The finding of several wolf-kills with poor fat stores could indicate that primary or secondary malnutrition was a factor in the animals"
deaths. However, it would take a statistical comparison between the fat stores of the deer at large and those of the wolf-kills to establish this.
The discovery that 13 percent of the fawns and 84 percent of the yearlings killed during January, February, and March had not yet shed their deciduous incisors and premolars, respectively, also fits well with the rest of our information. Evidently some unusual factor had caused the delay in tooth development and replacement. One possibility is that the animals were born in August or September, much later than normal. Although most deer in Minnesota are born in May and June, there are records of births in July and August. In addition, a fetus 181 to 200 days old was found in a doe killed on September 26 (Erickson _et al._ 1961).
An alternate explanation for the delay in tooth replacement is that the animals were suffering from malnutrition or nutrient deficiency. Severinghaus[35] has evidence that yearling bucks that have not replaced their deciduous premolars during November, and thus are aged at 17 months (Severinghaus 1949), generally have shorter, narrower antlers and fewer points than 18-and 19-month-old individuals. Degree of antler development in turn is considered related to nutritional state (Latham 1950). Thus it is reasonable to conclude that animals behind in tooth development and replacement, whether this is caused by age or diet, are physiologically inferior.
Most of the abnormal conditions discussed above pertain to the skeletal parts of wolf-kills. If the soft parts of a large number of kills could be examined thoroughly, one might discover a much higher incidence of diseases and other pathological conditions.
In conclusion, our data on both age and condition of wolf-killed deer show that at least during winter, wolves in our study area usually do not kill just any deer they discover, although they do try to.
Evidently, most deer can usually escape wolf predation. The most frequent exceptions are those 5-1/2 years old and older, those born late, those suffering from poor nutrition, those with abnormalities or pathological conditions, and possibly fawns.
The above conclusions parallel those of Murie (1944), Crisler (1956), Mech (1966a), and Pimlott _et al._ (1969) for wolves preying on Dall sheep, caribou, moose, and deer respectively, and further substantiate the claim by Mech (1970) that they can be extended to wolves preying on most, if not all, species of large mammals under most conditions. It is also apparent from the data presented above that deer over 5 years of age and those with abnormalities of the jaw or lower limbs represent such a small percentage of the total population that they are seldom taken by human hunters. In this respect, compet.i.tion between timber wolves and human hunters appears to be minimal in the study area.
FOOTNOTES:
[35] _C. W. Severinghaus. Unpublished data_.
[Ill.u.s.tration: _Figure 14.--Arthritis in right hind foot of specimen M-28. (Photo courtesy of University of Minnesota Veterinary Diagnostic Laboratory.)_]
[Ill.u.s.tration: _Figure 15.--Infection and fibrous ma.s.s in a front foot of specimen M-29. (Photo courtesy of University of Minnesota Veterinary Diagnostic Laboratory.)_]
[Ill.u.s.tration: _Figure 16.--Injury to left front foot of specimen M-196. (Photo courtesy of L. D. Mech)._]
[Ill.u.s.tration: _Figure 17.--Healed fracture of left hind leg of specimen M-227. (Photo courtesy of University of Minnesota Veterinary Diagnostic Laboratory.)_]
SUMMARY
White-tailed deer (_Odocoileus virginia.n.u.s_) killed by wolves (_Canis lupus_) during winter in a relatively unhunted wilderness area and in an immediately adjacent hunted area of Minnesota were compared with deer killed by hunters in the same general area, and with a hypothetical population. Deer killed by wolves were significantly older. Statistical comparisons also showed the following: (1) hunters generally killed an even s.e.x ratio of fawns, and a disproportionate number of adult bucks, (2) wolves took a higher percentage of female fawns than female adults, a disproportionate number of bucks in the wilderness area, and a higher percentage of does in the hunted area.
The latter fact evidently reflects the higher hunter success on males in the hunted area. Significantly higher incidences of abnormalities and pathological conditions of both mandibles and lower limbs were found in wolf-killed deer than in hunter-killed deer, and these conditions are described. It is concluded that wolf predation on white-tailed deer in the study area during winter generally is selective in that it tends to remove members of the prey population that are old, debilitated, or abnormal. Apparently these cla.s.ses of deer represent such a small percentage of the population that they are seldom taken by human hunters.
ACKNOWLEDGMENTS
This study was supported by Macalester College, the New York Zoological Society, the Minnesota Department of Conservation, the USDA Forest Service, and the U.S. Bureau of Sport Fisheries and Wildlife. Pilots Robert Hodge, Pat Magie, John Winship, Jack Burgess, Don Murray, and Walt Neumann aided substantially in obtaining jaws from wolf-killed deer. Students from the Macalester College Biology Department and personnel of the USDA Forest Service and the Minnesota Department of Conservation helped secure mandibles from both wolf-killed and hunter-killed deer. The interest of Mr. John E. Peninger and of many deer hunters in contributing the jaws is also greatly acknowledged.
Mr. David W. Kuehn sectioned the incisors of the deer jaws and determined their ages. Dr. Donald M. Barnes of the University of Minnesota Veterinary Diagnostic Laboratory examined the abnormal lower limbs, described their pathology, and provided photos of specimens used herein.
Mr. Wallace C. Dayton and Miss Elizabeth Dayton and the Quetico-Superior Foundation, all of Minneapolis, financed Mech during the preparation of this paper.
The following individuals read the ma.n.u.script and offered many helpful suggestions: Mr. R. L. Downing, Mr. C. W. Severinghaus, Mr. J. M. Peek, Dr. C. T. Cushwa, Mr. M. H. Stenlund, and Dr. R. R. Ream.
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