Wolf History, Conservation, Ecology and Behavior
Selection of Den, Rendezvous, and Resting Sites by Wolves in the Bialowieza Forest, Poland
Theuerkauf, Jrn ; Rouys, Sophie ; Jedrzejewski, Wlodzimierz, Canadian Journal of Zoology, 81 (January, 2003): 163-167 (5 pp)

We studied wolf (Canis lupus) selection of 19 den, 10 rendezvous, and 31 resting sites found between 1986 and 2000 in the Bialowieza Forest (Poland). Our objective was to determine whether wolves selected sites far from villages, forest edges, and roads, and whether these sites had dense ground cover for concealment. We also tested whether wolves selected a particular forest type for their den sites. Den and rendezvous sites were located at greater distances from villages, forest edges, and intensively used roads than random points. Locations of resting sites were not affected by these manmade structures. Wolves selected dry coniferous forests for den sites but also used other forest types. We concluded that the suitability of an area for pup raising depended mainly on the spatial distribution of forest, human settlements, and public roads, and to a lesser extent on habitat characteristics.
A Simple Reason for a Big Difference: Wolves Do Not Look Back at Humans, But Dogs Do
Miklosi, A.; Kubinyi, E.; Topal, J.; Gacsi, M.; Viranyi, Z.; Csanyi, V., Current Biology, 13/9, April 2003, 763-766

The present investigations were undertaken to compare interspecific communicative abilities of dogs and wolves, which were socialized to humans at comparable levels. The first study demonstrated that socialized wolves were able to locate the place of hidden food indicated by the touching and, to some extent, pointing cues provided by the familiar human experimenter, but their performance remained inferior to that of dogs. In the second study, we have found that, after undergoing training to solve a simple manipulation task, dogs that are faced with an insoluble version of the same problem look/gaze at the human, while socialized wolves do not. Based on these observations, we suggest that the key difference between dog and wolf behavior is the dogs' ability to look at the human's face. Since looking behavior has an important function in initializing and maintaining communicative interaction in human communication systems, we suppose that by positive feedback processes (both evolutionary and ontogenetically) the readiness of dogs to look at the human face has lead to complex forms of dog-human communication that cannot be achieved in wolves even after extended socialization.
Simulating the Effects of Wolf-Elk Population Dynamics on Resource Flow to Scavengers
Wilmers, C.C., Ecological Modelling, 177/1-2 (September 2004), 193 (16pp)
The reintroduction of gray wolves to Yellowstone National Park (YNP) provides a natural experiment regarding the effects of top predators on scavenger species. Fieldwork on the Northern Range of Yellowstone indicates that wolves facilitate carrion acquisition by scavengers, but it is unclear whether this represents a transient or permanent effect of wolf reintroduction. Here we present a wolf-elk model with human elk harvest and use it to investigate the long-term consequences of predator-prey dynamics and hunting on resource flow to scavengers. Our model shows that while wolves reduce the total amount of carrion, they stabilize carrion abundance by reducing temporal variation in the quantity of carrion and extending the period over which carrion is available. Specifically, the availability of carrion is shifted from reliance on winter severity and elk density to dependence on the strength of wolf predation. Though wolves reduce the overall abundance of carrion by lowering the elk population this reduction is partially offset by increases in the productivity of an elk population invigorated by removal of the weakest individuals. The result of this is higher carrion production per elk in the presence of wolves. In addition, this yields an ecological explanation for the phenomena that predators increase the robustness of their prey: namely that by reducing the effect of density-dependent resource competition among elk, those that remain, even some of the older animals, are better fed and healthier as a result. Our model also suggests that human hunting has no effect on the distribution of carrion across the year but is crucial in determining the long-term abundance of carrion because of the effect of hunting on elk population levels. By reducing the proportion of cows in the annual hunt, which have historically been high in order to control the number of elk migrating north of the park, managers can allow an adequate supply of carrion....
Snowmobile Activity and Glucocorticoid Stress Responses in Wolves and Elk
Creel, S.; Fox, J.E.; Hardy, A.; Sands, J.; Garrott, B.; Peterson, R.O., Conservation Biology, 16/3, 809 (6pp)
The effect of human activities on animal populations is widely debated, particularly since a recent decision by the U.S. Department of the Interior to ban snowmobiles from national parks. Immunoassays of fecal glucocorticoid levels provide a sensitive and noninvasive method of measuring the physiological stress responses of wildlife to disturbances. We tested for associations between snowmobile activity of wildlife to disturbances. We tested for associations between snowmobile activity and glucocorticoid levels in an elk (Cervus elaphus) population in Yellowstone National park and wolf (Canis lupus) populations in Yellowstone, Voyageurs, and Isle Royale national parks. For wolves, comparisons among populations and years showed that fecal glucocorticoid levels were higher in areas and times of heavy snowmobile use. For elk, day-to-day variation in fecal glucocorticoid levels paralleled variation in the number of snowmobiles after we controlled for the effects of weather and age. Also for elk, glucocorticoid concentrations were higher in response to snowmobiles than to wheeled vehicles after we controlled for the effects of age, weather, and number of vehicles. Despite these stress responses, there was no evidence that current levels of snowmobile activity are affecting the population dynamics of either species in these locations.
Sound Science Is Never Enough: Ethics, Science and Values in Wolf Recovery
Lynn, W.S., World Wolf Congress 2003

Biologists note that the restoration of wolves is primarily a social as opposed to an ecological issue. Wolves will thrive wherever there is sufficient prey, habitat and freedom from human persecution. This is an insightful point, and becomes more powerful when we recall how ethical norms condition our willingness to live with wolves. Even so, there is a mismatch between the science and ethics of wolf recovery. Both scientific and policy discussions of wolf recovery tend to emphasize "sound science". Sound science is supposed to be the evidence-based, theory-rich baseline for managing wolves in both wild and humanized landscapes. Yet, as previously noted, humanity's trouble with wolves is deeply informed by conflicting values, which have little or nothing to do with empirical data, quantitative models, or management techniques. Instead, they are deeply rooted ethical conflicts over our coexistence with large predators. Society needs, therefore, a "sound ethics" to complement our science, as well as to guide our efforts in wolf recovery. A sound ethics should, at a minimum, recognize the moral standing of wolves, highlight the moral significance of the science-based management of wolves, and emphasize the practical value of ethics in the restoration of wolves.
Spatial Dynamics of a Migratory Wolf Population in Winter, South-central Ontario (1990-1995)
Cook, S ; Norris, D ; Theberge, John,  Canadian Journal of Zoology, 77 (November, 1999): 1740-1750 (11 pp)

We examined the spatial distribution and movements of migratory wolves (Canis lupuslycaon) to a deer yard located adjacent to Algonquin Provincial Park, Ontario, during 5 winters from 1990 to 1995. Wolves from eastern and central Algonquin Provincial Park followed the annual migration of deer to yards located 13 km outside of the Park boundary. Spatial distributions were determined through mapping of telemetry locations and nearest neighbour analysis. We defined three spacing systems: consistent/high fidelity, clustered/moderate fidelity, and transitional/low fidelity. We found inconsistencies among packs in their adherence to these systems. Data indicate that areas of use changed quickly and tolerance levels among wolves in the deer yard were very high; alien wolves were recorded 163 times in close spatial and (or) temporal proximity. The social behaviour exhibited by this migratory population of wolves has never been recorded in a forested wolf-deer ecosystem. Factors that may contribute to this behavioural plasticity include food abundance, a high degree of genetic relatedness among wolf packs, and high rates of human-caused mortality.
Spatiotemporal Segregation of Wolves from Humans in Bialowieza Forest (Poland)
Theuerkauf, J.; Jedrzejewski, W.; Schmidt, K.; Gula, R., Journal of Wildlife Management, 67/4, October 2003, 706-716

Knowledge about the impact of human activity on the behavior of wolves (Canis lupus) is important to predict habitats suitable for wolf recolonization and for planning management zones. We tested the hypothesis that wolves live spatiotemporally segregated from humans. From 1994 to 1999, we radiotracked 11 wolves in 4 packs and monitored human activity in the Bialowieza Forest, Poland. Wolves avoided permanent human-made structures (settlements, forest. edge to arable land, roads, tourist trails) more in the day than at night. Wolf avoidance increased with increasing human use. Particularly large settlements and intensively used public roads reduced the area used by wolves. Wolves avoided human presence in the forest (traffic, forestry operations) by temporarily selecting areas where people were absent. One of the wolf packs selected a national park zone with restricted access (50 km(2)) as the core area of its home range in both day and night. Conversely, wolf packs living in a commercial forest with small nature reserves (less than or equal to 4 km(2)) did not select reserves in the day or night. We concluded that spatiotemporal segregation is an adaptation of wolves to coexist with humans while keeping their activity pattern optimized toward food acquisition. The distribution of areas with restricted human access, forest, settlements, and intensively used public roads are important factors determining the suitability of an area for wolves.
Succession Debt: Effects of Logging on Wolf-Deer Dynamics in Coastal British Columbia and Southeast Alaska
Darimont, C.T.; Person, D.K.; Paquet, P.C.; Bowyer, R.T., World Wolf Congress 2003

Logging in the temperate rainforests of coastal British Columbia initiates a pattern of forest succession and road acess by humans that has long-term adverse consequences for wolves and their deer prey. We call this "succession debt." Twenty-five to 40 years after clear-cutting, regenerating stands of even-aged conifers grow into a "stem exclusion" stage characterized by a dense canopy and depauperate under-story, which substantially reduces carrying capacity for deer. Once initiated, changes are irreversible and cannot be mitigated by adjustments to future forest management. Moreover, logging roads facilitate access to wolf and deer habitat. Long-term effects include declines in populations of deer and wolves, increased exploitation of deer and wolves by humans, greater probability that wolves will suppress deer numbers, and increased likelihood of conflicts between hunters and wolves for deer. Moreover, island populations in this archipelago are more vulnerable to disturbance from logging and less likely to be readily recolonized. We compare wolf-deer systems from adjacent areas of coastal British Clumbia and southeast Alaska. In sparsely populated and nearly pristine coastal northern British Columbia, human-caused mortality of wolves is low, and prey populations are likely at historical levels. In contrast, Prince of Wales Island in southeast Alaska has undergone extensive timber removal. Killing of 30-40% of the wolf population annually is common and we predict substantial declines in deer populations over the next 50 years. Resource managers need to consider long-term consequences of forest management on predator-prey systems and recognize that mitigation of those consequences may not be possible.