COMPORTAMENTO; ORGANIZAÇÃO SOCIAL; PREDAÇÃO; POPULAÇÕES

(Behaviour; social organization; predation; populations)

ARTIGOS

Goldman JA, 1995 An acoustic basis for maternal recognition in timber wolves (Canis lupus)? J Acoust Soc Am 97(3), 1970-1973 (1995)

Fico R, 1993 The impact of predators on livestock in the Abruzzo region of Italy. Rev Sci Tech 12(1), 39-50 (1993)

Morley JE, 1983 The effect of naloxone on feeding and spontaneous locomotion in the wolf. Physiol Behav 30(3), 331-334 (1983)

Peterson EK, 1981 Olfactory pedunculotomy induced anosmia in the wolf (Canis lupus). Physiol Behav 27(3), 543-546 (1981)

Lentfer JW, 1973 Notes on the captive wolf (Canis lupus) colony, Barrow, Alaska. Can J Zool 51(6), 623-627 (1973)

Beauchamp G K. Maller O. Rogers J G Jr. Flavor preferences in cats felis-catus and panthera-sp. Journal of Comparative & Physiological Psychology 91 (5). 1977. 1118-1127. Abstract In the 1st experiment domestic cats exhibited no preference, both in 24-h and 1-h 2-choice preference tests, for any of a variety of carbohydrate or artificial sweeteners regardless of whether a water or saline diluent was employed. A preference for sucrose or lactose dissolved in dilute milk compared with dilute milk alone was observed. This preference may be based on textural rather than flavor characteristics of the milk-sugar solution. In the 2nd experiment a similar lack of preference for carbohydrate sweeteners was found when using 5-min 2-choice preference tests with wild cats (Panthera). The 3rd and 4th experiments examined responses to solutions of hydrolyzed protein and individual amino acids and to emulsified fat mixtures. Solutions of hydrolyzed soy, lactalbumin, casein, L-alanine, L-proline and butterfat mixtures were all preferred to the diluent. A pattern of responses characterized by an avidity for protein and fat products was indicated, and no avidity for carbohydrate sweeteners may be typical of strict carnivores like cats.

McLeod, PJ; Fentress, JC Developmental changes in the sequential behaviour of interacting timber wolf pups. Behavioural Processes, 39(2), 127-136, 1997

McLeod, PJ Developmental changes in associations among timber wolf postures. Behavioural Processes, 38, 105-118, 1996

McLeod, P.J.; Moger, W.H.; Ryon, J.; Gadbois, S.; Fentress, J.C. The relation between urinary cortisol levels and behavior in captive timber wolves Can. J. Zool., 74(2), 209-16, 1996

Fentress, JC; McLeod, PJ Pattern construction in behavior. In W.P. Smotherman & S.R. Robinson (eds), Behavior of the fetus (pp. 63-76), Caldwell N.J.: Telford Press (1988)

Fentress, JC; Ryon, J; McLeod, PJ Coyote adult-pup interactions in the first three months Can. J. Zool. 65, 760-763, 1987

Fentress, JC; McLeod, PJ Motor patterns in development. In E. Blass (ed), Handbook of behavioral neurobiology: vol 8. Developmental processes in psychobiology and neurobiology (pp. 35-97), New York: Plenum Press (1986)

Fentress, JC; Ryon, J, McLeod, PJ; Havkin, GZ A multidimensional approach to agonistic behavior in wolves. In H. Frank (ed), Captive wolf research: advances, issues, and problems. (p. 253-374), Dordrecht, The Netherlands: Dr. W. Junk Publishing (1986)

TESES

Spaulding, Raymond Leon. DIET AND OBSERVER BIAS IN SCAT ANALYSIS OF GRAY WOLVES (CANIS LUPUS). Masters Abstracts International. Volume: 35-03, page: 0736. Abstract I analyzed 1,182 gray wolf (Canis lupus) scats, representing summer diet in 1988-1990, collected from 9 packs in northwest Alaska. Using a linear regression model, I determined the biomass of prey species consumed using relative estimated bulk and frequency of occurrence of prey types in scats. There were no significant differences $(P < 0.05)$ between methods in percent biomass of prey consumed and ranking of prey types. Caribou (Rangifer tarandus), moose (Alces alces), microtines, and birds composed 47.0, 21.9, 10.6, and 13.8%, of prey items, respectively. Ungulates composed 90.1%, birds 6.7%, and microtines 0.5% of biomass consumed. Ungulates composed 3.4%, birds 50.4%, and microtines 38.5% of numbers of individuals consumed. Significant $(P < 0.05)$ observer bias was found in the detection and frequency of prey types in the analysis of 1,052 scats by multiple observers.

Townsend, Susan Elizabeth. THE ROLE OF SOCIAL COGNITION IN FEEDING, MARKING, AND CACHING IN CAPTIVE WOLVES, CANIS LUPUS LYCAON AND CANIS LUPUS BAILEYI (AUDIENCE EFFECT, RANK, FOOD MARKING). Dissertation Abstracts International. Volume: 57-10, Section: B, page: 6129. Abstract In this study, I investigated whether captive wolves modify their behavior in order to convey information to or withhold information from conspecifics. Cognitive and ecological explanations are used to assess the significance of the "audience effect." Specifically, I measured how the presence or absence of conspecifics affected the frequency (a) of urination, (b) with which various urination postures were used, (c) of urine-marking food, (d) of caching and recovering of cached food, and (e) of urine-marking caches. These behaviors may be useful in conveying information about social rank (frequency of urine-marking and urine-marking posture), food ownership (urine marking-food), and location of food (caching and cache-recovery). Two separate captive groups representing two regional subspecies of wolves were studied for four research seasons (1991-1994). General activity, feeding, and caching behavior were recorded onto a hand-held microcassette recorder and by video-taping. As predicted, high ranking wolves urine marked significantly more often in the presence of conspecifics than in their absence. The sex and rank of a wolf determined which urination posture was used most frequently in the presence of an audience: the males used the raised leg urination posture, and, in two of three instances, high ranking females used the squat urination posture. Urine-marking food had no relationship to whom actually fed on the food; however, high-ranking individuals of one subspecies and all the Mexican gray wolves urine marked on food significantly more often in the presence of an audience than in the absence of one. Food-caching was performed more often in the absence of conspecifics than in their presence; however, food cache-recovery by individuals other than who had originally made the cache occurred in the presence of the individual who had originally made the cache about as often as in their absence. These results support in part the hypothesis that wolves modify their behavior in order to convey information to or withhold information from an audience of conspecifics.

Coscia, Elizabeth Maria. ONTOGENY OF TIMBER WOLF VOCALIZATIONS: ACOUSTIC PROPERTIES AND BEHAVIOURALCONTEXTS (CANIS LUPUS). Dissertation Abstracts International. Volume: 57-05, Section: B, page: 3397. Abstract This thesis concerns the ontogeny of vocal patterns in wolves, including their acoustic properties, contexts of expression and apparent functions. While it has long been recognized that wolves are among the most complex terrestrial social mammals and that their vocal behaviour contributes significantly to their social organization, objective and quantitative assessments of wolf vocal ontogeny have been lacking. In this study I have provided the first systematic evaluation of neonatal wolf vocalization patterns. Recordings were obtained from three litters of pups in a semi-natural, undisturbed den. The development of vocal behaviour during the pups' first six weeks of age was quantified using a multidimensional classification scheme. I catalogued 3396 sounds and 800 behavioural events and subjected them to detailed analyses. Vocalizations were divided into 11 classes based on their spectrographic structure. Properties used to distinguish vocal classes included: gross spectral type, duration and three frequency variables. Measures of frequency variation, inter-element-interval duration of sound series and relative amplitude were also taken for selected sounds. Eight vocal classes were quantified. Vocal classes were subdivided into three groups: early-appearing neonate-specific, early-appearing adult-structured and late-appearing adult-structured. Ontogenetic trends included changes in the relative proportion of specific vocal classes, decreases in fundamental frequency and combinations of specific sounds into patterns of non-random series. The behavioural context of vocal classes was examined with descriptive and statistical analyses of the relations between selected vocal classes and three behavioural categories: pup and adult behaviour at onset of pup vocal activity, and subsequent adult responses to vocalizations. The context of vocal classes distinguished as neonate-specific did not vary significantly with maturation, whereas the context of the adult-structured classes varied with changes in social and physical development. Interpretations of the expressive content and the function of pup vocal behaviour were based on the objective criteria used to describe behavioural changes in development.

Adams, Layne George. CALF PRODUCTION AND SURVIVAL IN THE DENALI CARIBOU HERD, ALASKA (PREDATION, RANGIFER TARANDUS, URSUS ARCTOS, CANIS LUPUS). Dissertation Abstracts International. Volume: 57-04, Section: B, page: 2246. Abstract Calf production and survival are important components of caribou (Rangifer tarandus) population dynamics. During 1984-95, I investigated female reproductive performance and calf mortality of caribou in Denali National Park, Alaska where they occur at low densities (0.3/km$^2$) and exhibit high calf losses. Age-specific natality increased from 27% at 2 years of age to 97% for 7-13 year-olds, then declined for females $ >= $14. Natality increased significantly (P $<$ 0.05) with body mass for 10-month-old females weighed in March, 6 months prior to breeding, and for females $>$1 year old weighed during the autumn breeding season. In most years, sexually mature females reproduced at high rates, regardless of lactation status during the breeding season. However, following unusual late winter snows and a short growing season in 1992, 1993 productivity declined significantly for cows with calves in autumn 1992. High calf losses to predation in 1992 were partially compensated for by the resultant improved 1993 productivity for cows that lost their 1992 calves prior to the breeding season. Calving began between 4 and 15 May and the peak of calving, or median calving date, occurred between 13 and 21 May. Body masses of females in autumn were significantly correlated with their subsequent parturition date, but female masses in late winter were not. The timing of calving for the population was significantly correlated with February-May snowfall prior to the breeding season. During 1984-87, 39% of radiocollared calves died within 15 days of birth, primarily from grizzly bear (Ursus arctos) and wolf (Canis lupus) predation. The rate of bear-caused mortalities declined significantly with age, and bears killed few calves $>$10 days old. Wolf predation was not significantly related to calf age but occurred primarily during about 6 days immediately following the peak of calving. Annually, the mortality rate for calves $leq$15 days old was inversely correlated with average birth mass. Annual losses to wolves were more strongly correlated with average birth mass than with spring wolf density. Even though the wolf population doubled during this study, the ratio of wolves to calves, or exposure of caribou calves to wolf predation, did not increase, when spacing by caribou at the wolf pack territory scale was accounted for.

Hayes, Robert D. NUMERICAL AND FUNCTIONAL RESPONSES OF WOLVES, AND REGULATION OF MOOSE IN THEYUKON (POPULATION ECOLOGY, CANIS LUPUS,ALCES ALCES, RANGIFER TARANDUS, OVIS DALLI). Masters Abstracts International. Volume: 34-04, page: 1496. Abstract Numerical and functional responses of wolves (Canis lupus) were studied in a 23,000 km$^2$ area of the east-central Yukon. Populations of wolf, moose (Alces alces) and woodland caribou (Rangifer tatandus caribou) were increasing following intensive reduction of wolf numbers. Snow-tracking surveys and radiotelemetry studies indicate that the wolf population recovered to pre-reduction densities within 4 years. Two hundred and ninety-one moose, 30 caribou and 1 mountain sheep (Ovis dalli) were found dead during my study. Wolves killed mainly young and old moose and most prey were not nutritionally stressed. Killing rates by 21 different wolf packs were studied during 45 periods in late winter. Kill rate of moose by wolves was negatively correlated with wolf pack size but was not correlated with moose density, prey searching rate, snow depth, observation rates, wolf:prey ratios, availability of alternate prey, or snowshoe hare abundance. Also, kill rate of moose calves by wolves was not correlated to wolf pack size, snow depth or calf availability each winter. Wolves in small packs had disproportionately higher kill rates on moose compared to wolves in large packs. I combine data from other studies and show that wolf predation could regulate moose to a single low density equilibrium (0.12 moose/km$^2$) in most wolf:moose systems in North America. Bear predation and changes to moose habitat quality have little effect on the stable equilibrium point, where moose are the primary prey of wolves. My model also indicates that no unstable upper density boundary exists beyond which moose could escape the regulating effect of wolf predation. (Abstract shortened by UMI.).

White, Katrin Anne Jane. TERRITORIALITY AND SURVIVAL IN WOLF-DEER INTERACTIONS (CANIS LUPUS, ODOCOILEUS VIRGINIANUS, PREDATOR PREY DYNAMICS). Dissertation Abstracts International. Volume: 56-07, Section: B, page: 3564. Abstract Territoriality is a commonly observed behaviour in mammals and gives rise both to spatial and temporal heterogeneities in land use. Despite the frequency and nature of this behaviour, there have been few attempts to model the phenomenon mathematically and none, as far as we are aware, which deal with the spatio-temporal aspects involved. Motivation for this research came from the extensive field studies carried out in northeastern Minnesota where wolf (Canis lupus) pack territories are maintained over many years. Separating the territories are buffer zones where wolves are scarce but where there are increased levels of scent marking. Moreover, the primary prey species, white-tailed deer (Odocoileus virginianus) are found most abundantly in the buffer regions which suggests that the spatial heterogeneity in predator density gives rise to refuges for the prey. We develop models based on simple movement rules for the wolves and on a basic predator-prey interaction. We begin by describing and analysing a single pack model--even in this simple case territories can be formed and we show how the size of the territory depends on pack size. When several packs interact and when deer are incorporated explicitly into the model system, we obtain density distributions which are consistent with the field observations. We investigate the predator-prey dynamics with seasonal reproduction by assuming that mortality occurs continuously throughout the year and that birth occurs as a discrete-time pulse into this otherwise decaying system. We also develop a simulation model to investigate the predator-prey dynamics in which case the model parameters can be estimated from the field data. The model systems are mathematically challenging and provide a new framework within which theoretical ecology can address the issue of territoriality and predator-prey interactions.

Garner, Dale Lee. POPULATION ECOLOGY OF MOOSE IN ALGONQUIN PROVINCIAL PARK, ONTARIO, CANADA (ALCES ALCES). Dissertation Abstracts International. Volume: 56-03, Section: B, page: 1171. Abstract Population dynamics of moose (Alces alces) were investigated in Algonquin Provincial Park based on long-term studies of moose population abundance and factors influencing demographics, 1976-94. In addition, intensive monitoring was conducted from 1990-93 on 76 radio-tagged animals (27 adult and 1 yearling female, 47 neonates) and 16 untagged neonates to determine reproductive performance, survival rates, and cause-specific mortality agents. Aerial survey methods indicate that the detection of $\pm$10% change in population abundance is possible. Reproductive performance from 1990-93 shows average production of 0.95 calves/cow and twinning rate of 18% which suggests Algonquin Park's population may be near or at ecological carrying capacity. Estimated yearling productivity was within the range reported in the literature for moose populations across North America. Annual survival rate for adults (0.94) and estimated yearling survival (0.88) correspond to rates observed in other populations across North America. However, annual calf survival (0.61) is high compared to other populations where both black bear (Ursus americanus) and wolves (Canis lupus) are sympatric. Winter weather has not played a significant role in moose population dynamics in the past 18 years. Winter tick (Dermacentor albipictus) causes mortality in both young and adults, but does not appear to have a significant influence on overall population change in either the current year or subsequent years. Long-term population trends and growth rates appear to be mediated primarily by ecological carrying capacity. Increasing visitor use of Algonquin Park and changes in policy allowing Native harvest may place additional stress on the population, thus warranting continued monitoring.

Loeven, Judith C. THE ONTOGENY OF SOCIAL PLAY IN TIMBER WOLVES, CANIS LUPUS. Masters Abstracts International. Volume: 33-04, page: 1176. Abstract Play behavior has been characterized both structurally, by describing its physical form, and functionally, by inferring its possible causation and evolutionary function. These defining characteristics rarely are stated clearly, and quantitative measures of play behavior are even less frequent. This study was one of the first to use quantitative methods to determine the temporal patterning of dyadic social behavior and the first to do so in play, providing a quantitative assessment of the ontogeny of social play behavior in captive timber wolves, Canis lupus, by following 6 dyads from age 2 to 30 weeks. Play's temporal characteristics were measured at several basic levels: the duration and patterning of all play, as well as the proportion and duration of its categories and types. These properties were then examined with regard to the individuals' development. By defining 8 play categories and types, play was also examined at a broader level than previously had been done for wolves. Both play's total duration and bout duration showed definite changes with the pups' development, as did the play categories and types. Changes in all measurements occurred rapidly until the pups were about 12 weeks old, after which few changes occurred. The appearance and proportions of play categories and types, as well as changes in bout length and total duration of play, indicated both increased physical ability and maturing social skills.

Weaver, John Lee. ECOLOGY OF WOLF PREDATION AMIDST HIGH UNGULATE DIVERSITY IN JASPER NATIONAL PARK, ALBERTA (CANIS LUPUS). Dissertation Abstracts International. Volume: 55-12, Section: B, page: 5146. Abstract There have been few studies of wolf (Canis lupus) predation in multi-species prey systems characteristic of the Rocky Mountains. Therefore, I investigated the ecology of wolf predation amidst high ungulate diversity in NE Jasper National Park, Alberta, from 1989 to 1992. In winter, estimated populations of 420 bighorn sheep and 300 elk occurred in large groups mostly in patches of grassland habitat interspersed among coniferous forests. An estimated population of about 350 deer ($\ge$2 mule deer for every white-tailed deer) was dispersed more evenly in various cover types across the landscape. About 30 moose occurred as singles or pairs in riparian shrub types or scattered through upland pine or spruce-fir forests. Amidst this rich diversity of ungulates, a pack of 4-9 wolves in winter preyed more than expected upon deer (especially white-tailed deer and male individuals of both species) and used moose, elk and bighorn sheep less than expected. With ungulate numbers considered in terms of groups, however, the wolf pack preyed upon ungulates in proportion to availability (i.e., no selectivity). Adult deer and elk comprised most of the biomass in the wolves' winter diet, yet the proportion of biomass of elk and bighorn sheep (especially adults) consumed was less than that available. Kill sites of deer were characterized by greater relative density of elk and moose than typical for the landscape. During the denning period, the wolf pack selected deer fawns and fed on adult elk and bighorn sheep less than expected; wolves continued to use the different species in proportion to availability of groups. During the rendezvous period, the pack switched from deer to calf and adult elk (about a 20% shift) with significant selectivity for elk calves. A re-examination of 9 studies across North America involving wolf predation amidst high ungulate diversity revealed that wolves preyed principally upon elk and deer. Elk represented the most profitable prey with minimal risk-of-injury whereas wide dispersion of deer facilitated frequent encounters. In this multi-species context, moose were less profitable due to risk-of-injury; opportunities for wolves to find bighorn sheep away from escape terrain seemed infrequent. Small group size, wooded or hilly terrain, and deep snow fostered wolf selection of elk. When elk were not as vulnerable, wolves preyed more intensively upon deer. I propose that wolves living amidst high ungulate diversity forage as 'expanding specialists' whereby they specialize on vulnerable individuals of large prey (elk and moose) yet readily generalize to common prey (deer). In a diverse prey system, wolf predation may shift among species depending upon annual fluctuations in winter severity or landscape changes. Wolf densities lower than predicted by Fuller's (1989) equation were all characterized by high ungulate diversity in which at least one of the prey species occurred in large groups.

Forbes, Graham John. WOLF-UNGULATE WINTER RELATIONSHIPS IN ALGONQUIN PARK, ONTARIO (CANIS LUPUS, ALCES ALCES, ODOCOILEUS VIRGINIANUS, DERMACENTOR ALBIPICTUS, CASTOR CANADENSIS). Dissertation Abstracts International. Volume: 55-08, Section: B, page: 3107. Abstract The interactions of a wolf-moose-deer-beaver predator-prey community were studied from 1987-1992 in Algonquin Park, Ontario, Canada. The goal of the study was to identify the impact gray wolves (Canis lupus) had on a multiple-prey system where the two largest prey species fluctuate in winter availability. The density, food habits, and movements from 57 radio-collared wolves were compared within three regions of varying ungulate abundance. Relative moose (Alces alces) and white-tailed deer (Odocoileus virginianus) availability was determined annually. The amount of deer in winter scats (12.3-32.5 percent) and the number of deer predation cases (n = 50 carcasses) was higher than expected in all three regions, even in areas where deer were considered to be very rare. Wolf use of deer was positively correlated with increases in winter deer abundance (r$^2$ =.66). Wolf density correlated with deer numbers (r$^2$ =.74). The winter consumption of moose by wolves mainly was derived from scavenging animals that had died in association with infestations of winter tick (Dermacentor albipictus). Beaver (Castor canadensis) apparently were important buffer prey species when deer numbers were low, or when deer seasonally emigrated from a wolf territory. Consumption of beaver, up to 15.1% of total winter prey biomass, was high in winter. Territory size of wolf packs studied averaged 149 km$^2$. Mean pack size in December was 6.0 wolves/pack and within-territory density was 4.3 wolves/100 km$^2$. Population density was estimated to be 2.7 wolves/100km$^2$. Extraterritorial movements were often related to deer migration to a wintering yard located outside of Algonquin Park. An overwintering deer density of less than 0.02 deer/km$^2$ appears to have prompted the emigration of wolves to the deer yard. A total of 37 non-dispersal movements were recorded in 2 years (1990-1991, 1991-1992). Wolf movements were more extensive during the winters of 1990-1991 and 1991-1992 when few deer wintered in the Park. The annual mortality rate of wolves was 30-37%. The greatest cause of mortality was hunting and snaring (45%), and rabies (21%). Most mortality occurred outside of the Park (62%), often in association with winter movements by wolves out of the Park in search of white-tailed deer. Wolves were shot incidentally by hunters in pursuit of deer or were snared by local residents living in an area that is a winter concentration area (deer yard). Assessment of the total mortality was hindered by rumoured wolf kills and collars which were damaged by people killing the wolf. Trapping records for the vicinity adjacent to the Park indicated a consistent trapping pressure and trapping levels dependent upon the changing financial value of the pelt. The protection and management of wolves in Algonquin Park is presented in terms of cross-boundary issues and the proposed re-introductions of wolves into other park environments. In 1993, based on mortality levels and the value of wolves to the Park, managers decided to protect wolves from excessive mortality during winter months in the main area used by wintering deer outside the Park. The Carnivore Conservation Area strategy is introduced as a framework for incorporating wolves into a regional cross-boundary approach for managing this wolf population. (Abstract shortened by UMI.).

Goldman, Jill Andrea. AN ACOUSTIC BASIS FOR MATERNAL RECOGNITION IN WOLF PUPS (CANIS LUPUS). Masters Abstracts International. Volume: 32-06, page: 1701. Abstract A litter of four wolf pups was born on May 10, 1990. Pawnee (the mother) and Ursula (adult female) entered the den on different occasions and attended to the pups during the first five postnatal weeks. The adult squeak vocalization often accompanied specific caregiving activities. Squeaks from outside the den began when the pups were three weeks old. Pups often exited the den in response to squeaks but it was apparent that not all squeaks elicited pup-exit responses. Squeaks were grouped into four conditions: (1) Pawnee's squeaks inside the den, (2) Ursula's squeaks inside the den, (3) squeaks from outside that elicited pup-exit responses and, (4) squeaks from outside that elicited pup no-exit responses. Fundamental frequency (f$_( o)$) was exposed as one acoustic parameter that distinguishes the two senders' squeaks and could potentially be used by the pups to facilitate auditory recognition of their mother. Chi square analyses confirmed that; (1) two adults can be distinguished based solely on their f$_( o)$ distributions, and (2) the squeak f$_( o)$ distribution that pups exited to was significantly different from that which they did not exit to. Evidence supporting this acoustic basis for mother recognition came from the comparison between the squeak f$_( o)$ distribution that elicited pup exit responses and Pawnee's f$_( o)$ distribution. The former was wholly subsumed by the latter, and was concentrated at the lower end of Pawnee's distribution. This concentration may reflect the pups' response bias toward those of their mother's squeaks whose f$_( o)$s were most likely to distinguish her from other attendant wolves.

Lamothe, Alain R. WINTER FOOD HABITS AND FOODCHAIN TRANSFER OF METALS IN WOLVES (CANIS LUPUS) OF THE KEEWATIN DISTRICT, NORTHWEST TERRITORIES. Masters Abstracts International. Volume: 32-06, page: 1586. Abstract Winter feeding habits of wolves (Canis lupus) from the Keewatin District, Northwest Territories, were examined through analysis of stomach contents. Carcasses were collected from Inuit hunters at four different locations (Eskimo Point, Rankin Inlet, Baker Lake, and Repulse Bay) during the winters of 1987 to 1989. A total of 237 stomachs were analysed. Observations indicate that caribou (Rangifer tarandus) constituted the most frequent item (92.7%), while moose (Alces alces) remains were less frequent (5.3%). Other items present in low frequencies were muskoxen (Ovibos moschatus), wolf, birds, grasses, and indigestible material (rocks, garbage, and woody shrubs). Geographic differences were noted and appeared to be related to latitudinal ranges of prey species. Whereas caribou comprised an almost exclusive food source for wolves in central and northern portions of the study area, moose constituted a secondary prey species utilized by wolves to the south. On both a weight and volume basis, caribou constituted the principle (94.6% and 94.8%, respectively) food item present in the stomach contents. Temporal changes were noted in the frequency with which caribou were preyed upon by wolves. Empty stomachs occurred in approximately 40% of the animals harvested each year suggesting that a substantial portion of the population had not fed for at least several hours prior to collection. Tissue metal (cadmium, lead, nickel, zinc, copper, and iron) burdens were examined in the above wolves and other key components of this Arctic foodchain (soil-vegetation-caribou-wolf) in an attempt to assess the extent of environmental contamination and trophic bioaccumulation within the Keewatin District. Female caribou sampled near Eskimo Point showed tissue metal levels typical of values reported for other ungulates from relatively non-contaminated areas. Metal levels in soils, lichens, and willow sampled along a transect northwest of Eskimo Point were generally low. (Abstract shortened by UMI.).

Levi, Uriel Abraham. THE SPATIAL LOGIC OF ANIMAL CULTURE: IMPLICATIONS FOR BIOEXHIBIT DESIGN (WOLVES). Dissertation Abstracts International. Volume: 55-01, Section: A, page: 4613. Abstract The study of wolf (Canis lupus) socio-space suggests that members of a wolf pack distribute themselves in both linear (axial), and clustered (convex) arrangements. These arrangements appear to be linked to the ability of wolf packs to function as coordinated hunting units in an unpredictable environment. Convex arrangements facilitate the intense social interaction necessary to establish and enforce role relationships within the pack. Axial arrangements reinforce these relationships and facilitate the environmental awareness necessary for effective hunting. The topological nature of these patterns provides the foundation for reducing the scale of habitats in artificial exhibits, while preserving their socio-spatial properties. Bio-exhibit site-plans that provide wolf packs with opportunities to move between convex and axial arrangements, will correspondingly provide them with opportunities to exercise species-appropriate social behavior. Further, the configurational correlates of this social behavior establish the foundation for developing visitor interfaces that make this social behavior accessible, thereby, providing visitors with a better understanding of the wolf as a social animal.

Ballard, Warren Baxter, Jr. DEMOGRAPHICS, MOVEMENTS, AND PREDATION RATES OF WOLVES IN NORTHWEST ALASKA (CANIS LUPUS). Dissertation Abstracts International. Volume: 54-11, Section: B, page: 5458. Abstract During 1987 through 1992, 85 wolves (Canis lupus) were captured, radio-collared, and relocated from aircraft 1,123 times in northwest Alaska. Wolf packs usually did not follow migratory caribou (Rangifer tarandus) but maintained year-round resident territories that averaged 3,652 km$^2.$ During years when caribou were absent and moose densities were low, $\le$25% of the wolf packs moved 64 to 272 km to the caribou wintering grounds. Wolves used different slopes, aspects, and habitats in summer versus winter. Twenty-five percent of the radio-collared wolves dispersed. Annual finite rates of increase ranged from 0.64 to 1.43. Annual wolf survival rates averaged 0.59. There were differences in survival rates among years. Sixty-one percent of the wolves died. Hunting was the main cause of death (69%) followed by rabies (21%). Rabies was a significant natural limiting factor. This wolf population could sustain mortality rates of about 53% annually. Caribou and moose composed 51 and 42%, respectively, of the observed wolf prey. Adjusted for prey size, each pack killed 1 adult moose equivalent per 6.7 days. Wolf pack sizes and adjusted kill rates and kgs of available prey per wolf per day were correlated from several areas across North America. When caribou were present they were the principal prey. However, when caribou densities were $<$100/1,000 km$^2$ wolves preyed upon moose. Wolves preyed upon relatively healthy caribou and moose that were in marginal condition. Wolves were killing about 6-7% of the caribou herd and from 11 to 14% of the moose population annually. Existing wolf predation may have serious impacts on resident, low-density moose populations. During spring 1990 I tested the line-intercept method of sampling tracks for estimating wolf densities for a known wolf population (i.e., 48 wolves). The population estimate based upon line-intercept sampling was 50.7 (80% CI = 33.4 to 67.9) suggesting that the survey method provided relatively accurate population estimates. I placed 23 satellite transmitters on wolves aged 10-months to 8 years with no apparent adverse effects on them. Accuracy of 1,855 relocations at 9 sites averaged 336 and 728 m for best and worst quality relocations, respectively. Satellite telemetry has potential for providing improved data sets for evaluation of wolf territory sizes and movements.

Dale, Bruce Williams. WOLF-CARIBOU RELATIONSHIPS IN A MULTIPLE UNGULATE PREY ECOSYSTEM (CANIS LUPUS, RANGIFER TARANDUS, ALCES ALCES, OVIS DALLI). Masters Abstracts International. Volume: 31-04, page: 1655. Abstract Winter wolf (Canis lupus) predation and functional response in wolf - caribou (Rangifer tarandus) dynamics were investigated in a multiple ungulate prey ecosystem in Gates of the Arctic National Park, Alaska. Prey selection, prey availability, prey switching, kill rates, and food availability for 4 wolf packs were estimated in March 1989, March 1990, and November 1990. Estimates for these study periods reflected near record, average, and early winter snow conditions, respectively. Wolves killed predominately caribou even if moose (Alces alces) or Dall sheep (Ovis dalli) were more abundant. Prey selection varied with study period; however, per wolf kill rates and food availability did not. Length of intervals between kills was correlated with pack size and the biomass of the previous kill. Kill rates indicated a destabilizing Type II functional response. Modeling with a linear numerical response revealed wolf predation to be an increasingly important limiting factor at low caribou densities. However, little potential for regulation of caribou by wolves was observed.

Woods, John G. ECOLOGY OF A PARTIALLY MIGRATORY ELK POPULATION (CERVUS ELAPHUS, ALBERTA). Dissertation Abstracts International. Volume: 53-11, Section: B, page: 5596. Abstract In this thesis I investigate the ecology of a partially migratory ($<$100% of the animals migrate) population of elk (Cervus elaphus) in the Canadian Rockies. I raio-tagged elk in a 330 km$^2$ area of the Bow River valley (BRV) in Banff National Park, Alberta and followed them for 36 months. Elk movements to adjacent watersheds expanded the study area to 2900 km$^2$. My goals were to describe the seasonal movements and made by BRV elk and to reach some understanding of the causes of these movements. The M/R (migrant/resident) ratios for adult bulls and adult cows were 4.3 and 0.5 respectively. Classified ground counts revealed that adult bulls made up only 11% of the population and that the overall M/R ratio for the population was 0.6. Migrations did not take elk beyond the foraging range of timber wolves (Canis lupus), their principal predator. Three cows changed migration status between years and some migrants were sympatric with residents during the rutting (breeding) season. These observations in elk is a conditional ESS (evolutionarily stable strategy). Although 1 adult radio-tagged bull dispersed, individual annual home-ranges of the remaining elk overlapped from year-to-year. Elk were strongly philopatric to winter, summer, and rutting ranges. There were no differences detected in the relative philopatry of bulls and cows, or of migrants and residents. Bulls had average 1-way migrations of 30 km horizontally and 840 m vertically. Cows had average 1-way migrations of 36 km horizontally and 1079 m vertically. The net energy and time investments for these movements were calculated and judged to be trivial. Elk on both high and low elevation ranges ate primarily grasses and sedges during the autumn, winter, and spring, and the leaves of deciduous shrubs during the summer. Similarities outweighed differences between high and low elevation ranges. Elk population characteristics (numbers, composition, survival, recruitment, predation, parasites, animal morphology) were measured during 1985-1991 and compared with similar data gathered during 1944-69. In most respects, the population has not changed over these years and little is known of density-dependent processes.

Hillis, Tracy L. THE DEMOGRAPHY AND ECOLOGY OF THE TUNDRA WOLF, CANIS LUPUS, IN THE KEEWATIN DISTRICT, NORTHWEST TERRITORIES. Masters Abstracts International. Volume: 30-04, page: 1173. Abstract Four hundred and twenty-five wolf carcasses were studied. Males were larger than females in most features. Differences in sexual dimorphic development were consistent with the hypothesis that female wolves partition resources differently than males. Seasonal changes in condition indices indicated that the timing of fat deposition in females differed from that in males. Condition indices in males were maximal in mid-winter, while female fat deposits increased throughout the fall and continue to develop into the spring. Yearly variation in wolf reproduction is predominantly controlled by changes associated with implantation and foetal survival in utero. The yearly changes in age structure was not explained by hunting, but appeared to represent an expanding predator population responding to high prey densities. Differences in wolf abundance during different phases of the prey cycle suggest that the wolf population numbers fluctuate in association with primary prey, further supporting the hypothesis that wolf populations fluctuate in response to these populations. (Abstract shortened by UMI.).

Williams, T Mark. SUMMER DIET AND BEHAVIOUR OF WOLVES DENNING ON BARREN-GROUND CARIBOU RANGE IN THE NORTHWEST TERRITORIES, CANADA. Masters Abstracts International. Volume: 30-03, page: 0667. Abstract Wolves (Canis lupus) denning on the ranges of the Bluenose and Bathurst barren-ground caribou (Rangifer tarandus groenlandicus) herds in the Northwest Territories relied mainly on caribou as a food source during the denning period. Adults ate small mammals and birds more often than their pups. On the range of the Bluenose herd, 19 of 20 dens found by the location of radio-collared wolves were in the boreal forest-tundra transition zone, consistent with reports that most wolves den along tree line. General habitats and the use of vegetative cover varied between these dens; however all dens were dug into sandy soil. I recorded the activities of wolves associated with two different dens. More food was available to the adults observed in 1987 relative to those in 1988 based on the rate at which food was returned to the den. Associated with differences in food delivery rates were differences in activity patterns of adults and pups, rates of play behaviour, the tendency of non-breeding adult wolves to help, and over-summer pup survival. (Abstract shortened by UMI.).

Jhala, Yadvendradev Vikramsinhji. HABITAT AND POPULATION DYNAMICS OF WOLVES AND BLACKBUCK IN VELAVADAR NATIONAL PARK, GUJARAT, INDIA. Dissertation Abstracts International. Volume: 52-04, Section: B, page: 1799. Abstract The problem of allocating limited resources for conflicting needs of 2 endangered species, Canis lupus pallipes and Antelope cervicapra at Velavadar National Park was addressed by optimization models. Data were collected from February 1988 to December 1990. The semi-arid grassland was dominated by Dichanthium annulatum and Sporobolus virginicus. Two-year rainfall sequence explained 96% of the variability in the peak above-ground plant biomass. Blackbuck relied primarily on grasslands for food. Pods of the4 exotic Prosopis juliflora comprised 10% of the diet and were important during summers and drought years. Dry matter intake, estimated for capative blackbuck feeding natural forage, dropped in summers and increased during monsoons and witers. Average annual intake was 86 gms/kg$^(0.75)$/day. Forage digestibility and quality were low in summers. Fecal ether extract and fecal nitrogen were useful for predicting forage quality. Droughts and floods caused blackbuck population declines. The population takes 4-5 years to recover. Life-table analysis revealed a bimodal mortality pattern for males. Males between 7 to 9 years were at a high risk of wolf predation. Wolves used to shrubland for lying-up, denning and for rendezvous sites. Important wolf habitat was marked on a habitat map of the park. Average pack size was 6 wolves. Litters were born in December-January. Mean litter size was 5.5 pups. Pup mortality was estimated at 28%. Food habits of wolves was determined from scats. Blackbuck were the major prey (88%) followed by rodents (3.3%) and hare (2.3%). Consumption, estimated from fresh kills, was 0.07 kg/kg of wolf/day. A ratio of 158 blackbuck to 1 wolf was considered essential to sustain predation. The objective of increasing the forage-based carrying capacity of the park was optimized by linear programming. Limitations of the annual budget, park area, wolf habitat needs, stochastic rainfall and associated forage production, and desired wolf: blackbuck ratio were model constraints. The optimization results recommended maintaining grasslands free of Prosopis, to cut shrublands in areas other than critical wolf habitat for increasing grasslands, and to develop saline regions into shrublands. Managing well-dispersed, pruned, Prosopis groves would provide pods for blackbuck, wolf habitat, and allow grass growth. A 25-30 year shrub rotation was considered appropriate for improving the saline alkali soils.

Hiestand, Nancy Laura. A COMPARISON OF PROBLEM-SOLVING AND SPATIAL ORIENTATION IN THE WOLF (CANIS LUPUS) AND DOG (CANIS FAMILIARIS). Dissertation Abstracts International. Volume: 50-11, Section: B, page: 5371. Abstract In the 10-15,000 years since the domestication of the dog from the wolf, the dog has been molded by artificial selection to conform to the needs of humanity, whereas the processes of natural selection have resulted in current wolf behavior. To better understand the behavioral differentiation of the two species, two adult and seven six and one-half month old hand-reared timber wolves were compared with forty adult German Shepherd dogs on their ability to manipulate objects sequentially and to orient in three-dimensional space. The initial task was to pull a single rope suspended from the ceiling. In later tests this problem increased in complexity by the addition of ropes and by changing the spatial configurations. Adult wolf performance was consistently successful across all tests and requirements. Juvenile wolves had little difficulty with one and two rope tests, but showed a decrement in ability on the three rope test. The behavior of the dogs segregated into four profiles: Non-Responders (6 dogs), One Rope Responders (15 dogs), Two Rope Responders (14 dogs), and Three Rope Responders (5 dogs). It has often been proposed that one result of artificial selection on the dog is that it retains characteristics that are typical of juvenile wolves into adulthood. This makes the dog in effect a "neotenous wolf". This study was the first to compare juvenile and adult wolves with adult domestic dogs on two putative neotenous behaviors: complex problem solving and spatial orientation. Indeed, on the most difficult task, the three rope test, adult dogs did perform like juvenile wolves. However, many other behavioral characteristics either are the same as, or are accelerated in, the dog when compared to the wolf. Thus I must agree with Zimen (1981) and J. P. Scott (personal communication) that in general the dog cannot be considered a neotenous wolf.

Page, Richard Edward. THE INVERTED PYRAMID: ECOSYSTEM DYNAMICS OF WOLVES AND MOOSE ON ISLE ROYALE (MICHIGAN). Dissertation Abstracts International. Volume: 50-09, Section: B, page: 3820. Abstract Thirty years of studying the wolf-moose ecosystem on Isle Royale National Park, Michigan, USA has presented an unprecedented opportunity to empirically test concepts in ecology and to advance the understanding of density-dependence, predator-prey interactions and the stability of ecosystems. Many of the population regulation processes on the island are similar through 3 trophic levels after accounting for the expected differences of scale. Wolves (Canis lupus lycaon) are essentially self-regulated below the limits set by the availability of their primary prey, moose (Alces alces andersoni). The size of a wolf pack is determined by the summation of individual dispersal strategies of its members. Since wolf populations are ultimately regulated by their food supply, at high moose densities they establish a social regulation by setting pack territory density initially below a food limitation. Moose are regulated both by predation from the trophic level above, and by food availability in the level below. Age structures reconstructed from dead moose confirmed an increase in mean age and also indicated a predator pit at low density in the 1950's, as suggested from mainland studies. Decades later, rapidly escalating functional and numerical responses of wolves at high moose density result in a truncation of the moose recruitment curve, promoting instability and making cyclic population fluctuations likely. Moose predation on balsam fir resulted in long-term recruitment failure and an over-mature age structure as in the moose population itself. It would be expected that survival rates would increase and reproductive rates decline with steps up the trophic pyramid. Longevity is actually greatest in the fir population, then moose, then wolves. The inversion of rates of increase and other population parameters in the Isle Royale wolf-moose-fir ecosystem is primarily responsible for the dramatic cyclic behavior seen in this system. The organizing influence of predation in population regulation and community dynamics is clearly evident. The majority of North American moose populations are in a predator pit induced by wolf predation well below a moose food limitation. (Abstract shortened with permission of author.).

Paquet, Paul Charles. BEHAVIORAL ECOLOGY OF SYMPATRIC WOLVES (CANIS LUPUS) AND COYOTES (C. LATRANS) IN RIDING MOUNTAIN NATIONAL PARK, MANITOBA. Dissertation Abstracts International. Volume: 50-07, Section: B, page: 2803. Abstract Although wolves and coyotes appear to be sympatric throughout portions of their geographic ranges little is known about their spatial interactions. Most researchers have implied that in regions where both species occur, coyotes either avoid the areas where wolves are most active or are displaced by wolves. Spatial segregation has been attributed to antipathy of wolves toward coyotes as well as inability of coyotes to compete successfully for essential resources such as food. In Riding Mountain National Park wolves (Canis lupus) and coyotes (C. latrans) overlapped temporally and spatially. The movements of coyotes relative to wolves were not random. Coyotes sought out active wolf areas, and followed wolf tracks rather than avoiding them. The movements of wolves were not altered by the presence of coyotes. Urine marking behavior of the two species was similar with both canids using the same scent mounds. Coyotes increased their rate of marking significantly in response to wolves. Wolves were less responsive to coyotes. Wolves did not use urine marks to demarcate territorial boundaries. The behavior does not appear to be a consequence of sympatry with coyotes. Competition between wolves and coyotes was minimized by differential use of the available food resources. Wolves preyed primarily on elk (Cervus elaphus), and secondarily on white-tailed deer (Odocoileus virginianus), and moose (Alces alces) yearlings. Coyotes preyed on deer and rarely, young elk. Although wolves occasionally killed coyotes, coyotes followed wolves and scavenged at their kills. The benefits accrued from scavenging apparently compensated for the associated risks. This relationship could change if the nature of the food supply is altered. A hypothesis is proposed that suggests the association, and therefore the distribution, of wolves and coyotes is influenced primarily by: (a) the availability and use of large ungulate prey species by wolves, and (b) the availability of wolf-provided carrion for coyotes to scavenge. It is concluded that the proximate reason why coyotes are excluded from some areas occupied by wolves is that they are unable to scavenge for food, and secondarily because of direct aggression by wolves.

Crabtree, Robert Lawrence. SOCIODEMOGRAPHY OF AN UNEXPLOITED COYOTE POPULATION (WASHINGTON). Dissertation Abstracts International. Volume: 50-06, Section: B, page: 2253. Abstract We estimated socio-demographic parameters of a natural, unexploited coyote (Canis latrans) population in the shrubsteppe of southcentral Washington from 1984 to 1988. A large proportion of the population was sampled (145 individual captures). Adult coyotes were categorized by social class based on physical characteristics, space use, site fidelity, movements patterns, and social interactions with other coyotes. Adult coyotes exhibited a high degree of spatial structuring according to well-defined social classes and subclasses. Territorial social groups (67% of the spring prehelping population) contained an older-aged alpha pair and an average of 1.4 young adult associates. Loners or nonterritorial individuals comprised 33% of the spring population and were divided into 2 major social classes: solitary residents (18%) with younger and older subclasses and nomads (15%). The average range size of social group members, solitary residents, and nomads was 14.5, 54, and 220 km$^2$, respectively. Territories were contiguous, non-overlapping, and uniform in size. Loners were located on the periphery of and in narrow corridors between territorial ranges. A healthy class of 2 to 4 year-old mature loners form a reservoir of potential territorial replacements, followed by a young group of 1 and 2 year-old nomads. Additional intraspecific "pressure" may result from old-aged solitary residents that were former territorial owners. Successful breeding occurred at ages 2 through 6 for alpha females ranging in ages 2 through 11 years. Reproductive failure among territorial groups was 27% and mainly involved breeding adults age 6 years and older. Population productivity was very low at 0.62 pups per female, whereas average litter size was 5.6. Pups weighed 27% below average and incurred a 58% mortality rate during the first 14 weeks following birth. The overall annual survival rate was 0.90 and was strongly a function of social class (P $<$ 0.001). Emigration from the population was low (16%) and consisted primarily of pups and associates. Immigration was low (6%) and limited mainly to younger nomads. Density averaged 0.41 and 0.38 coyotes per km$^2$ for an absolute density estimate and complete enumeration, respectively. The ALE coyote population was stable and appeared to have a high degree of intraspecific strife. We propose a theory of population regulation similar to that described for wolves (Canis lupus) and discuss the possible compensatory responses to exploitation.

Jenks, Susan Margaret. Title BEHAVIORAL REGULATION OF SOCIAL ORGANIZATION AND MATING IN A CAPTIVE WOLF PACK. Dissertation Abstracts International. Volume: 50-05, Section: B, page: 1739. Abstract Reproduction is restricted in wolf (Canis lupus) packs: generally a single female breeds successfully each season. This study examines patterns of agonistic relationships among individuals, and the influence of these social dynamics on intrasexual competition, mate preferences, and the reproductive success of individuals in a family group of wolves. The pack was studied over the final four years, 1980-1984, of a nine year project that was initiated in 1975 when a pair of wolves and their first litter of offspring were released into an 80 x 80 foot enclosure. Distributions of aggressive, submissive, defensive and sexual behavior were examined to determine behavioral profiles of individual wolves, dominance relationships among individuals, and patterns of mating behavior, during the four three-month breeding seasons encompassed by the study. Pack social structure was analyzed in terms of a dominance hierarchy and social differentiation among individuals. The influence of social structure on patterns of mating behavior were examined in terms of age, sex, and the changing social and genetic context of the pack over the four years. During this study, agonistic relationships regulated mating behavior. The dominant male and female were the only wolves observed to breed in each year. High-ranking wolves interfered more often with the sexual activity of others, emitted and received the highest frequencies of sexual behavior, and participated more frequently in aggressive coalitions. Transition from a pack characterized by parents and their offspring to one characterized by siblings and the offspring of one pair of those siblings resulted in increased levels of intrasexual competition. The sexual behavior of subdominant females was severely inhibited. During the course of this study, changing social dynamics resulted in social status transitions and breeder rotation. The dynamics of these transitions involved successful offspring to parent, and sibling to sibling challenges. The relationship between these interactions and the reorganization of the social and breeding structure of this pack suggests that the nuclear family sociogenetic context of wolf packs is a directing force for both stability and change and regulation of reproduction among pack members, based on feedback systems involving age and long-term social relationships.

Schotte, Cynthia Strom. THE DEVELOPMENT OF SOCIAL ORGANIZATION AND MATING PATTERNS IN A CAPTIVE WOLF (CANIS LUPUS) PACK. Dissertation Abstracts International. Volume: 49-12, Section: B, page: 5110. Abstract A pair of zoo wolves and their eight week old pups were released in an 80 x 80 foot enclosure with an opaque perimeter fence and were observed for five years. They were fed daily and had continuous access to water. They were not handled or tamed and disturbances were held to a minimum. Some or all of the pups from the single litter born each spring were culled from the group. Observations were made regularly, primarily by the author. During the peak of the breeding season, the wolves were under almost continuous observation during daylight hours. At other times of the year, they were observed five or six days a week on a sampling basis during periods of peak activity. Complementary and overlapping methods were used to record the observations. The methods, which were often used in combination, sometimes by several observers working simultaneously, included: written notes; dictation onto cassette tape; and motion picture or videotape recordings of samples of complex interactions. Every fifteen minutes, the location and activity of each animal was indicated on a schematic diagram of the compound. More than, 5,500 pages of written data and 30 hours of film were generated. For the purpose of this dissertation, the data for the five breeding seasons were subjected to detailed analysis and supplemented by data collected outside the breeding seasons. The results showed that the pack organization did not mature until the breeding season in which the first litter experienced puberty (as two year olds). At that time, the original parents (designated as X and Y) became the dominant female and male, and they held those positions throughout the five years. X was the pack's leader. The mature pack organization was characterized by affiliation, cooperation, leadership, social roles and restricted breeding. All pubertal and postpubertal females showed external signs of estrus each winter but only X displayed sexual behavior and mated. All pubertal and postpubertal males showed sexual behavior each winter. Y and two of his sons mated. X preferred Y in all breeding seasons but did not mate exclusively with him.

Fuller, Todd King. WOLF POPULATION DYNAMICS IN NORTHCENTRAL MINNESOTA. Dissertation Abstracts International. Volume: 49-06, Section: B, page: 2040. Abstract During 1980-1986, 81 wolves (Canis lupus) were radiomonitored in northcentral Minnesota. Winter territories averaged 78-153 km$^2$ and all had $\le$0.72 km/km$^2$ of drivable roads. In winter, mean pack size ranged from 4-9, lone wolves comprised 20-30% of the population, and annual density averaged 33-69 wolves/1,000 km$^2$. Annual finite rate of increase averaged 1.04 and was correlated with mean number of pups per pack in fall. Litters averaged 6.1 pups at birth and 3.2 by mid-November, then comprising 46% of pack members. Annual survival of pups was 0.40. Annual survival of radiomarked resident wolves (0.67) was greater than for non-residents (0.50). Most deaths (80%) of these wolves were caused by humans but some (10%) were due to other wolves. From 0 to 30% of radiomarked wolves left their home territories each month, most often during September-March; wolves left temporarily 0 to 6 times before permanently dispersing. Modeling indicated that 24% of adults, 66% of yearlings, and 8% of pups dispersed each year. Wolves may increase their reproductive fitness by dispersing as yearlings, but remaining in packs as older adults. Relatively more fawns (54%) and old (median age = 9.5 years for females, 3.5 years for males) white-tailed deer (Odocoileus virginianus) were killed by wolves in winter (n = 74). Estimated kill rates in January-February (x = 22 days/kill/wolf) did not vary with snow depth. Beaver (Castor canadensis) occurred in 20-47% of scats during April and May, but during June and July, newborn deer fawns occurred in 25-60%. Overall, deer provided 79-98% of biomass consumed each month; adult wolves consumed an estimated 19 deer/year, including 11 fawns. A consensus of studies indicate that wolf numbers are directly related to ungulate biomass; per capita biomass availability affects reproduction and survival. Annual rates of increase vary directly with mortality rates, and harvests exceeding 28% of the winter population often result in population declines. Summer food resources other than large ungulates may importantly affect pup survival, the major factor in wolf population growth. Equations useful for management incorporate wolf densities, annual ungulate kill/wolf, ungulate densities, potential rates of increase for ungulates, and harvest levels of ungulates by humans.

Gauthier, David Andrew. POPULATION LIMITATION IN THE BURWASH CARIBOU HERD, SOUTHWEST YUKON. Dissertation Abstracts International. Volume: 46-08, Section: B, page: 2545. Abstract A four-year field study (1978 to 1982) in the Kluane region of the southwest Yukon Territory (61(DEGREES)N, 139(DEGREES)W) focused on factors affecting changes in the abundance and distribution of a herd of caribou (Rangifer tarandus caribou). The research examined five predictions arising from the hypothesis that predation, primarily by wolves (Canis lupus), was the major limiting factor to herd growth. The influences of forage, climate and disease on herd growth were also assessed. Seasonal censuses (calving, rut and winter periods) and other surveys were undertaken to apportion changes in caribou numbers to changes in mortality, recruitment or ingress/egress, and to assess seasonal distribution. Thirty-nine caribou were radio-collared and used in capture-recapture analysis. Estimates varied from a minimum herd size of 330 to a maximum of 560. Calves comprised 20% of the herd in calving periods, 19% during ruts and 18% during midwinters. An average of 56% of calves died within five months of birth. Natural-mortality rates of adult/subadults varied between 6% and 9%. The results of visual examination of live-captured caribou (n = 39) and blood analyses (n = 15) showed no evidence of external disorders, viral antibodies, infections, nutritional deficiencies or anemias suggesting summer forage conditions were not resulting in deterioration of caribou physical condition. Caribou primarily occupied sedge/grass and shrub habitats over all seasons, increasing their occupancy of sedge/grass habitats from calving to rut and winter. An analysis of fecal pellets showed that caribou primarily consumed shrub species, but substantially increased their use of lichens in winter. Two wolf packs varying in total number from ten to fourteen animals preyed primarily on ungulates (65% of all prey consumed) and secondarily on small mammals (35%) over all seasons. The biomass consumption of prey determined through wolf fecal analysis showed that caribou were the dominant prey type (35% of all prey consumed) followed by moose (30%) and hare (21%). Ungulate calves comprised only 20% of prey types in the calving period. Wolves consumed caribou and moose in proportion to their available biomass during calving but disproportionately consumed caribou in rut and winter. Wolf predation accounted for 56% to 64% on average of total annual caribou mortality over 1981 and 1982. It is concluded that predation was likely the most important factor responsible for limiting herd growth. Wolves were the dominant predator, principally important in reducing caribou numbers outside of the immediate calving period. (Abstract shortened with permission of author.).

Mcleod, Peter John. ASPECTS OF THE EARLY SOCIAL DEVELOPMENT OF TIMBER WOLVES (CANIS LUPUS). American Doctoral Dissertations. Source code: X1988.

Havkin, Zvika. FORM AND STRATEGY OF COMBATIVE INTERACTIONS BETWEEN WOLF PUPS (CANIS LUPUS). American Doctoral Dissertations. Source code: X1981.

Lockwood, Randall. AN ETHOLOGICAL ANALYSIS OF SOCIAL STRUCTURE AND AFFILIATION IN CAPTIVE WOLVES (CANIS LUPUS). Dissertation Abstracts International. Volume: 37-04, Section: B, page: 1961.

Mech, Lucyan David. THE ECOLOGY OF THE TIMBER WOLF (CANIS LUPUS, LINNAEUS) IN ISLE ROYALE NATIONAL PARK. Dissertation Abstracts International. Volume: 23-10, page: 3578.

Field, Rebecca. VOCAL BEHAVIOR OF WOLFE (CANIS LUPUS): VARIABILITY IN STRUCTURE, CONTEXT AND ONTOGENY. American Doctoral Dissertations. Source code: X1978.

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