<b>Theberge, J. B., and M. T. Theberge</b>. 2004. The Wolves of Algonquin Park: A 12-Year Ecological Study. University of Waterloo, Department of Geography, Waterloo, Ontario, Canada, 168 + v–xii pp. ISBN 0-921083-67-x, price (paper) $23.50 Canadian (approximately $25.00 US), order from S. Castela (scastela@fesmail.uwaterloo.ca).

This monograph on the gray wolf (Canis lupus) in Ontario's Algonquin Provincial Park summarizes results and implications of a 12-year study of the species in a park renowned as protected wolf habitat. In 1958, Algonquin Provincial Park became the 3rd area in North America where wolf populations were legally protected from harvest by humans (preceded by Denali [Alaska] and Isle Royale [Michigan] national parks in the United States). In this volume Theberge and Theberge present much original data from a field study conducted in 1987–1999, supplemented by inclusion of major portions of student theses that dealt with aspects of wolf ecology as well as previously published papers from peer-reviewed journals. This unusual format provides a comprehensive review of information gained by the authors during 12 years of fieldwork, as well as a comparison with data gathered by Douglas Pimlott and colleagues during a pioneering study in 1959–1963 (Pimlott et al. 1969), a study that laid the foundation for research that senior author John Theberge conducted with Pimlott in the 1960s on wolf vocalizations (Theberge and Falls 1967) and behavior in summer (Theberge and Pimlott 1969). This monograph was published by the University of Waterloo, not necessarily known as an outlet for major publications on mammals. However, there are good reasons why the authors may have chosen this nontraditional outlet for this significant publication on wolves. Probably most important is simply a desire to provide a single source for research findings that cover a wide range of topics in wolf ecology, including anecdotal information of the type that is rarely found in peer-reviewed journal papers. An additional reason is that the fieldwork was severely challenged because the work generated much politically charged policy discussion, elevating emotion and attention, both positive and negative. Among the acknowledgements of important contributions of field assistants, graduate students, pilots, funding sources, and the like, a university attorney also is mentioned, providing an indication of the level of controversy that surrounded this research. I suspect the authors wanted to present their comprehensive data and interpretation in 1 place without further delay. The significance of the study reported here is 2-fold. First, the study area contains a form of wolf that has helped launch a major resurgence of scientific interest in the ecological and genetic implications of hybridization among species of Canis. A new species, the eastern wolf (Canis lycaon), has been proposed (Wilson et al. 2000) that includes the Algonquin wolf, thought to be the original form of gray wolf that spawned dispersing populations to Eurasia in the Pliocene and itself survived in North America south of the Pleistocene ice sheets. This small form of wolf shows many ecological and genetic similarities to the endangered red wolf (Canis rufus), which historically occupied the southeastern Gulf Plain and much of the east coast of the present United States. The red wolf was endangered by hybridization with coyotes (Canis latrans); indeed, the species was declared extinct in the wild in the 1970s and rescued by a captive-breeding effort. Similarly, the putative eastern wolf is compromised by genetic hybridization with coyotes, especially to the south of Algonquin Park where intact forest gives way to mixed farm–forest habitats with high road density and greater human-induced mortality. Algonquin Provincial Park thus provides a critical area where an evolutionarily unique form of wolf survives in a reasonably secure wild habitat. The greater human impacts to the south of the park provide context for the 2nd significant aspect of this study. The research reported here provided much detail on the migratory behavior of wolves that developed in recent decades (after the study by Pimlott and colleagues), as wolves began to follow white-tailed deer out of the park to 2 deeryards adjacent to the park's southern edge. These changes coincided with a significant decline in deer in the park as forested habitats grew older. However, what resulted was a significant increase in human-caused mortality of wolves that were shot, trapped, and especially snared as they entered deeryards outside the park. The refugium that should characterize wildlife habitat in Algonquin Park turned out to be inadequate. Responding to the challenge stimulated by this research, as delivered by conservation organizations, high-level provincial politicians acted over a period of many years to provide a buffer outside of the park boundaries where wolves would be protected. The final political solution was a 6-mile-wide buffer strip surrounding the park where wolves were permanently protected, probably a 1st for parks or preserves around the world. The brief summary of these actions provided at the end of the monograph does not attempt to convey the acrimony and controversy that were part of the story, but the authors provide some of this in another book published earlier (Theberge and Theberge 1998). The most important data in this monograph are annual survival rates and causes of mortality recorded for 137 wolves that were radiocollared inside the park. In 10 of 11 years, mortality exceeded 25%, and ranged as high as 48%. About two-thirds of all wolf mortality was human-caused, especially in the townships south of the park. Fifty-eight percent of wolves killed by humans died in 2 deeryards outside the park. Wolf population density, which declined during the study, was maintained primarily by immigration. Chronic population decline continued at a slow rate, and a workshop conducted in 2000 by the Captive Breeding Specialist Group of the Species Survival Commission, International Union for the Conservation of Nature, concluded that the population was unsustainable in the absence of immigration. The implications of harvest for population dynamics are especially relevant in several states in the United States where the wolf has recently been removed (or proposed for removal) from the list of endangered species, areas where management by states will almost certainly expose wolves to increased harvest pressure. In Algonquin, human-caused mortality led to reduced wolf pack size—from almost 5 to 3 in 1989–1999 (pack size averaged 5.9 in 1959–1963), and wolf density declined because there was no compensatory increase in recruitment. Population density fluctuated annually depending on the level of human-caused mortality, and was unrelated to reproductive parameters. Included are interesting details about a significant outbreak of rabies and resulting mortality of wolves in 1990–1991, as well as a photograph of the heavily worn teeth of an old male that was estimated to be 15 years old, possibly the oldest documented wild wolf. The morphological data on eastern wolves, and the historical changes in size over a 35-year period, provide valuable historical context for this species of wolf. All skull measurements declined during 1963–1987, and body weights of females likewise declined. The authors speculate that increased genetic introgression of C. latrans during this period underlies the smaller body size they recorded. Nevertheless, Algonquin wolves are clearly larger than heavily hybridized Canis located south of the park. Using scats collected in Algonquin and in the region to the north of the park, the authors argue that wolves in the park population also are smaller than C. lupus to the north (contra Grewal et al. 2000), based on the assumption that scat size is proportional to body size. The volume might have benefited from more uniform peer review, which appears to have been limited to the published papers that are the foundation for parts of the monograph, and (to a lesser degree) unpublished university theses. Two examples will suffice. Yearling recruitment was calculated in a somewhat convoluted manner because of the small number of captures per year and the vagaries of age determination. Kill rates of moose, deer, and beavers were estimated indirectly by a long series of serial assumptions and calculations that started with prey remains in scats combined with assumed metabolic needs of wolves. The resulting wolf-induced prey mortality rates provided in this monograph, the holy grail of wolf–prey studies, should be used sparingly because of the numerous untested assumptions involved. Who should purchase this volume? Certainly all academic and wildlife agency libraries in the geographic range of the gray wolf should have this monograph on their shelves, as well as wolf specialists and all those interested in behavior, conservation, and management of wild Canis. This is the scientific magnum opus of long-time field researchers who have decades of experience with wolf research and have not shied away from some of the most important conservation battles involving wolves in Canada. Douglas Pimlott would be pleased.

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