Upper  Adam pack for the whole study period (LA:  April 11 - November.20; UA:  April 10 - September 10 , 1978)  x  73  Page  Figure 15  100, 90, 50 per cent home ranges of the Lower Adam  (LA) and the Upper Adam  (UA) packs during the denning period (April - July, 1978) Figure 16  75  100, 90 and 50 per cent home ranges of the Lower Adam  (LA) and Upper Adam  packs during the post-denning (LA: August - November; UA:  (UA)  period August -  September, 1978) Figure 17  Movements of the adult male  77 (86) from  the Lower Adam pack during a 10-hour and 5-hour period Figure 18  Q "I  Movements of a lone adult male wolf (November 1, 1978 - June 1, 1979)  xi  84  ACKNOWLEDGEMENTS  It would not have been possible to gather and compile the information in this report without the generous support of several individuals, agencies, and institutions. David Shackleton, Assistant Professor, University of British Columbia, provided unending assistance and encouragement throughout all the phases of my work. Drs. Fred Bunnell, Ian McTaggart Cowan, Daryll Hebert, and Ray Peterson together with Dr. Shackleton orig inally appro  and encouraged Financial support was generously provided by  the National Research Council of Canada, University of British Columbia, British Columbia Fish and Wildlife Branch, British Columbia Parks Branch, British Columbia Resource Analysis Branch, Wild Canid Research and Survivial Centre, MacMillan Bloedel Company, the Mittlenatch Society and the World Wildlife Fund  (Canada).  Susan Shaw was my assistant during the major phase of the field work.  Not only did she help to  collect a good deal of the data, but together we spent many happy hours wandering through pristine forests searching for wolf activity sites and enjoying the beauty of the wilderness and its wildlife inhabitants. xii  David  Williamson gave generously of his time and knowledge to reduce these taslcs.  He shared a wealth of practical  knowledge with me and I will forever be grateful to him for helping to create a good "Mountain Lady".  Keith McKillican,  of Island Airlines, flew me to make most of the aerial observations and locate radioed wolves.  His ease in  handling the plane under adverse weather conditions, in narrow valley systems gave me utmost confidence in his ability.  Thanks are also extended to the other pilots and  staff of Island Airlines. Dr. Don Bowen offered many valuable  suggestions  during the early phases of my work, and Dr. Alton Harestad gave me invaluable help with data analysis.  Dr. L. David  Mech passed on much of his knowledge concerning capturing and radio-tracking wolves during my visit with him in Minnesota. Despite my father's perplexity when I announced I was going off to the woods to trap wolves, he was always encouraging and helped me to endure the stress that was sometimes a part of the research. Other individuals who contributed to the study include Holly Cleator, Donald Eastman,'Susan Fleck, Doug Janz, Deborah Lebrocq, Kevin Lloyd, Inu Nelnor, Jane Packard and Frank Roberts I deeply appreciate the support extended to me by all during the course of the research. xiii  ~  1.  INTRODUCTION  The wolf (Canis lupus) is a large canid widely distributed throughout the northern hemisphere.  From  the earliest times, it has held the interest of Homo sapiens and since the early 1940's has been the subject of numerous detailed studies both in North America  (Murie, 1944;  Mech, 1966; Pimlott el al., 1969; Carbyn, 1974; Haber, 1977) and Eurasia (Makridin, 1962; Pulliainen, 1965).  Within its  distribution the species is thought to be composed of 32 sub-species, 24 of these occurring in North America  (Mech,  1970) . The Vancouver Island wolf (Canis lupus crassodon Hall) was described in 1932, and designated as a subspecies unique to the Island from its apparent distinct pelage colour and carnassial teeth characteristics However, its taxonomic status is currently  (Hall, 1932). questionable,  as adequate cranial samples were not available to Hall (1932), Joliceuer  (1959) or Lawrence & Bossert  (1967) .  Although no accurate figures exist which deal with wolf numbers, it is believed that this race has gone through "violent fluctuations" and has been "virtually extirpated on either two or three occasions since the early 1920's" (Cowan, pers. comm.).  Since the early 1970's  there have been reports of an apparent increase in wolf numbers (Hebert, pers. comm.). Concurrently there has been increased access into remote areas of probable prime wolf habitat throughout the Island, to the extent that only one pristine watershed remains.  Coupled with this increased  access, an awareness of the wolf has developed in various segments of the human population.  Without accurate data  on the population dynamics of the wolf this apparent increase in numbers can only be speculative, and its management practically  impossible.  This study was the first on Vancouver Island to investigate wolf behaviour and ecology.  The major objec-  tives were to provide baseline information dealing primarily with the food habits and social organization of two adjacent wolf packs.  For the former objective the wolf's major  food sources, the relative proportions of occurrence in their diet, and seasonal variation in prey species and ageclass utilization were investigated.  Aspects of social  organization examined included wolf densities, pack distributions, pack sizes and compositions, home range sizes, den and rendezvous site characteristics and, daily and seasonal movements. As there was substantial information on the ecology of the wolf in other regions with contrasting or similar habitats and prey complexes, certain predictions were made  regarding the ecology of the Vancouver Island race.  It was  suggested that because elk, deer, and beaver are large mammalian prey species for wolves elsewhere, these species would also constitute the major portion of the diet for the Vancouver Island race.  Seasonal variation in utilization  of prey species and type was also predicted.  Specifically  that young ungulates would be used more frequently than adults early in the summer when they were vulnerable and abundant.  Finally, steep watersheds, a common topographical  feature in the study site, were expected to be influencial in limiting the activities of wolf packs to separate watersheds.  2.  The study site  STUDY AREA  (Figure 1) is situated on the  northeast portion of Vancouver Island, 115 km southeast of Port Hardy and 80 km north of Campbell River, British Columbia  (latitude 50°N, longitude 125° 15' W).  The  major watersheds are the Adam and Eve Rivers which flow in a north-northwesterly  and northeast-northwesterly  respectively through the area.  direction  They converge one kilometre  from the ocean and ultimately drain into Johnstone The study site encompasses approximately  Strait.  530 km 2 .  Access to the site is afforded by logging  roads.  A major highway to Port Hardy, opened in 1979, traverses the area along valley floors.  The terrain is characterized by  steep and narrow valley systems ranging in elevation from sea level to 2,158 m. Logging practices within the area have altered natural habitat types, resulting in clearcut and second growth zones in valley bottoms and side hills.  Pristine  habitat remains at river headwaters and upper reaches of side hills within watersheds.  Habitat types within these  areas include pristine forests, alpine, and high elevation wet meadows.  The study site includes three biogeoclimatic  zones - the subalpine mountain hemlock, coastal Douglas fir and coastal western hemlock zones 1969-70) .  (Krajina and Brooke,  Figure 1  Map of the study site  7.  Within the study site trees commonly form closed forest stands where mountain hemlock (Tsuga mertensiana), amabilis fir (Abies amabilis), yellow  (Chaemacyparis  nootkatensis) and red cedar (Thuja plicata), and western hemlock (Tsuga heterophylla) are the dominant Tree line occurs at 1373 m.  species.  The major faunal components  are shown in Table 1. Table 2 provides data on snowfall, precipitation and temperature recorded at sites near  the study area.  Precipitation and temperature data show general trends over a thirty year period.  Table 1.  Major Mammal Species Inhabiting the Study Area (From Cowan & Guiguet, 1975).  Wolf  Canis lupus crassodon  Cougar  Felis concolor vancouverensis  Wolverine  Gulo luscus vancouverensis  Black bear  Ursus americanus vancouveri  Marten  Martes americana caurina  Racoon  Procyon lotor vancouverensis  Mink  Mustela vison evagor  Otter  Lutra canadensis pacifica  Black-tailed deer  Odocoileus hemionus columbianus  Roosevelt elk  Cervus elaphus roosevelti  Beaver  Castor canadensis  Shrews  Sorex spp. (2)  Bats  Myotis spp. (4)  Marmot  Marmota caligata vancouverens is  Squirrel  Tamiasciurus hudsonicus  Mouse  Peromyscus maniculatus  Vole  Microtus townseudi laingi  Muskrat  Ondatra zibethica osoyoosensis  Weasel  Mustela erminea onguinae  Red Fox  Vulpes fulva  leucodontus  lanuginosus interdictus  9.  Table 2.  Chatham  Metereological Data from Two Stations in the Area Surrounding the Study Site. Data were Collected over the Period 1941 to 1970.  Point  Mean d a i l y temperature  Latitude  3 6  .5  SO 20 H  39.6  41.5  Longitude  45.0  125  53.2  26 ^  56.8  Ele.va.tlon  60.4  59.8  65 f,t.  55.4  48.7  42.1  38.4  CF")  Mean d a i l y maximum t e m p e r a t u r e  39.2  43.u  ts.y  Mean d a i l y minimum t e m p e r a t u r e  33.7  36.1  3/.1  No. o f days with f r o s t Mean r a i n f a l l Cinches)  9  4  8 - 5 9  6  4 .92  Q  6Q  Q  „ „ ^'-s  46  6 3 6  3  .3  6 6  s 3  .4  61.0  52.5  45.0  41.3  .i  49.7  44.8  39.2  35.5 7  4 9 - 9  5 3 - 4  0  0  0  0  *  2  *  O  6 7  6.04  4.68  3.66  3.16  2.97  3.63  5.02  10.91  9.70  11.69  3.7  12.0  Mean snowfall  14.6  4.0  3.5  0.3  0.0  0.0  0.0  0.0  0.0  0.0  Mean total ppt.  10.05  7.32  6.39  4.71  3.66  3.16  2.97  3.63  5.02  10.9110.0712.89  12  12  12  12  12  13  13  13  13  No. o f y r s . record  AlsDit  of  12  50 35 N  Latitude.  Bay  Longitude.  126  56 ill  Elevation  169  13  it.  13  ASL  temperature  37.4  40.6  41.4  45.4  50.7  54.6  57.4  57.9  54.8  49.0  42.4  39.2  maximum ^ t e m p e r a t u r e  40. 7  44.7  46.8  51.7  58.1  61.4  64.2  64.6  60 . 9  53.7  46.0  42.4  Mean daily minimum temperature  34.1  36.4  35. y  ,g g .38.»  No. o f days with f r o s t  x  Mean r a i n f a l l (inches) Mean snowfall Mean total ppt. No. o f y r s . record  of  6  .45  10.3 7.47 23  o  6  6  43  2  47  •^  50.6  7  1  *  0  51.2 °  48.7 0  44.3  38.8 *  0  36.0 3  7  4.47  4.98  3.31  2.15 2.09  2.06  2.49  4.44  8.44  7.76  7.99  3.8  2.1  0.1  0.0  0.0  0.0  0.0  1.7  5.0  23.0  4.86  4.30  3.32  2.15 2.09  2.06  2.49  4.44  8.44  7.95  8.48  22  22  22  22  22  22  22  Z3  23  22  0.0  22  10.  3.  3.1  FOOD HABITS  Introduction During the past four decades, a number of wildlife  studies have focussed upon the relationships between wolves and their prey.  In different geographical regions with  varying habitats and prey species, the diet of the wolf has received much attention.  Their food habits have been deter-  mined mainly by the analysis of scats, and also by direct observations of wolves interacting with and feeding upon their prey.  Major studies in which the diet of the wolf  has been investigated include those of Murie, 1944; Cowan, .1947; Thompson, 1952 ; Mech, 1966; Pimlott et al. , 1969; Kuyt, 1971; Carbyn, 1974; Van Ballenberghe et al., 1975; and Haber, 1977. These studies have shown that the wolf relies mainly upon ungulates for food.  Depending upon the prey  complex and habitat, general trends have been revealed regarding major versus minor prey, and seasonal variation in their utilization of prey species and type. Murie, who was the first to do a detailed study on wolves, found that in Mt. McKinley National Park, Alaska, they preyed upon caribou (Rangifer articus stonei), Dall sheep COvis dalli dalli), ground squirrels  (Citellus  parryii ablusus), marmots (Marmota caligata caligata), and  11.  voles  (Microtine S££.).  The summer diet  (May 1 to September  30) consisted mostly of calves and lambs, while adult animals were taken primarily during the winter to April 30).  (October 1  Ungulates constituted approximately 60 per  cent of the diet. In the Rocky Mountain National Parks, Cowan showed that the winter diet was made up of 49, 15, 9, 6 and 6 per cent elk  (Cervus canadensis), mule deer  hemionus), moose  (Odocoileus  (Alces americana), bighorn sheep  canadensis), and caribou  (Rangifer arcticus)  (Ovis  respectively.  Elk, deer, moose and bighorn occurred in 42, 14, 4, and 8 per cent respectively of the summer scats. found in 17 per cent of the summer scats. americanus), horse  Beaver were  Goat  (Oreamnois  (Equus caballus), rodents and other small  mammals were taken in limited Thompson's  quantitites.  (1952) work in northern Wisconsin  demonstrated that over 90 per cent of the wolf's diet was made up of white-tailed deer snowshoe hare  (Odocoileus virginianus), while  (Lepus americanus), and voles  pennsylvanicus and Clethrioromys  (Microtus  gapperi) served as secondary  food sources. A shift in the diet occurred one summer and fall during an abundance of voles in the area.  He thought  that wolves preyed upon a higher proportion of fawns versus adults during the summer (May 21 - August Thompson commented on the possible increased  31).  vulnerability  12.  of fawns after June 26 when they began to move about more, as opposed to their earlier period of life when they remained somewhat  secluded.  Moose were the major prey species of wolves on Isle Royale National Park, Michigan.  Mech (1966) found that  they occurred in 76 per cent of the scats while beaver constituted most of the remainder of the diet. Pimlott et al. (1969) working Algonquin Provincial Park, Ontario, reported that wolves relied principally upon white-tailed deer and utilized moose and beaver as secondary prey species  (these  occurred in the diet 80.5, 8.5, and 7.1  per cent respectively) . Hares and rodents were taken in small  riumbers.  In one area of the park where there was  an overabundance of beaver, wolves relied mainly upon this species for that year.  The summer diet  30) consisted mostly of fawns  (July 1 to September  (71% of. all deer) and calves  (881 of moose). In the Thelon River area of the Northwest Territories, Kuyt  (1971) showed that wolves relied almost  exclusively upon caribou.  Birds, microtines, and fish were  occasionally taken, particularly during the summer when small animals became important in areas' temporarily without caribou.  The winter diet was much less varied. Carbyn (1974) , working in Jasper National Park,  Alberta, found that elk, mule deer, moose, and sheep  13.  occurred in 46, 30, 8 and 2 per cent respectively of the summer scats  (May 1 to September 30).  During the winter  (October 1 to April 30) the same species occurred in 11, 66, 8 and 4 per cent of the scats. A study in northeastern Minnesota by Van Ballenberghe et al. (1975), revealed that white-tailed deer comprised the greater proportion  (55%) of summer scats while  moose and beaver constituted 13 and 10 per cent respectively of the diet.  In winter deer and moose occurred in 66 and  21 per cent of the scats respectively. Haber (1977) working in the same area as Murie (1944) found the summer diet of wolves to consist of caribou, moose, and sheep. young of the year.  Many of these animals were  During the winter wolves utilized  primarily older animals of these species. In the most recently published study of food habits, Theberge et al. (1978) observed that wolves in Algonquin Park, Ontario, have changed their food habits following a decline in deer (0. virginianus) and an increase in beaver.  Changes in occurrence of these prey species  followed their relative abundance.  They also found  differences among scat contents collected at rendezvous sites and in other areas, with  respect to beaver, deer and  moose, their major prey species.  14.  In these studies the diet of the wolf has been estimated  by calculating the percentage occurrence of vari-  ous prey species in scats.  However, this method is prone  to error due to the varying surface:  volume ratio of prey  eaten, which tends to overestimate the proportions of smaller animals eaten and underestimate those of larger animals  (Shelton, 1966; Mech, 1966, 1970; Kruuk, 1978;  Carbyn, 1974). wolves  The results of a study performed on captive  (Floyd et al., 1978) make it possible to determine  the relative total weight of prey species or type fed upon.  This method is more meaningful in describing the  quantitative relationship between wolves and their prey than the former.  This is particularly evident for the  summer diet of the wolf which often contains substantial proportions of young ungulates of varying sizes.  3.2  Methods Fresh scats were collected along regular travel  routes of wolf packs at intervals of five to seven days, and from den and rendezvous sites after packs had vacated them. A sample of each scat was collected and labelled with date and location.  Samples were stored in plastic bags at out-  door temperatures and later frozen.  A sample rather than  the whole scat was collected because the remainder of the  15.  scat was used as an attractant for capture purposes. Although wolf scats commonly contain one prey item, this method may not detect minor prey items. Scat samples were sterilized at 110°C for 3 hr. to kill the ova of Echinococcus granulosus To remove debris from hair shafts and then  identified  (Bowen, 1978).  samples were moistened  macroscopically on the basis of maximum  diameter, length, colouration, and pigment banding of guard hairs.  For comparison purposes a known collection of prey  species hairs, including those of both adults and young, was prepared.  More specifically, the criteria used to  identify prey species' hairs are as follows: Beaver:  Hairs characteristically brown along the shaft with spatulate tips.  Deer:  Adult hairs have a subterminal yellow or white band not exceeding 4 mm. in length. Fawn hairs have a maximum diameter less than 115 Jd.  Elk:  Adult hairs have a subterminal yellow or white band greater than 13 mm.  Rump patch  hairs have no subterminal band but may be easily distinguished from" those of deer which do possess a subterminal band. Calf hairs have a maximum diameter greater than 115 u. and less than 210yu.  16.  Food habits were analyzed by percentage occurrence of prey species or type in scats,  and by relative total  weight of prey type fed upon after the method of Floyd et al. (1978).  The regression equation used for the latter  method is as follows: y = 0.3 8 + 0 . 0 2x where y equals the kilograms of prey per collectable scat, and x the average weight (kg) of a given prey type.  This  method compensates for error in percentage occurrence calculations due to the varying surface:volume ratio of prey eaten which tends to overestimate proportions of smaller animals eaten and underestimate larger prey.  3.3  Results Analysis of 616 scat samples combined from both  packs for the total study period shows that the three major prey species were black-tailed deer, Roosevelt elk, and beaver (Table 3).  Comparison of the relative weights of  various ungulate age classes demonstrates that adult and calf elk replaced black-tailed deer fawns  as the second  most important food types (Table 3). Figure 2 displays monthly diets calculated from the combined samples of the two packs.  Young ungulates  were the major food items in the summer diet (June 1 to  17.  Table 3.  The diet of Vancouver Island wolves (Canis lupus crassodon) determined by analysis of scats from two packs collected between February and November, 1979. Relative weights of prey were determined following Floyd et al. (1978) .  CDPrTP, SPECIES  N1  N  RELATIVE OCCURRENCE m  RELATIVE WEIGHT (%)  13  2.1  0.7  Adult Deer  364  59.1  63.2  Deer Fawns  126  20.4  8.1  Adult Elk  27  4.4  14.1  Elk Calves  86  14.0  13.9  Beaver  1 - Number of scats in which each species was found.  Figure 2  Seasonal changes in the diets of wolves obtained from combined samples from both packs, estimated by relative weight (A) and relative occurrence (B).  No scat samples were  obtained in September. scats.  N shows number of  19  36  20  73  89  30  0  41  16  glOO  Feb Mar Apr May Jun Jul Aug Sep Oct Nov  KEY  beaver  adult deer  fawn  •  adult elk  August 31).  After this period it was not possible to  determine how important the above age class was,due to the development of adult pelage characteristics.  The decline in  utilization of beaver occurred after July when young ungulates became available.  Observations made in December, 1977  and January, 1978 showed that beaver are also utilized at this time. Figure 3 shows scats collected at den and rendezvous sites. mid-July.  Both packs occuppied dens from early May until The Upper Adam pack occupied  a rendezvous site  between mid-August to mid-October and was visited for scat collection after this time.  The Lower Adam pack's  rendezvous sites were not visited although attempts were made to find these sites.  Scats from both types of homesite  reveal a higher proportion of adult deer than those of monthly diets as shown in Figure 2.  Beaver occurred only  in den site scats, while a small proportion of adult elk occurred only at the Upper Adam rendezvous site. The seasonal diets of each pack viewed separately, shows various differences (Table 4).  Although the Upper  Adam pack utilized adult elk from February to May, there was no detectable use of this species by the' Lower Adam pack until June and July when use of adult and calf elk coincided.  An  increasing use of elk calves and fluctuating use of deer fawns is shown by both packs during the summer period.  Figure 10  Diets of wolves determined from scats collected at den (D-S) and rendezvous (R-S) sites.  Den sites occupied from  early May until mid-July and rendezvous sites mid-August to mid-October.  The  rendezvous site data is for the Upper Adam pack only, those for densite represent both packs. scats.  N. shows number of  22.  RELATIVE OCCURRENCE {%)  {X) 1 H D I 3 M  3AI1V13H  Table 4.  Diets of the Upper and Lower Adam wolf packs, expressed in terms of the relative weights of each prey species in the scats (after Floyd et al. 1978). Scats from den sites represent the period of occupancy from early May to midJuly, 1978 and rendezvous site scats represent the period mid-August to midOctober, 1978. RELATIVE WEIGHTS 1 OF EACH PREY SPECIES BY MONTH AND LOCATION  SPECIES  PACK  Upper Adam Beaver  Adult Deer  Deer Fawns  Adult Elk  MARCH  APRIL  MAY  0.9  0.8  0.9  2.2  7.5  8.5  1.9  49.7  63.2  55.9  51.5  100.0  92.5  91.5  48.4  Lower Adam Upper Adam  80.5  Lower Adam Upper Adam  -  Lower Adam Upper Adam  .  -  -  -  18.6  49.5  35.9  Number of Scats Collected  Upper Adam Lower Adam  Upper Adam Lower Adam  -  -  -  -  29  JULY 0.8  AUG -  -  NOV  DEN SITE  RENDEZVOUS SITE  1.5  -  85.1  50 .0 36.6  32.7  11.5  15.5  15.4  32.9  6.3  58.0  66.7  72.1 10.1  -  -  94.4  0.5  14.3  20.0 42.0  9.0  10.4  15.8  37.7  64.5  25.3  56.7  57.1  25  26  11  20  46  18  6  10  9  53  43  12  1 - Relative weights expressed as percentages.  OCT  -  41.9  Lower Adam  Elk Calves  JUNE  FEB  33.3  -  3.3 13.7  -  57 41  5.1  16  125  76  24 .  Although patterns of use did not coincide, adult deer utilization also seemed to fluctuate for both packs during this same period.  3.4  Discussion The abundance, distribution, and seasonal  availability of various prey species and age-classes appears to be reflected in the diet of the two packs studied. Another factor possibly contributing to dietary variations was the relative difference in pack size. No accurate data were obtained on the density and distribution of deer, but the limited observations made on this species indicated that they were both highly abundant and widely distributed throughout the study site.  The fact  that they were the major prey of wolves may be attributed to these factors.  The homogenous distribution of deer is  in contrast to that of elk, which display a "clumped" or heterogeneous distribution. Elk were less important in the diet of wolves, except during the summer period when calves constitute a higher proportion of the herds.  In  addition to their clumped distribution, elk undergo long range seasonal migrations which often take them into adjacent wolf pack ranges.  Due to these seasonal migra-  tions and their clumped distribution, elk are probably  taken opportunistically by wolves, particularly during the period when calves are not abundant. Beaver illustrate a further type of distribution which may be described as fixed relative to ungulate species. They were least important in the wolves' diets.  In contrast  to other regions where they are preyed on mainly during the summer (Cowan, 1947; Pimlott et al., 1969; Van Ballenberghe et al. , 1975 ; Theberge et al., 1978) beaver contributed to the diet of wolves in this study during the winter and spring period.  This is likely related to the mild climate  on Vancouver Island which leaves beaver habitat free from ice and snow, and thus the beaver vulnerable at this time. When comparing the summer versus winter diet, it is apparent that the wolves in this study vary their use of prey species and age-classes on a seasonal basis.  Deer  fawns and elk calves constitute the bulk of their diet during the summer, at which time the occurrence of beaver as a food source is almost non-existent.  This may reflect the  higher proportion of young ungulates present during this season, and possibly their greater availability and vulnerability to wolves, due to their size and inexperience at either outrunning or warding off predators.  Seasonal  variation in prey species and age-class utilization is well documented in previous studies (Cowan, 1947; Thompson, 1952; Pimlott et al., 1969; Van Ballenberghe et al., 1975).  The diets of individual packs show distinctions particularly during the summer period.  The larger,  Upper Adam pack, preyed on a higher proportion of adult deer during the early summer while the smaller, Lower Adam pack, depended more upon young ungulates.  This may be  related to the larger pack being able to capture  larger  size prey as has been demonstrated with coyotes (Bowen, 1978).  The high proportion of young ungulates in the  smaller pack's diet may be reflecting the inability of fawns and calves to escape, as contrasted to adults which often succeed in outrunning wolves.  The winter and spring  diets showed a greater proportion of adult elk taken by the large pack particularly during March, April, and May, when there were no apparent occurrences of this species in the smaller pack's diet.  Another possibility explaining  these individual pack diet variations is a difference in prey base within each pack's home range.  Sufficient infor-  mation was not available to test this hypothesis. Scat contents from den and rendezvous sites showed some differences in relative proportion and weights compared to scats collected from the rest of the home ranges. Comparable differences for rendezvous sites and other areas were shown for wolves in Algonquin Park (Theberge et a_l. , 1978) where the higher proportion of beaver in rendezvous site scats was attributed to the fact that beaver colonies  27 .  were adjacent to rendezvous sites.  However, no comparable  arguement can be made for the Vancouver Island .scat results which showed much higher use of adult deer at the den and rendezvous sites compared to other areas. In relation to other studies, the diet of the Vancouver Island race reveals both similarities and contrasts.  Variations in diet between regions probably reflect  the unique prey complexes the wolf deals with, and the distinct nature of the habitat which its activities and those of its prey are subjected to.  In particular, the diet  of the wolves studied resembles that of wolves in Ontario, Minnesota, and the Rocky Mountains. The largest prey present within the study site is elk, a position held by moose in Ontario and Minnesota (Pimlott et al., 1969; Van Ballenberghe et al., 1975). Otherwise, the prey complex is the same and proportions of deer and beaver which occur in the wolf's diet in these areas are similar to those found in the Vancouver Island race's diet.  Minor differences are shown by the presence  of moose in both the summer and winter diet of Minnesota wolves (Van Ballenberghe et al., 1975) . This may be attributable to the unavailability of beaver during the winter when streams and ponds are frozen over.  A further  consideration is a difference in prey ecology.  Elk herds  undergoing seasonal migrations may be contrasted to moose  28 .  which are mainly solitary non-migrators. to create a  This would tend  homogenous prey distribution in Ontario and  Minnesota (Mech, pers. comm) as opposed to the partial heterogenous distribution on the island. How the diet of the wolf is influenced by these factors is beyond the scope of this thesis but nevertheless noteworthy. Carbyn's (197 4) work showed that elk were taken in greater proportions during the summer and that male deer were the major food item in the wolves' diet. Cowan's (1947) data from the same region tend to conflict with Carbyn's showing instead elk as the major prey species. However, Cowan noted that elk were overabundant in the study site, so perhaps wolves were responding to this factor as has been demonstrated elsewhere (Thompson, 1952; Pimlott et al., 1969) . Although the prey complex in the Rocky Mountains is vastly different to that on Vancouver Island, making comparisons difficult, it is significant that in one of the above studies male deer were the major food source for wolves, with elk secondary and taken in greater proportions during the summer. If the diet of the two packs "studied is representative of others throughout the island it appears that this race demonstrates foraging patterns which are consistent with those of other regions.  The factors which  29.  contribute to variation between areas to be climatic differences and the unique ecology of prey species which inhabit distinct habitats.  30 .  4.  4.1  SOCIAL ORGANIZATION  Introduction The manner in which wolves are organized amongst  themselves and in relation to their environment has been investigated by several researchers (Murie, 1944; Cowan, 1947; Mech, 1966; Pimlott et al., 1969; Clark, 1971; Carbyn, 1974; Van Ballenberghe et al., 1975; Haber, 1977). Aspects of their social organization which have been focussed upon include pack distributions and sizes, wolf densities, home range sizes, daily and seasonal movements, and homesite (den and rendezvous sites) characteristics. Data have been obtained by maintaining close contact with wolves in their natural environments, either by intensive ground work often with the aid of aircraft, or by radiotelemetry . Table 5 summarizes the findings of several researchers on various aspects of wolf social organization in North America.  It is quite apparent that  estimates of home range size and wolf densities reported by those authors,  vary considerably between different  regions and even within areas.  Table 5.  Reference  Summary of results from previous studies of the social organization of wolves in North America. Location  Murie, 1944  Mt. McKinley, Alaska  Cowan, 1947  Banff and Jasper Park, Alberta  Burkholder, 1959  Alaska  Mech, 1966  Isle Royale, Michigan  Algonquin Park, Ontario  Clark, 1971  Baffin N.W.T.  Carbyn, 1974  Jasper Park, Alberta  Haber, 1977  Pack SiZe  1800  7A 5P  3  50 5000  Pimlott et al. 1969 Joslin, 1967  Van Ballenberghe et al. 1975  Home Range S i Z e d  Island,  Minnesota Mt. McKinley, Alaska  [4662.0]  [129.S] [12950.0]  4-5 10  205  [531.0]  2-3  105  [272.0]  15-21  40, 50, 60, 120 [103.6, 129.5, 155.4, 310.8] 1798.8 [4660]  7-8  15  _ Wolf Density*-  600-1000 [1554.0-2590.0]  Renovated fox dens formed in sandy or loamy soil. Within spruce and cotton wood stands or open riverside site entrances 15" x 20" to 16" x 25".  1 per 10-12 [25.9-31.1]  Sandy knolls, old beaver Houses, and rock over hangs.  1 per 500 [1295.0]  Information not given.  1 per 35-53 [90.7-137.3] 1 per 10-14 [25.9-36.3]  Adults of both sexes; "P":  Underground burrows, tree bases, hollow logs and rock caveS.  1 per 120 [311]  Expanded fox dens in sandy soils, rock overhangs or crevices. Within coniferous forests or mature aspen stands.  5-8  1 per 4.0-6.8 [10.4-17.6]  Sandy slopes, balsam and jack pine stands.  12-15  1 per 40-66.6 [103.6-172.5]  Elevated areas within spruce or willows. Southerly aspects. Several main entrances, interconnecting tunnels and chambers 10-20'.  Pups, less than 12 months of age.  c - Densities in wolves per m i 2 and [per k m 2 ]  South facing slope with entrance 17" x 28" and inside tunnels 12" in diameter.  1 per 6 [15.5]  a - Areas of home ranges in mi 2 and [ k m 2 ] b - "A":  Features  1 per 138 [357.4]  10-14 34.0-94.2 [88-244]  Den Site  32 .  4.2  Methods 4.2.1  Capture  Procedures  Individuals were captured in steel-jawed traps (#4 Newhouse Woodstream Corporation) with attached steel chains and "drag logs".  Sets were distributed over a wide  area, buried in trails and road sides frequently used by wolves.  Fresh wolf scats and urine were used as attrac-  tants.  Traps were checked daily at dawn, on foot, snow-  shoe or with a 4-wheel drive vehicle. Captured animals were immobilized with a mixture of phencyclidine hydrochloride (Sernylan, Parke, Davis and Co., 100 mg/ml) and promazine hydrochloride [Sparine, Wyeth Laboratories, 50 mg/ml), at a dosage rate of 1.0 mg/kg of estimated body weight for the former and an equal concentration of the latter.  Drugs were administered  intramuscularly to the upper hind leg region with a syringe on a pole. Radio-transmitters, with frequencies in the 150 mH z range, were secured around the animals' necks by bolting collar ends together.  Transmitters were  equipped with activity switches, weighed 345 grams, and had a theoretical life expectancy of four years (Wildlife Materials, Carbondale, Illinois).  Weights of wolves were taken with a spring balance scale to the nearest pound and later converted to kilograms. Standard body measurements, including total length, tail length, heart girth and hind foot length were taken. Permanent upper and lower canine teeth lengths were measured from gumline to tip, along the labial midline with vernier calipers.  Ages of wolves were estimated by tooth wear  CBowen, pers. comm.).  Photographs of incisors and canines  were taken for later comparison and checking age estimates. Colour coded ear tags (Ketchum Manufacturings Sales Ltd. , Ottawa, Ontario) were inserted into individual's right ears and lip tattoos applied to both sides of the lower lip for permanent future identification. Blood samples were drawn from the lateral saphenous vein in the hind-leg, or from the cephalic vein in the upper fore-leg.  Between 10-40 cc of blood were taken, 5 cc  for whole blood studies, 4 cc for Cortisol, and the remainder centrifuged for serum.  Samples were refrigerated, then  flown to the Puget Sound Institute of Pathology in Seattle for analysis on the same or following day. Blood samples were analyzed for the following constituents:  hemoglobin (Hgb), number, of red blood cells  (RBC), hematocrit (Hct), mean corpuscular volume (MCV), mean corpuscular hemoglobin concentration (MCHC), mean corpuscular hemoglobin (MCH), number of white blood cells (WBC),  and concentrations of cholesterol, calcium (Ca), phosphorus (P), bilirubin, serum urea nitrogen (BUN), glucose, lactic dehydrogenase (LDH), alkline phosphotase, serum glutamic oxalacetic transaminase (SGOT), creatinine, tryglyceride, sodium (Na), postassium (K), chloride (CI) , carbon dioxide (CC^) , total protein, albumin, thyroxine, and Cortisol.  Blood smears were used for anti-distemper antibody  analyses. Rectal temperatures of immobilized animals were periodically monitored, their overall conditions noted, and 2-4 cc of penicillin administered intramuscularly as standard procedure.  Once animals were "processed" they were  placed out of sight in shade, and left to recover on their own once they began to show head movement.  4.2.2  Radio-Location Methods Data on daily seasonal movements, temporal and  spatial distribution of packs, home range sizes, and pack sizes and compositions, were obtained by radio-telemetry methods. air.  Radio-tracking was performed from the ground and  Telmetry equipment utilized, included a portable  receiver (TRX-24, Cat. No. Clb) , a three-element Yagi antenna (Cat. No. C7d. Wildlife Materials), two fourelement Yagi antennae (Cushcraft Corporation, Mancester, New Hampshire), and a headset (Model H10-00, David Clark Company Inc., Worcester, Ma.).  Ground tracking involved taking several compass bearings from known map locations on each located individual.  When animals were inactive and at a den or rendez-  vous site, three compass bearings were taken. When they were active or away from known activity areas, five to six bearings were taken. Date, time of day, and animal state (active/inactive) were noted along with compass bearings. Bearings were later plotted on 20 chain (l":l/4 mile; 1": 402 metres) company maps (MacMillan Bloedel) and each individual's location determined by triangulation.  Polar  co-ordinates of locations were then calculated, and punched onto computer cards for analysis. For aerial tracking a Cessna-180 wheel or float plane was used.  Initially, a single three-element antenna  was used to locate wolves.  To improve precision two four-  element antennae were vertically mounted to wing struts, pointed slightly down relative to the wing's plane, and directed out under the wing.  Antennae cables were fed  through air vents and connected to a coaxial switch, receiver and headset.  When the general location of a wolf  was determined the plane was flown at an altitude ranging from 305 to 610 m. until transmitter si-gnals were heard. The pilot was then directed to fly in that direction which yielded equal signal strength on each antenna.  The  area from which the loudest signal was received was flown  36 .  over.  Circling and simultaneous switching to alternate  antennae enabled the tracker to determine whether the radioed animal was within the inner or outer circle flown by the plane.  The animals' location was then determined  to within a 200 m. radius by flying decreasing circles around it.  Final locations were made as close to the  ground as possible.  When animals were not directly  observed the date, time of day, animal state, habitat type and identification of monitored individuals were noted. For direct observations the same information was noted in addition to numbers of wolves observed, and activities at time of observation (interacting, resting, travelling). When the general location of animals was unknown the plane was flown at altitudes of 2,743 to 3,048 m. until signals were heard.  Once the animals' location was  found the plane descended and the former method was followed.  All aerial locations were plotted on 20 chain  maps and handled the same as ground locations.  On the few  occasions that weather conditions prevented low altitude tracking, locations were then determined from higher altitudes (1,524 - 1,829 metres).  4.2.3  Den and Rendezvous Sites Den and rendezvous sites were determined by repeated  radio locations from these sites, and were examined after packs had moved to other areas.  Descriptions were made  37 .  of den interiors, site locations, activity areas, site trails, and habitat type and vegetation in the immediate vicinity.  Site aspect, observations of other species and  their signs, were also noted.  Photographs were taken at  each site, particularly of den interiors, activity areas, and site trails.  4.3  Results 4.3.1  Weights, Measurements and Metabolic Profiles Weights and measurements of the nine wolves  captured during this study are shown in Table 6.  Adult  males were heavier and larger in body dimensions than adult females. Results of blood and serum analyses are shown in Tables 7, 8 and 9.  Problems of haemolysis or inadequate  sample size limited analyses for two individuals.  No  blood sample was collected from the small male pup. 4.3.2  Pack Sizes and Composition  The visual density of the habitats in the study area did not allow for many direct observations of packs. Therefore, information on pack sizes and composition is limited, and supplemented by indirect observations of radio-tagged individuals, animal tracks and sign, howling and signs of activity around den and rendezvous sites.  Table 6. Weights (kg) and measurements (mm) of nine Vancouver Island wolves (Canis lupus crassodon) captured between April 10 and November 1, 1978, near Kelsey Bay, British Columbia. Pack and Wolf S  tIdentlty j  e  x  Age  1  Total  Total  Tail  Weight  Length  Length  Hind Foot Length  Heart  Canine Length  Girth  Upper  Lower  Upper Adam  (87)  M  A  38.6  1803  495  267  550  24.5  Lower Adam  (86)  M  A  32.7  1800  483  241  787  28.3  25.1  M  A  36.7  1727  394  279  792  24.5  23.0  Lone Wolf  M  A  36.6  1780  460  300  770  26.0  23.5  Average  M  A  36.18  1777. 5  458 .0  271.8  752.3  25.83  23.87  2.484  35.18  45.07  24.62  62.21  1.795  1.097  10.0  1100  270  210  480  8.9  6.1  21.8  1490  380  250  610  19.2  19.3  Upper  Adam  2  Standard Deviation 3  -  Lower Adam  M  P  Lower Adam  F  P  Lower Adam (97)  F  A  31.8  1638  432  254  660  26.8  22.0  Upper Adam (90)  F  A  31.4  1715  432  264  729  23.0  18.5  Upper Adam (4)  F  Y  28.2  1638  392  254  711  22.8  19.7  Average  F  A/Y  30.47  1663.7  418.7  257.3  700.0  24.20  20.07  1.973  44.456  23.09  5.77  35.79  2.254  1.779  Standard Deviation  1 - Age classes:  4  A - Adult, P - Pup of year, Y - Yearling.  2 - Animal died shortly after handling. 3 - Estimated from denning period to be 4 - Estimated to be  months old.  months old.  Table 7.  Haemotology of some Vancouver Island wolves.  Weight (kg)  WOLVES  Hgb (g  RBC (10 6 /mm3)  Hct m  MCHC (Vol %)  WBC 103/mm3  Adult Males 60.2  84  28.8  12.9  62.9  76  33.1  15. 1  38.2  69  32.8  9.1  47.2  71  33.3  23.0  6 .37  47.8  75  31.4  11.2  12.5  5 .52  38.2  69  32.8  12.9  16. 1  6.35  49.4  72  Upper Adam  (87)  38.6  17.4  7 .22  Lower Adam  (86)  32.7  20.4  8.27  Upper Adam*  36.7  12 . 7  5.52  Lone Wolf  36. 7  15.6  6.64  21. 8  14.9  31.4 28 . 2  Female Pup Lower Adam  Adult/Yearling Females Upper Adam  (90)  Upper Adam (4)  * - This animal died shortly after handling  32.9  17.8  Serum chemistries of Vancouver Island wolves.  Table 8.  WEIGHT (kg)  CHOLESTEROL Cmg 1)  Upper Adam (87)  38.6  280  Lower Adam (86)  32.7  230  Upper Adam*  36.7  167  Lone Wolf  36.7  95  WOLVES  Ca (mg  P (mg t )  Bilirubin (mg . )  (mg \)  BUN  Glucose  0.8  32  140  Adult Males  Adult/Yearling  10 . 8  31.4  184  Lower Adam (4)  28.2  200  29  11.0  S.8  9.S  6.1  0.2  34  4.3  2.5  0.1  10.2  4.0  3.8  4.6  Females  Upper Adam (90)  4.4  0.3  (mg \)  100  LDH  ALK.PH. SGOT CREATININE TRIGLYCERIDE  (iu/l) (IU/DCIU/I) (mg  240 270  72  140  QNS  77  300+  1.0 0.8  160  186  56  155  QNS  53  76  25  130  0.6  0.2  43  116  130  55  186  0.7  0.2  15  115  215  59  350 +  0.3  »)  * - Animal died shortly after handling. 1 - Inadequate Sample.  O  41 .  Table 9.  Serum proteins and hormones.of Vancouver Island wolves.  WEIGHT  WOLVES  (kg)  TOTAL (g %)  ALBUMIN  THYROXINE CORTISOL  (g %)  ( g %)  ( g %)  Adult Males Upper Adam  (8 7)  38.6  8.0  4.4  1.9  QNS 1  Lower Adam  (8 7)  32.7  7.3  4.3  1.3  16.0  Upper Adam*  36.7  5.4  3,4  QNS  9.6  Lone Wolf  36. 7  3.4  1.9  0.7  31.4  6.6  3.7  2. 5  5.8  28.2  6.2  3.8  1.0  4.6  QNS  Adult/Yearling Females Upper Adam  (90)  Upper Adam (4)  1 - Insufficient sample. * - Animal died shortly after handling.  42 .  A.  The Lower Adam pack was estimated to con-  sist of an adult male, an adult female, and three pups (one female, one male and one unknown) which were born during May, 1978.  Observations of this pack were as follows: The adult male was captured on April 11, 1978,  and recaptured on June 23, 1978.  The adult female was  captured on April 14, 1978. The adult male was observed on May 24, 1978 at 09.50 hrs. in a clearcut adjacent to the denning area. At this time he moved off into the timber began to howl and was joined shortly by another wolf, thought to be the adult female. On June 22, 1978 at 09.30 hrs. the adult male and female were observed travelling through a clearcutsecond growth area approximately 1 km from their den. On July 12, 1978 at 22.30 hrs. the adult female was radio-located in a small roadside timber stand close to the den.  She responded to my howling with growls and  "yippy" barks.  Shortly after she howled a single animal,  thought to be adult male, began howling from the densite, followed by howling of pups.  When I returned to this area  at 23.14 hrs. there were tracks of a single animal crossing the road towards the den. A male pup was captured on July 28, 1978 and a female pup on October 29, 1978.  43 .  On August 29, 1978 at 17.00 hrs. the pack (adult male, adult female and three pups) was observed from the air on a small rock outcrop approximately 2 km from their den.  The adults were resting, and the pups romping and  playing among themselves. On October 9, 1978, the adult male was observed from the air resting in an open area on the banks of the Eve River. Members of the pack were not observed again until late November when the adult male was found shot in the field.  The female pup could not be radio-located at  this time and the radio on the adult female was no longer functioning.  Tracks of two animals were consistently  seen on snow covered roads in this pack's territory in January, 1979. It is possible that this pack consisted of more than five animals, however, the above information led me to believe the pack was composed of two adults (the breeding pair) and three pups.  The amount of sign seen  within their territory and around their den would support this belief. B.  The Upper Adam pack was believed to have  originally consisted of ten animals.  An adult male  from this pack died shortly after handling at the trapsite,  44 .  leaving nine pack members.  Obersvations of this pack were  as follows: An adult male was captured on April 10, 1978, an adult female on April 30, 1978 and yearling female on May 3, 1978. On May 24, 1978 at 12.00 hrs. the yearling female was observed alone travelling through a clearcut  area.  On July 24, 1978 at 07.00 hrs. the adult male was observed running down a grassy slope to cross a road. When he spotted my vehicle he turned and moved away from the road onto a small knoll and began to howl.  Other  pack members, including pups, responded from across the river.  It was not possible to determine numbers on the  basis of howling. On September 3, 1978 nine members of this pack were aerially observed at a rendezvous site from 12.3013.00 hrs. played.  Some individuals were resting while others  Individuals observed were thought to include  five pups, one other adult/yearling and the three radiotagged members. On September 6, 1978 the two radio-tagged females could not be located and five individuals were aerially observed including three pups and two adult/yearlings. The radioed male was located approximately two kilometres  45 .  from his rendezvous site but could no longer be located anywhere within the territory the following day. On February 16, 1979 at 12.30 hrs. nine wolves were observed together in a gravel pit approximately 4 km. from their densite of 1978 (Marika van Alderwegen, pers. comm.). On March 22, 1979 six members of this pack were aerially observed lying in the middle of a frozen lake (Keith McKillican, pers. comm.). On the basis of these observations, the Upper Adam pack was believed to originally consist of two adult males, one adult female, one yearling female, two unknown and four pups.  The aerial observation of September 3,  1978 is believed to be incorrect regarding pups versus adults, such that there were only 4 pups in this pack. The age classification of radio-tagged pack members is based upon weight and tooth wear  4.3.3  (Don Bowen, pers. comm.).  Home Range Sizes and Distributions The concept of territory is not used here  because investigation of defense of an area by a pack was not part of this study.  Home range is used instead  to describe the areas used by the wolves of the two packs, as determined by radio telemetry locations.  The defini-  tion  "... that  of home range is that of Burt  (1943):  46 .  area traversed by an individual(s) in its normal activities of food gathering, mating and caring for young." Home range  sizes were calculated by the convex polygon  method and plotted using a computer program developed by Harestad  (1979).  Home range areas were calculated for  100, 90, and 50 percent of the nearest  locations.  If an animal does occupy a home range, a plot of home range area against the number of locations should asymptote.  This is referred to as the "observation-  area" curve  (Odum and Kuenzler, 1955).  Unfortunately,  the time period covered in this study did not encompass an entire annual cycle, and animals of both packs did demonstrate seasonal shifts in spatial distribution, including their core areas  (50 percent home ranges).  The observation area curves shown in Figures 4 and 5 indicate that during the denning period an asymptote was reached for all individuals for their 100 per cent home range boundaries.  Following denning, further  increases  in home range size and corresponding changes in the asymptote level occurred.  These changes coincided for  most animals, with shifts in core area use, and may also reflect a general increase in movements between rendezvous sites, as the pups became more mobile. The observation-area curves for 90 per cent home range boundaries, show a different pattern  (Figures 4 and  Figure  4  Observation area curves for Lower Adam adult male's  (86) and adult female's  100 and 90 per cent home ranges.  (90)  Vertical  broken lines separate denning and postdenning periods.  48.  100  j o o  9 0  " V 9 0  ™r  20  r—~r——i  30  40  50  r  60  r  70  i  80  Number of Locations  i  90  i  100  Figure 4  Observation-area curves for Upper Adam adult male's  (87), adult female's  yearling female's home ranges.  (97) and  (4) 100 and 90 per cent  Vertical broken lines  separate denning and post-denning periods.  50 .  51 .  5), with a drop in area at or around the time the wolves vacated the den sites.  This decrease in 90 per cent home  range area indicates that during this shift from dens towards rendezvous sites, the wolves limited their movements within the previous 90 per cent areas, rather than travelling throughout the home range or moving into previously unoccupied  areas.  The data in Figures 4 and 5, further indicate that the cumulative home ranges  of both packs, considered  from the aspect of the annual cycle, may not have reached their maximum size. whole study period  Thus the home range sizes for the (Table 9) are likely underestimates of  the annual home range areas used by each pack. A.  Lower Adam Pack  Home range sizes for the two adult wolves monitored in this pack are shown in Tables 10 and 11, and together with plots of home ranges suggest that the adult male area than did the female  (Figures 6, 7 and 8)  (86) occupied a much larger  (90) .  Even though the radio  transmitter of female 90 eventually malfunctioned, the denning period should be comparable for these two animals.  The larger home range size of the male com-  pared to the female may reflect the fact that as the only adult member of the pack, he had a large share of  Table  2 10. Individual and pack home range sizes (km ) for the whole study period combined for two packs of Vancouver Island wolves.  WOLF AND PACK IDENTITY  PERIOD  NUMBER OF LOCATIONS  HOME RANGE SIZES 100%  90%  50%  75.0  33.8  3.3  18 . 5  15.1  1.7  Lower Adam: Adult Male  Apr 11-Nov 20  100  Apr 14-Aug 29  35  Apr 11-Nov 20  135  75.0  33.8  3.7  Apr 10-Sept 10  72  43.8  24.8  3.0  Apr 30-Sept 3  69  47.8  26.3  2.8  (4) May 3-Sept 3  72  63. 0  45.0  3.9  213  64. 0  43.6  3.9  (86)  Adult Female  (90)  Pack  Upper Adam: Adult Male  (87)  Adult Female  (97)  Yearling Female Pack  Apr 10-Sept 10  2146 Table 11.  Individual and pack home range sizes (km ) shown for denning (April 11 to July 11) and post-denning periods of two packs of Vancouver Island wolves. (Lower Adam: July 12 to November 20; Upper Adam: July 12 to September 10).  POST-DENNING PERIOD  DENNING PERIOD  WOLF AND PACK IDENTITY  Number of Locations  Home Range Sizes 50% 100% 90%  Number of Locations  Home Range Sizes 100% 90% 50%  Lower Adam: Adult Male (86)  40  43.1  20.9  1.1  60  56.3  32.0  3.2  Adult Female  25  16.4  15.1  0.3  10  10.5  2.5  0.1  65  43. 2  25.3  0.7  70  56.3  29. 0  3.3  43  42.1  20.9  1.6  29  20. 7  15.5  0.9  37  32.9  25.0  1.8  32  26.6  17.6  1.4  39  57.4  40.0  2.4  33  27.0  17.6  1.4  119  60.9  41.8  2.5  94  28 .1  22.7  1.7  (90)  Pack  Upper Adam: Adult Male  (87)  Adult Female  (97)  Yearling Female Pack  (4)  F i g u r e 10  100, 90 and 50 per cent home ranges of the adult male  (86) and the adult female  (4)  from the Lower Adam pack for the whole studyperiod  (86:  April 11 - November 20; 4:  April 14 - August 29, 1978). Orientation of axesis arbitrary but allows direct comparisons between all home range plots.  I  Figure  10  100, 90 and 50 per cent home ranges of the Lower Adam adult male during the denning (D) and post-denning  (PD) periods  (D:  April - July; PD: August - November, 1978).  F i g u r e 10  100, 90 and 50 per cent home ranges of the Lower Adam adult female during the denning (D) and post-dcnning to July; PD:  (PD) periods  August, 1978).  (D:  Apr  60 .  hunting to provide food for the female who was restricted in her movements because of the young pups at the den. Concentrations of locations for both animals in the denning and post-denning periods  (Figures 7 and 8) correspond to  the den site and the rendezvous sites occupied by them (Figure 9) . B. Upper Adam Pack Data on the sizes of home ranges  (Tables 10 and  11) of the three animals: monitored in this pack .show that the yearling female  (4) ranged through a larger home range  area than either of the two adults, particularly during the denning period.  All three wolves occupied similar  sized home ranges in the post-denning period. Home range plots for the three wolves of the Upper Adam pack are shown in Figures 10, 11, 12 and 13. The adult  (97) and yearling  (4) females have remarkably  similar location plots, particularly for their 90 and 50 per cent home range locations.  In fact during the post-  denning period they had essentially identical distributions.  They were actually located together on almost  every occasion in the post-denning period and very frequently during the denning period. The adult male  (87) while generally similar  in his distributions to the two females, was quently located with them.  infre-  His 50 per cent boundaries  F i g u r e 10  Home ranges  (100, 90 and 50 per cent)  of Lower Adam  (LA) and Upper Adam  (UA)  packs for whole study period superimposed on topographic map. rendezvous sites  Den sites  ( ^ ) and  ( • ) are shown.  Numbers  by rendezvous sites indicate order of use during the post-denning period.  For Lower  Adam 1-2 denotes alternating use of these two sites.  Rendezvous site 2 for Upper  Adam was last site radio-collared wolves located, site 3 had tracks after wolves shot.  Figure  10  100, 90 and 50 per cent home ranges of the adult male  (87), adult female  and yearling female  (90),  (4), of the Upper  Adam Pack for the whole study period (87:  April 10 - September 10; 97:  April 30 - September 3; 4: September 3, 1978) .  May 4 -  64.  5  I i f i 5  5  Figure 10  100, 90 and 50 per cent home ranges of the Upper Adam adult male  (87) for the  denning  (D) and post-denning  periods  (D:  (PD)  April - July; PD:  September, 1978).  August  i  1  1  5  1  r  F i g u r e 10  100, 90 and 50 percent home ranges for the Upper Adam female denning (D:  (D) and post-denning  April - July; PD:  1978).  (97) during the (PD) periods  August - September,  69.  Figure 13  100, 90 and 50 per cent home ranges of the Upper Adam yearling female denning (D:  (D) and post-denning  April - July; PD:  1978) .  (4) for the (PD) periods  August - September,  PD  71 .  did coincide with theirs, and locations throught the postdenning period were similar for all three animals.  This  similarity in distributions of the three pack members, together with the similar size of post-denning home ranges (Table 1 ) suggests a greater pack cohesion at this time compared to the previous denning period when pups would not be mobile. ground  Den and rendezvous sites located on the  (Figure 9), corresponded to the concentrations  of locations shown in home range plots  (Figures 10, 11,  12 and 13) . C.  Both Packs  Home range sizes of the two packs differed, with the Lower Adam male range  (86) occupying a larger home  when the whole study period was considered  (Table 10) and during the post-denning period However, observations of Lower Adam male  (Table 11) .  (86) extended  into November, while those for the Upper Adam pack were curtailed in September when animals were shot.  And,  although the observation-area plot for the Lower Adam male  (86) had asymptoted during the post-denning period  (Figure 4), those of the Upper Adam adult yearling  (97) and  (4) female appeared to be still increasing  (Figure 5). The observation-area curve of the Upper Adam adult male (87) did suggest an asymptote had been reached, however, in the post-denning period.  72 .  The relative distributions of the home ranges occupied by the two packs during the whole study period are shown in Fig.14.  Distances between pack home range  boundaries were approximately 1, 9 and 19 km for the 100, 90 and 50 per cent home range plots respectively 14).  Similar or greater distances when denning  15) and post-denning separately.  (Fig.  (Fig. (Fig.  16) periods are considered  Thus it appears that, at least for the radio-  collared individuals, the two packs held distinct, nonoverlapping home ranges during the study period, the core areas  (50 per cent plots) being the most  separate.  Whether this separation of core areas was by "design" or due to limitations of suitable home sites is unknown.  4.3.4  Wolf Densities Wolf density within, the study area, particularly  within the Upper pack's home range is likely to be higher than actual figures for reasons which will be discussed later.  On the basis of the available data the density 2 within the bower pack's range is one wolf per 15 km and one per 6.4 km 2 within the Upper pack area. 4.3.5  Daily and Seasonal Movements A.  Lower Adam Pack  The two radio-tagged members of- this pack were  F i g u r e 10  100, 90 and 50 per cent home ranges of the Lower Adam pack Adam pack (LA:  (LA) and the Upper  (UA) for the whole study period  April 11 - November 20; UA:  10 - September 10, 1978).  April  74.  +5  T  5 km j—i—I—L.  0 -  -3  o  i  r-  r  t  1—  5  1  r  Figure  10  100, 90 and 50 per cent home ranges of the Lower Adam  (LA) and the Upper Adam  (UA) packs during the denning period (April - July, 1978).  76 .  Figure 16  100, 90 and 50 per cent home ranges of the Lower Adam duri  (LA) and Upper Adam  the post-denning period  November;  (UA) packs (LA: August-  UA: August-September, 1978).  79 .  together when located 68.6 per cent of the time (24 out of 35 locations). To test the hypothesis that animal activity (active or inactive) was independent of time (day or night) a chi-squared value was calculated for both samples.  For  April through September 06.00 - 21.00 hrs. was described as day and 21.00 - 06.00 hrs. night.  For October and  November 08.00 - 18.00 hrs. was termed day and 18.00 08.00 hrs. night.  The above hypothesis was rejected (Chi2=  4.41, df=l, p<.05); it appeared that animals were more active at night. Shifts in the home range location of this pack were evident.  During April until. mid-July their activities  were centered around the den with occasional forays into other areas of their range.  During the post-denning period  they alternated between two core areas*, the first a site along the Eve River approximately 1.5 km from the den, and the other a higher elevation area 6 km from the den.  The  former site was characterized by coniferous and alder/ willow stands and expansive sand/gravel bars along the river.  The latter was within a pristine conifer stand at  1000 meters of elevation. Unfortunately, logistical constraints allowed only two occasions during which the movements of a wolf were monitored continuously for an extended period of  80 .  time.  The adult male (86) was followed for 10 hours on  September 14, (Fig. 17).  and for 5 hours on September 15, 1978  During these times he covered approximately  1.7 km in the 10 hour period and approximately 2.4 km in the 5 hour period.  These distances do support the differ-  ences in  versus nocturnal activity levels  daytime  described above.  The 5 hour period covered part of the  night, during which the animal travelled more than it did in a 10 hour daylight period. B.  Upper Adam Pack  The three radio-tagged members of this pack were together when located 45 per cent of the time (31 out of 69 locations).  The two females were together 88.4 per cent  of the time (61 out of 69 locations). The activity of animals in this pack was also dependent of time (Chi2 =15.38,  df = 1, p < .05).  This pack centered their activities primarily around the den from April until mid-July although extensive movements away from this homesite were evident for all tagged members.  After this time they moved to a  riverside site 6 km from the den, then to a rendezvous site 3 km from the latter.  The former area  was located  in a clearcut/second growth zone beside a creek flowing into the Adam river.  The latter was a typical rendezvous  Figure 17  Movements of the adult male (86) from the Lower Adam pack during a 10-hour and 5-hour period  06.30 - 16.30 hrs ( O )  14/9/78  18.30 - 23.00 hrs ( A )  15/9/78  last fix first fix  83 .  site as described in the homesite section.  Although not  borne out by radio-location data, direct observations of individuals and their sign suggests that the remaining members of the pack moved to a rendezvous site located 2 km from the den sometime during  December.  Differenential  utilization of the home range is also apparent for this pack. 4.3.6  Movements of a Lone Wolf This animal, an adult male, moved over such a  vast area of the Island that his movements and locations had to be obtained mainly from the air.  His movements are displayed  in Figure 18. He was captured during the early winter (November 1978) in an area believed to be between the two resident packs' ranges.  He remained in the general area  for three days then moved into the northern section of the Upper Adam pack's range. area.  They were still in the immediate  A month later he was located in another watershed  about 80 km directly southwest, and then a week later he had moved to the headwaters of the Adam river, a direct distance of 50 km.  Three days later he was located in the  Gold River valley 70 km to the south.  His next move took  him into the Salmon River 100 km from his last location.  Figure  18  Movements of a lone adult male wolf (November 1, 1978 - June 1, 1979)  C.S. - capture site  t  86.  It was possible to trade him on the ground here, at which time tracks in the snow indicated thatfoehad been following deer along an old logging grade.  He remained in this  watershed for at least two weeks, then moved 85 km farther south into the Campbell River watershed and stayed there for three weeks.  In this area he was observed on the  middle of a frozen lake feeding upon a deer (yearling confirmed on ground), which he had just recently (fresh blood) chased onto the lake and killed.  The throat of  the deer was the first part to be fed upon.  The snow  cover on the lake was crusted allowing the wolf to run on top while the deer broke through. circled above him,  Initially as the plane  he did not appear disturbed, but after  some time began to drag the carcass towards the lake's edge.  He then abandoned it and wandered off to lie down  in a sparsely treed area at the lake's edge.  Sometime  during the afternoon, or evening he dragged the kill about .3 km into a thick alder stand 25 meters from the water.  He was located resting a few meters from the  carcass and occasionally chasing off ravens which continuously landed on and picked at the carcass.  The night pre-  vious to the kill he was located in -a patch of second growth, inactive, at the lake's edge. After this time he moved in a southerly direction primarily along the east coast of Vancouver Island passing  87 .  through several watersheds inland from the coastal towns. When last located he was in an area just outside of Victoria where he remained for at least a week. When moving through mountainous terrain he seemed to travel the easiest route through mountain passes and along watersheds.  He was often located near one of  two major powerlines, which run from Campbell River to Victoria, and these would have afforded an easy travel route.  Several sections of the line revealed other wolf  sign and are probably usedby them in a similar manner to logging roads and old grades. 4.3.7  Den Site Characteristics A. Lower Adam Den The den was located in the base of a western  red cedar windfall which measured 24.5 meters in length. At its widest point the den entrance was 1.5 metres, separated into two passages by a central hump which extended 1.4 meters into the den.  The left passage  measured .98 meters and the right .56 meters into the den.  The left passage measured .98 meters and the right  .56 meters in height.  Where the den floor levelled out,  the single passage was .33 meters in height and .27 meters in width.  Farther into the den the passage opened  out being large enough for a human to sit with ample head  88.  and leg room.  The immediate entrance floor was dry, but  became increasingly damp farther into the den.  Several  pup scats were collected just inside the entrance and wolf hair was observed in several places inside the den. At the entrance level the den floor was 6 meters from ground level.  The outer circumference of the tree  base measured approximately 11 meters.  The den entrance  o  aspect was 292 W. The den site was located in a primary growth timber stand at 150 m. elevation which jutted out into clearcut and second growth hemlock zones. species were western hemlock and red cedar.  Dominant tree Shrubs  observed included red huckleberry and blueberry (Vaccinium spp.), salmonberry (Rubus spectabilis), devil's club (Oplopanax horridus) , and skunk cabbage (L.ysichitum americanum).  The ferns noted were oak fern (Gymnocarpium  dryopteris), sword fern (Polystichum munitum), deer fern (Blechnum spicant), and bracken (Pteridium aquilinum). A small stream ran through the timber stand in which the densite was located.  Numerous boggy sites were  observed which would likely be ponds during the wetter months when the pack utilized the den.  Another stream  with nearby ponds ran through the clearcut adjacent to the timber stand.  The nearest major river, the Eve, was  approximately 2 km from the densite.  89 .  Four major trails led to and from the den.  Two  Tan parallel to the denning log from behind the entrance, and two radiated in from the front at either side of the den entrance.  These trails branched into many side trails  throughout the forested site. A total of 126 scats were collected in the immediate vicinity of the den. The remains of two deer (lower jaws and femurs) were gathered in the surrounding area. Throughout the timber stand and near the den, were numerous squirrel chewings (hemlock cones), bear scats, and fresh deer tracks and pellets.  Small "digs"  at tree bases were observed possibly indicative of marten and squirrel.  A marten was observed near the den.  Fresh  elk pellets were observed in the clearcut adjacent to the timber stand. A well worn area forming a semi-circle of approximately 1 m. radius, suggestive of an activity site, was observed in front of the den.  To one side of  the den another windfall with well worn areas around and under it, was noted.  It appeared to provide cover for  at least two adult/yearling wolves.  Several pup and adult/  yearling scats were collected around this windfall.  90 .  Adjacent to the den entrance was a lateral seepage site supporting skunk cabbage and devil's club.  The  ground sloped up with increasing distance from the den, the forest floor being rather soft and spongy underfoot with numerous windfalls throughout.  Several of these  windfalls were worn down on the top giving the impression they had been utilized as resting/travelling/vantage points.  Throughout the surrounding area the tracks of pups  and adults were observed, particularly under windfalls and in boggy sites.  Scats of adults were often found on  windfall tops, and those of pups on ground level. B.  Upper Adam Den  Due to the nature of the habitat and topography in the area, the denning area utilized by the Upper Adam pack was not examined until November 22, 1978.  At that  time snow cover precluded making the desired detailed observations. The denning area was situated on a north facing ridge within an area of pristine timber at approximately 7 50 m altitude.  Several excavated caves were found at  the bases of red cedar and mountain hemlock (Tsuga mertensiana) trees.  These caves measured on the average  .4 meters in height and width.  The ground was well worn  around tree bases and wolf hairs were observed both  inside and at cave entrances.  Also noted were numerous  areas around den entrances w^ere wood had been chewed. The densite was located within a primary growth timber stand approximately 2 km from logged areas. Dominant tree species included red cedar, western hemlock and amabilis fir. s££.  The major shrubs were Vaccinium  Ground cover consisted of mosses, and arboreal  lichens were supported by dominant tree species.  Grasses  and sedges were the primary components in nearby high elevation wet meadows. Three high elevation wet meadows were close to the denning area, and two streams flowed in a northeast direction to the Adam River approximately 3 km from the site. One major trail ran up the ridge to and from the site and others radiated out into the timber.  Adja-  cent to the high elevation wet meadow nearest to the denning area a small activity area was observed. Several scats were collected around the bases of trees where the ground had been worn from animal activity. Fifty-four the site. surrounding  s c a t s were c o l l e c t e d at and around  Deer and elk remains were observed in the area while searching for the denning site.  92 .  Squirrel chewings were frequent about the site and fresh deer and old elk signs were observed in the surrounding area. 4.3.8  Rendezvous Site Characteristics A.  Lower Adam site  Although the rendezvous site of this pack was searched for, it was not found. B.  Upper Adam Site  Pack members were radio-located or aerially observed at this activity site from mid-August until midOctober, 1978. The site was situated in a high elevation wet meadow surrounded by pristine timber at approximately 800 m. altitude.  Numerous other meadows and bogs were  situated nearby.  The length of the meadow (90 m x 30 m)  ran in a southeast-north-west direction.  Several small  creeks flowed southwest into the Adam River about 3 km from the site. Trees common within the surrounding timber included mountain hemlock, amabilis fir, yellow cedar, western white pine and western yew. were supported by dominant trees.  Arboreal lichens  Major shrubs within  the meadow were Vaccinium spp. , and labrador tea.  93 .  Grass and sedge species were the dominant constituents of the meadow vegetation. Pack members appeared to predominately utilize only a small corner of the site for activity purposes. The main activity area was hidden from view of the remainder of the meadow by several stunted trees and snags. In this corner the ground was well worn around a small raised mound (5 m x 3m).  Digging was noted around the  mound and at the bases of numerous nearby trees.  One  major trail ran along the southwest edge of the meadow leading into the main activity area.  Several trails  radiated out from this area into the timber. Sixty-two scats were collected around the site and the remains of some deer kills observed in the surrounding timber. Observations around the site confirmed that both elk and deer were present near the site. and pellets of both species were noted.  Fresh tracks Another examina-  tion of this site in mid-November confirmed that deer were again present. Although contact with this pack was lost during September, 1978 it is known that they utilized another rendezvous site with similar features to the above during  December, 1978 through April, 1979.  This site was close  to their densite of 1978, therefore it appears they were again frequenting areas near to the denning area.  4.4  Discuss ion Morphological data from the nine wolves captured  in this study show that in comparison to similar data from other parts of North America, the adult Vancouver Island wolves of both sexes are heavier than those from Ontario (Pimlott et_ al. , 1969; Kolenosky and Stanfield, 1975), Quebec (Huot et_ aJ., 1978) and northeastern Minnesota (Van Ballenberghe, 1975).  And, while similar in weight  to Stenlund's (1955) sample from northeastern Minnesota, are lighter than animals from Alaska (Rausch, 1967) and the orthwest Territories (Fuller and Novakowski, 1955; Kelsall, 1968).  However, in terms of body dimensions  the Vancouver Island wolves measured here, show larger total lengths and tail lengths than those reported by Van Ballenberghe (1975) for northeastern Minnesota, and by Fuller and Novakowski (1955) and by Kelsall (1968) for wolves from the Northwest Territories.  Hind foot  lengths and upper canine lengths are similar to those reported by Van Ballenberghe (1975), while hind food lengths appear smaller than data from the Northwest Territories (Fuller and  Novakowski 1955, Kelsall 1968). Only one other published report presents data on metabolic profiles of wolves, and these are for pups (Seal et al., 1975).  Changes with age in certain  blood parameters have been reported for domestic dogs (Bulgin et_ al. , 1970).  If these are considered in com-  parisons with the data for wolf pups presented by Seal et al. (1975) and the adult Vancouver Island wolves studied here, it is apparent that the trends in total serum protein, serum albumin, haemoglobin concentration and RBC are comparable.  Seal et al. (1975) suggested that  relative levels of BUN and cholesterol between their samples for two years, indicated differences in nutritional state of their wolf pups.  The adult Vancouver  Island wolves had higher BUN values but lower cholesterol values than the wolf pups reported, by Seal et al. (1975). Both these values for wolves are higher than those reported by Bulgin et al. (1970) for beagles. Aspects of the social organization of the two wolf packs investigated in this study may be summarized as follows:  The Lower Adam pack, consisting of five  animals (including pups), utilized a- 75 km  2  home range, 2  with a resulting density of one wolf per 15 km . Within this home range they demonstrate seasonal shifts in core areas, which were related to activities around den and rendezvous sites.  They denned in a red cedar windfall  96 .  within pristine coniferous timber, and alternated their post-denning movements between two rendezvous sites, one a riverside site, the other at a higher elevation in coniferous timber.  The larger Upper Adam pack, estimated to  be composed of ten individuals (including pups), ranged within a 64 km2 home range, with a density of one wolf per 6.4 km2.  The smaller home range size of the large  pack, may be an underestimate, because the period over which they were followed was less than the period the small pack was monitored. densities,  Similarly, the lower calculated  may be due to the fact that only three of the  large pack were radio - collared, and thus other adult members may have included other areas in their home ranges, thereby increasing the home range size estimates for the whole pack.  Upper Adam wolves demonstrated similar sea-  sonal shifts in home ranges, moving from a densite located within a primary stand of timber at high elevation, to rendezvous sites at other locations. It was originally  hypothesised that the topo-  graphy within the study site would limit the distribution of packs to separate watersheds.  The distributions of  these two packs, together with incidental observations of other wolves in adjacent areas, suggest that this hypothesis is rejected in general, and applies only where  97 .  valley systems have very steep sides.  The two home  ranges observed, occurred within the same general watershed, and also encompassed other tributary drainages.  An  alternative hypothesis to be tested in the future, is that the wolves are limited in their distributions by a combination of suitable home sites, and availability of food sources, particularly high densities of black-tailed deer. In comparison to previously published studies of Nearctic wolves, those studied on Vancouver Island appear to hold smaller home ranges, and occur at higher densities  (Table 5).  However, the lack of year round  observations on Vancouver Island due to the illegal shooting of the radio-collared wolves, probably resulted in an underestimate of annual home range size as presented here. The actual estimates did approach those reported for the mid-latitude forested regions (Mech, 1966; Pimlott et al., 1969,; Van Ballenberghe et al., 1975), but were vastly different from the large home ranges estimated for wolves inhabiting Alaska and the Northwest Territories (Murie, 1944; Burkholder, 1959; Clarke, 1971; Haber, 1977). Pack sizes are somewhat difficult to compare between studies and between areas, as previously published studies have determined them using different methods. They are somewhat variable throughout North America  98 .  (Table 5) both between and within regions.  Furthermore,  the durations of the various studies also varies considerably.  Data gathered over longer periods may be more  accurate for estimating pack size, but significant fluctuations in pack numbers can vary over relatively short periods (Haber, 1977). There appears to be a general trend for wolf densities to be higher in the mid-latitudes (Cowan, 1947; Mech, 1966; Pimlott et al., 1969; Van Ballenberghe et al, 1975) as compared to those of the arctic and subarctic zones (Murie, 1944; Burkholder, 1959; Clark, 1971; Haber, 1977). Some combinations of differences in the densities and seasonal availabilities of prey species, probably accounts for this variability (Mech, 1970).  Although no  adequate data are available from the Vancouver Island study area, it is hypothesised that the relatively small home ranges the two packs occupied, and their high densities compared to other studies, reflects a higher prey density, particularly with regard to the main prey black-tailed deer.  Further studies of the Vancouver Island wolf at  study sites varying in black-tailed deer densities, would be required to test this hypothesis -of the influence of prey density on wolf home range size and density.  Homesites of both packs displayed good drainage, structural suitability, nearby water sources, and a clear view of the surrounding areas.  Prominent trails, activity  and resting sites, and animal signs (scats and tracks) were common features of homesites.  They were generally situated  towards the centre of home ranges, and spatially distributed several kilometers from the adjacent pack's sites. Characteristics of den sites vary in specific details among the various studies undertaken in North America. However, all generally display good drainage, structural suitability, a close source of water, and clear views of the surrounding areas (Murie, 1944; Cowan, 1947, Joslin, 1967; Pimlott et al., 1969; Mech, 1970; Clark, 1971; Carbyn, 1974; Haber, 1977).  All these features were asso-  ciated with the Vancouver Island wolves' den sites examined. In some studies, wolf dens were found to utilize previous dens of other species, and were sometimes re-excavated (Murie, 1944; Clarke, 1971; Haber, 1977). It is suggested that pristine conifer stands covering suitable substrate, or with large windfalls, are important for successful denning by Vancouver Island wolves.  Such areas are therefore necessary to preserve  during forest cutting operations.  100 .  Rendezvous sites show common features throughout the literature.  They are invariably situated in open  meadows bordering timber stands, and have signs of activity and resting sites, together with other signs 1967, Haber 1977).  (Joslin  Utilization of more than one rendez-  vous site within a pack's home range, similar to those reported here were found by Joslin (1967) for Ontario wolves.  However, these latter animals were found to  move more frequently between rendezvous sites than the Vancouver Island wolves. Observations of diurnal activity in the Vancouver Island wolves demonstrated that they were more often found resting during the daylight hours, but were significantly more active at night.  This together with  the limited observations collected during the two continuous monitoring periods, suggest that where studies involve only daylight observations or daylight radio telemetry locations, home range size and distribution may be significantly underestimated.  More intensive work  during the night period should provide a quite different picture from our present level of understanding, and have implications for theoretical considerations of home range sizes (Harestad and Bunnell, 1979).  101 .  The lone wolf captured and monitored during this study clearly does not fit the previous ideas about lone individuals  (Mech, 1970).  This animal was  apparently alone throughout the period he was followed, and ranged over a large portion of Vancouver  Island.  Although his movements appeared in part related to the north-south oriented, cleared powerlines, major intervalley movements were also apparent among his locations in the northern part of the Island. The social organization of wolves appears to be dependent upon the nature of the areas they inhabitat and the ecology of the prey species within the particular region.  Wolves inhabiting the mid-latitudes of North  America demonstrate similarities in social organization, while those found in the higher latitudes depart from these patterns.  The Vancouver Island race demonstrates  patterns which are consistent with the habitat and prey base.  102 .  5.  FINAL DISCUSSION  As man continues to alter the original state of his surroundings, the interrelationships between the environment and its wildlife inhabitants are receiving increasing attention.  One area of particular concern has  been the dynamics of predators and their prey.  During  the past two decades numerous field studies have focussed upon the behaviour and ecology of the large predators CMech, 1966; Hornocker, 1970; Kruuk, 1972; Schaller, 1972; Haber, 1977).  These studies and others have demonstrated  that the foraging habits of predators are closely related to both their own and their prey species' social organization and behaviour.  Diet and hunting technique have been  shown to differ according to the environment and prey species in question  (Schaller, 1972; Kruuk, 1975).  In  addition, the degree of sociality within certain carnivore species appears to be related to their exploitation of varying sized prey species  (Schaller, 1972; Bowen, 1978).  Conclusive data are not available from this study to clearly demonstrate the above principles. However, to a minor degree certain suggestions can be made.  Deer constitute the bulk of the wolves' diet on a  103.  yearly basis.  This has been shown in other environments  which support a similar prey complex (Pimlott et aJ., 1969; Carbyn, 1974; Van Ballenberghe et al., 1975; Theberge et al, 1978) .  The homogeneous distribution and  high abundance of this prey species appears to be reflected in the wolves' diet as opposed to elk and beaver which are alternate and usually secondary food sources.  Utilization  of elk may be opportunistic, owing to their clumped distribution, relatively low abundance and hence lower predictability compared to deer.  Beaver are predictably  distributed  but low in biomass and are preyed upon through the winter and spring months. larger  It appears that beaver constitute a  proportion of the diet during that time of year  when ungulate young are not available.  The occurrence of  of this species in the diet of wolves during this period is likely reflecting the relatively mild climate on the Island.  In more mountainous and eastern habitats this  species is preyed upon during the summer. The summer diet of wolves consists mainly of ungulate young.  Seasonal variation in utilization of  prey type and species is clearly demonstrated.  Wolves are  known to vary their diet on a season&l basis in other environments which support both a similar and unique prey complex (Murie, 1944; Pimlott et al. , 1969;  104.  Kuyt, 1971; Clark, 1971; Van Ballenberghe et al. , 1975; Theberge et al., 1978). Differences in the diet of the two packs studied here suggests a relationship between larger group foraging efficiency and their greater ability to prey on larger prey species captured.  This has been demonstrated for  other carnivore species  (Kruuk, 1972; Schaller, 1972;  Bowen, 1978).  Another possibility is a difference in the  prey distributions in the two packs' home ranges.  The  diet of the larger, Upper Adam pack, showed a higher proportion of adult elk on a yearly basis, whereas the smaller, Lower Adam pack, showed occurrences, of this species only during the summer.  Furthermore, the larger pack preyed  initially upon adult deer in the summer, while the smaller pack fed primarily upon ungulate youne.  The late summer  diet of the larger rack consisted of fawns, calves, and adult elk while that of the smaller showed mainly fawns and adult deer.  These findings sueeest that the total nrey  complex could be taken by the larger pack throughout the year as a result of their greater numbers, while the smaller pack was restricted to a partial prey complex and was seasonally dependent upon the largest prey species, elk. Theberge et al. (1978) report changes in diet related to changes in prey species abundance  in Ontario.  105.  Judging from the howling of packs as they left their den sites, presumably for hunting purposes, individual were often separated and seemed to travel away from dens in a fanned-out formation.  Once they had reached  valley bottoms or a particular site, it appeared that they located one another by howling.  This aspect of wolf  social behaviour is thought to act in coordinating their social hunting activities  (Theberge and Falls, 1967).  Both packs studied here displayed seasonal shifts within their total home ranges, and utilized core areas within these areas.  With the exception of the Upper pack's  denning area, both packs frequented rendezvous sites located at a higher elevation from sections of the home ranges used prior to this time.  These movements may have  been related to those of elk and deer.  The seasonal  migrations of the former are made to areas of higher elevation while deer appear to use forested areas bordering on clearcut zones at this time of year.  Fresh sign of  both prey species was noted around these rendezvous sites. However, there were no adequate data to demonstrate a direct relationship between the movements of wolves and their ungulate prey. Data on home range sizes, pack sizes, and hence densities are .limited by the lack of information, particularly  106 .  Mech, 1970) .  Previous researchers have thought that dens  were selected on the basis of drainage, water supply, suitable digging structure, early disappearance of snow, and a view.  Rendezvous sites are believed to be selected  for similar reasons.  I believe that site selection is  mainly based upon the presence of an abundant and close food source.  The other factors noted above may vary in impor-  tance within different habitats.  For instance, wolves in  higher latitudes which seem to hunt mostly by sight  (Clark,  1971) , appear to choose sites which provide a good view, an easily dug substrate, early disappearance of snow and a nearby water supply.  In forested areas dens often lack  such a view.  This may be related to their hunting a great  deal by smell  (Mech, 1970).  The elevated location of dens  appears to be a common factor in choice, which may result from their desire to be on a raised area.  This would  naturally provide good drainage and hence a dry  site, but  does not necessarily imply that areas are selected for this reason.  An adequate accessible water supply would also be  necessary for the lactating female (Mech, 1970) . Wolves probably base their choice of site mainly upon food supply.  Depending upon the habitat  involved  the other factors noted above are likely to influence their site choice secondarily.  A further possiblity is that  107.  sites are chosen in areas that are spatio-temperally separated from those of adjacent packs.  It would be  advantageous if not crucial for packs to separate themselves and maintain their daily and seasonal patterns with the least amount of contact with neighbouring packs.  Inter-  specific strife often resulting in the death of animals is known to occur with wolves  (Mech, 1977).  Throughout Vancouver Island and within the study site the habitat is constantly being altered by logging and and  other practices to the extent that the total "landscape  of regions is changed by the formation of clearcut and second growth zones which replace pristine timber.  The  effects of these practices upon the dynamics of wolves and their prey are not definitely known.  However, the vast  clearcut areas provide additional forage for ungulate specie which probably promote an increase in their numbers (Harestad, pers. comm.), and in turn also cause an increase in wolf numbers over time.  In timbered areas where  wolves formerly hunted by smell only they could .now hunt by both sight and smell.  Another equally important factor is  the human hunting pressure upon deer in these open areas, which in combination with wolf and cougar predation may act to depress deer numbers.  Furthermore, the occurrence  108.  of open (logged) areas may act against the wolf.  Consider-  ing five out of the eight radioed wolves in this study "disappeared", mostly by  shooting, during the deer hunting  season, it appears that their chances of survival are not enhanced in accessible areas of altered habitat. The interrelationships between the environment and its wildlife inhabitants are worthy of much attention. To successfully manage wildlife species it is necessary to determine the ecological needs of the predators and their prey species both in areas of altered habitat and those which remain in a pristine state.  109.  LITERATURE CITED  Bowen, W.D. 1978. Social Organization of the Coyote in Relation to Prey Size. Unpubl. Ph.D. thesis. University of British Columbia, Vancouver, 229 pp. Burkholder, B.L. 1959. Movements and Behaviour of a Wolf Pack in Alaska. J. Wildl. Manage., 23(1): 1-11. Bulgin, M.S., S.L. Munn, and W. Gee. 1970. Hematologic Changes to M Years of Age in Clinically Normal Beagles. J. Amer. Vet. Med. Assoc., 157: 1064-1070. Burt, W.H. 1943. Territoriality and Home Range Concepts as Applied to Mammals. J. Mammal., 24(3): 346-352. Carbyn, L.N. 1974. Wolf Predation and Behavioural interactions with Elk and Other Ungulates in an Area of High Prey Diversity. Can. Wildl. Serv. Publ., 233 pp. Clark, K.R.F. 1971 Food Habits and Behaviour of the Tundra Wolf on Central Baffin Island. Unpubl. Ph.D. thesis, University of Toronto, Toronto. Cowan, I.McT. 1947. The Timber Wolf in the Rocky Mountain National Parks of Canada. Can. J. Res., 25: 139-174. Cowan, I.McT. and C.J. Guiguet. 1975. The Mammals of British Columbia, B.C. Provincial Museum, Dept. of Recreation and Conservation. Handbook No. 11. 414 pp. Floyd  T.J., L.D. Mech, and P.A. Jordan. 1978. Relating Wolf Scat Content to Prey Consumed. J. Wildl. Manage, 42(3): 528-532.  Fuller W.A. and N.S. Novakowski. 19.55. Wolf Control Oper; tions, Wood Buffalo National Park, 1951-52. Can. Wildl. Serv. Wildl. Magmt. Bull., Ser. 1, No. 11. Haber  G.C. 1977. Socio-ecological dynamics of wolves and prey in a subarctic ecosystem. Unpubl. Ph.D. thesis University of British Columbia, Vancouver, 786 pp.  110.  Harestad, A.S. 1979. Seasonal Movements of Black-Tailed Deer on Northern Vancouver Island. Unpubl. Ph.D. thesis, University of British Columbia, Vancouver, 184 pp. Harestad, A.S. and F.l. Bunnell. 1979. Home Range and Body Weight - a Reevaluation. Ecology, 60: 389402. Hornocker, M.G. 1970. An analysis of Mountain . Predation upon Mule Deer and Elk xn the Idaho Primitive Area. Wildl. Monogr., 21: 39 pp. Huot  J., D. Banville and H. Solicoeur. 1978. Etude de 'la predation par le loup sur le cerf de v i r g m i e dans la region de l'outaouais, Quebec. Minist. Tour., Chasse et Peche, Dir. Rech. Faun, 77 pp.  Jolicoeur, P. 1959.  on Evolution, 13. 285-zyy  Multivariate Geographical Variati  in the Wolf Canis lupus 1.  Joslin, P.W.B. 1967. Movements and Home Sites at Timber Wolves in Algonquin Park. Am. Zool., 7: 279-Z86. Kelsall, J.P. The Migratory Barren-Ground Caribou of Canada. Can. Wildl. Serv. Monogr. Ser., 3. 340 pp. Kleiman, D.G. and J.F. Eisenberg. 1973 Comparisons of Canid and Felid Social Systems from an Evolutionary Perspective Anim. Behav., 21: 65/-Oi>y. Kolenosky G. and R.O. Stanfield. 1975. Morphological and Ecological Variation among Gray Wolves (C. l u p u s ) of Ontario, Canada. In "The Wild Canids: TfielrSysternatics, Behavioural Ecology and Evolution", £ d - M . Fox, Van Nostrand R e m h o l d Co., New York. 508 pp. V  v T and R C Brooke. 1969/70. Ecology of A s t e r n North America. Dept of Botany, University of British Columbia. Vol. 2 (.1 W J •  Kruuk, H. 1972. The Spotted Hyena. University of Chicago Press, Chicago. 33b pp.  111. Kruuk, H. 1975. Functional Asspects of Social Hunting by Carnivores. In "Function and Evolution in Behaviour", eds. G. Baerends,.C, Beer and:,A. Manning, Clarendon Press,-Oxford. Kuyt, E. 1971. Food Studies of Wolves on Barren-Ground 'caribou Range in the Northwest Territories. Canadian Wildl. Ser., Progress Notes #23. Lawrence, B. and W.H. Bossert. 1957. Multiple Character Analysis of Canis lupus, Catrans and f a m i l i a n s , with a Discussion of the Relationship of C a m s niger Am. Zool., 7: 223-232 Makridin, V.P. 1962. The Wolf in the Yamal North. Zhur. 41: 1413-1417. Mech, L.D. 1966. The Wolves of Isle Royale. 'part. Serv. Fauna Series No. 7, 210 pp. -Mech, L.D. The Wolf. '384 pp. Mech  Zool.  U.S. Nat.  Natural History Press, New York.  L.D. 1977. Productivity, Mortality and Population 'Trends of Wolves in Northeastern Minnesota. J. of Mammal., 58: 559-574.  Murie  A 1944. The wolves of Mount McKinley. Fauna of the National Parks of the U.S. Series. No. 4. 204 pp.  Odum, E.P. and E.J. Kuenzler. 1955._ Measurement of Territory and home range size M b m a l s . AUK, 72: 128-137. Pimlott D H. J.A. Shannon and G.B. Kolenosky. 1969. _ The Ecoiogy of the Timber Wolf in Algonquin Provincial Park Dept. of Lands and Forests, Ontario. 92  iiniSSS:  112.  Schaller, 1972. The Serengeti Lion: A Study of Predator-Prey Relations. University of Chicago Press. Chicago. 480 pp. Seal  U.S., L.D. Mech, and V. Van Ballenberghe. 1975. 'Blood Analyses of Wolf Pups and Their Ecological and Metabolic Interpretation. J. Mammal., 56: 64-75.  Shelton, P.C. 1966. Ecological Studies of Beavers, Wolves and Moose in Isle Royale National Park. thesis. Purdue University, 308 pp.  Ph.D.  Stenlund, M.H. 1955. A Field Study of the Timber Wolf (Canis Lupus) on the Superior National Forest, Minnesota. Minnesota Dept. Conserv. Tech. Bull., 4: 5 5 pp. Theberge, J.B. and J.B. Falls. 1967. Howling as a Means of Communication in Timber Wolves. Am. Zool. /: 331-338. Theberge, J.B., S.M. Oosenbrug, and D.H. Pimlott. Site_ and Seasonal Variations in Food of Wolves Algonquin Park, Ontario. Can. Field-Nat., 92: 91-94. Thompson, D.Q. 1952. T r a v e l , R a n g e and Food Habits of Timber Wolves in Wisconsin. J. Mammal., 33. 49L 442. vaT1 Ba 11 enberehe V. 1975. Physical Characteristics of 'an Timber Wolves in Minnesota! In "Proceedings of the 1975 Predator Symposium" Eds. R.L Phillips and C. Jonkel, Mont. For. Cons. Expt. Sta., U. Mont., Missoula.  Wildlife Monograph #43: 43 pp.  *  hl  T r r 1 978 The Importance of a Relative " s h I r t a g ; if Food in A n L a l Ecology. Oecologia 33: 71-86.  


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