Part 7

However, it appears that the higher harvest of bucks by human hunters does markedly affect the sex ratio of the deer population in the hunted area, for the wolf-kill of adults in that area contained a significantly higher percentage of does (56 percent) than did the wolf-kill of adults in the wilderness area (29 percent).

Evidently the hunter harvest is not heavy enough to affect the age structure of the deer population to any marked degree, for no significant difference in age structure was found between the wolf-kill in the hunted area and that in the wilderness area (table 1). This does not conflict with the conclusion that hunting affects the sex ratio of the deer herd, because it would take much less to influence a population characteristic having two classes (sex) than one having 14 (age).

One additional difference in the sex ratio was found between two other subsamples of the wolf-kill--that is, the wolf-kill before and after an unusually high snow accumulation, which reached its peak about February 1, 1969 (table 1). Of a total of 77 animals killed before this snow condition occurred (including those from previous years), 38 percent were females. Of 44 animals killed after the heavy accumulation, 57 percent were females. One possible explanation for this is that females may normally be less vulnerable to wolf predation, for Kelsall (1969) has shown that they probably have a lighter weight-load-on-track than males. Thus when snow conditions changed greatly, making deer generally much more vulnerable to wolves (see Mech _et al._, p. 35), a preponderance of does suddenly might have become available. There is some evidence that does may be generally less vulnerable under most conditions, for all seven of our wolf-killed deer over 10 years old were females, and the oldest was over 14.

Condition of Wolf-Killed Deer

Because the data show that wolves in our study area tend to kill a disproportionate number of older deer, it is not surprising to discover that wolves also tend to capture a disproportionate number of individuals with abnormalities and pathological conditions (table 8). The explanation for such selection is obvious in regard to the abnormalities of the lower limbs (figs. 14-17): deer with injured or abnormal limbs simply cannot run as fast or as agilely as normal animals (table 7). Our observations show that deer usually depend on their alertness and speed to escape approaching wolves (Mech 1966b, Mech _et al._, p. 1). Any trait or condition that tended to interfere with either alertness or speed would decrease an individual's chance of escape.

It is more difficult to explain how dental abnormalities or pathological conditions of the mandible (figs. 8-10) would predispose an individual to wolf predation. However, in the case of dental abnormalities the genetic or environmental conditions that caused the abnormality might also have caused some other trait that increased the animal's vulnerability. Or the abnormal condition itself may have caused a further, more critical, disruption of the animal's physiology or behavior, which in turn predisposed it to wolf predation.

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, competition between timber wolves and human hunters appears to be minimal in the study area.

FOOTNOTES:

[35] _C. W. Severinghaus. Unpublished data_.

SUMMARY

White-tailed deer (_Odocoileus virginianus_) 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 sex 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 classes 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 manuscript 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.

LITERATURE CITED

Crisler, Lois. 1956. Observations of wolves hunting caribou. J. Mammal. 37: 337-346.

Downie, N. M., and Heath, R. W. 1959. Basic statistical methods. 289 p. New York: Harper and Bros.

Erickson, A. B., Gunvalson, V. E., Stenlund, M. H., Burcalow, D. W., and Blankenship, L. H. 1961. The white-tailed deer of Minnesota. Minn. Dep. Conserv. Tech. Bull. 5, 64 p.

Gilbert, F. F. 1966. Aging white-tailed deer by annuli in the cementum of the first incisor. J. Wildl. Manage. 30: 200-202.

Kelsall, J. P. 1969. Structural adaptations of moose and deer for snow. J. Mammal. 50: 302-310.

Klein, D. R., and Olson, S. T. 1960. Natural mortality patterns of deer in southeast Alaska. J. Wildl. Manage. 24: 80-88.

Kuehn, D. W. 1970. An evaluation of the wear method as a criterion for aging white-tailed deer. M.S. Thesis., Univ. Minn.

Latham, R. M. 1950. Pennsylvania's deer problem. Penn. Game News, Spec. Issue 1. (Cited from: Allen, D. L. 1962. Our Wildlife Legacy.)

Maguire, H. F., and Severinghaus, C. W. 1954. Wariness as an influence on age composition of white-tailed deer killed by hunters. N. Y. Fish and Game J. 1: 98-109.

Mech, L. D. 1966a. The wolves of Isle Royale. U.S. Nat. Park Serv. Fauna Ser. 7, 210 p.

Mech, L. D. 1966b. Hunting behavior of timber wolves in Minnesota. J. Mammal. 47: 347-348.

Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. 389 p. New York: Natural History Press, Doubleday.

Mech, L. D., Frenzel, L. D., Jr., Karns, P. D., and Kuehn, D. W. 1970. Mandibular dental anomalies in white-tailed deer from Minnesota. J. Mammal. 51: 804-806.

Murie, A. 1944. The wolves of Mount McKinley. U.S. Nat. Park Serv. Fauna Ser. 5, 238 p.

Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The ecology of the timber wolf in Algonquin Provincial Park. Ont. Dep. Lands and Forests Res. Rep. (Wildl.) 87, 92 p.

Ryel, L. A., Fay, L. D., and Van Etten, R. C. 1961. Validity of age determination in Michigan deer. Mich. Acad. Sci., Art, and Letters 46: 289-316.

Severinghaus, C. W. 1949. Tooth development and wear as criteria of age in white-tailed deer. J. Wildl. Manage. 13: 195-216.

Severinghaus, C. W. 1955. P. R. Rep. W-28-R-9: Job 1A, April 13, 1955.

Shaw, S. P. 1951. The effect of insufficient harvests on an island deer herd. N.E. Wildl. Conf. (Mimeo).

Siegel, S. 1956. Non-parametric statistics for the behavioral sciences. 312 p. New York: McGraw-Hill.

Stenlund, M. H. 1955. A field study of the timber wolf (_Canis lupus_) on the Superior National Forest, Minnesota. Minn. Dep. Conserv. Tech. Bull. 4, 55 p.

THE EFFECT OF SNOW CONDITIONS ON THE VULNERABILITY OF WHITE-TAILED DEER TO WOLF PREDATION

L. David Mech, L. D. Frenzel, Jr., and P. D. Karns

Wolves (_Canis lupus_) and deer (_Odocoileus virginianus_) having evolved together, no doubt have become adapted to contending with each other's physical abilities. Thus it is not surprising to learn that deer which succumb to wolf predation are generally weaker, older, or abnormal compared with the total deer population (Pimlott _et al._ 1969, also see Mech and Frenzel, p. 35).

However, the structural and behavioral adaptations of both species must have evolved under environmental conditions that are average or usual; otherwise, an adjustment of wolf to deer populations, and vice versa, could not have been maintained over long periods. This implies that extreme or unusual conditions might sometimes occur, to which either the wolf or the deer is poorly adapted.

One of the most important environmental factors that can influence the interactions of wolves and deer is snow. The total fall, depth on the ground, and the density are all aspects of snow that may vary considerably and affect the ability of wolves to capture deer. Recent studies of wolves and deer in northeastern Minnesota (see Mech _et al._, p. 1, also Mech and Frenzel, p. 35) afforded us opportunities to investigate the relationships between snow and the interactions of wolves and deer.

METHODS

Two principal methods of study were used in this investigation. The first involved recording the snow depth and support quality ("penetrability") in feet and tenths of feet (Verme 1968). Snow measurements were taken during the winters of 1966-67, 1967-68, and 1968-69, in which large differences in snow conditions existed. Ten such measurements were made weekly near Isabella, Minnesota, in an open aspen (_Populus tremuloides_) stand away from influences that might have caused drifting or other unusual snow conditions; the measurements were averaged. Penetrability was determined with Verme's snow-compaction gauge--a 3-foot piece of 1-1/8-inch (outside diameter) copper tube filled with lead to total 3 pounds, which gives a weight per area of 211 gm./cm.^2. To obtain a measurement, the pipe is held vertically with its lower end just flush with the snow, and then is released. The depth to which it sinks is considered the penetrability of the snowpack by a walking deer.

Although the snow conditions measured at Isabella are not representative of the entire study area, year-to-year comparison in the Isabella area should also apply generally throughout the region.

The second technique used in this study was observing the movements of wolves and deer. This was usually done from low-flying aircraft, and was facilitated by the use of radiotracking, as described by Mech _et al._ (p. 1). Close inspection of wolf-killed deer was made from the ground (Mech and Frenzel, p. 35).

RESULTS AND OBSERVATIONS

Snow measurements for each winter are shown in figures 1 through 3. The winter of 1968-69 was the most extreme of the three in terms of accumulated snow, and was generally regarded as having one of the heaviest snowfalls and accumulations on record for the study area. Snow depth on the level near Isabella reached 3.9 feet at one time, and from January 3 to April 4 it exceeded 2.4 feet. The highest snow level reached during 1966-67 was 2.4 feet, and the highest level reached during 1967-68 was 1.4 feet. In the vicinity of Ely, some 30 miles from Isabella, the 1968-69 peak accumulation was 39 inches, the highest accumulation since 1948-49 when records were first kept.[36] Thus we consider the winters of 1966-67 and 1967-68 to be within the normal range for the study area, and the 1968-69 winter as being most unusual (fig. 4).

The snow penetrability in 1966-67 remained high throughout January, February, and March. During the following winter, penetrability fluctuated more, but even at its greatest, it was relatively unimportant to deer because the total snow depth was so low. During 1968-69, however, penetrability was a very important aspect of snow condition. It was so high during late January and early February, when snow accumulation was also at its peak, that a walking deer would be expected to sink in 2.5 to 3.5 feet. Snow penetrability then decreased through February and March to a point where a walking deer would sink in approximately 0.6 foot on March 21. However, because snow accumulation remained so high through February and March, the lower penetrability during late February and March still afforded no relief to running deer, because they must exert forces several times as great as when walking. On the contrary, the low penetrability (which is an indirect measure of density) could be expected to hinder a running deer in deep snow, for it would cause much more resistance.

Deer movements, like snow conditions, varied greatly during the three winters of the study. During the first two winters, deer were generally found singly and in groups of two to six, often around the shores of lakes but also scattered about inland. In late January and February 1967, running deer were observed sinking deeply into snow, but their movements still did not seem to be hindered, no doubt because of the high penetrability (low density) of the snow that year (fig. 1).

However, during late January, February, and March of 1969 the deer were much more concentrated, mostly in conifer swamps, along southwest-facing slopes, or on lakes. Although groups of two or three animals could be found in scattered inland "pockets" throughout the winter, groups of five or six were not uncommon on lakes during January. The tendency to concentrate continued to increase, and on February 6, as many as 11 deer were observed on one lake; by March 13, group size had increased to as high as 22 deer in the same area. Throughout February and March, heavy concentrations of deer tracks covered most wilderness lakes, further evidencing much greater use of shorelines than had occurred in the two previous winters (fig. 5).

No doubt deer tended to concentrate on lakes because travel inland became so difficult. On January 28, two deer were seen plowing through snow up to their necks. Although the snow began settling in February, and the penetrability decreased, by late February running deer still plunged chest-deep and had to hesitate at every bound. These conditions persisted until about March 26, by which time a surface crust strong enough to hold a running deer had formed.

In considering wolf mobility in snow, two types of movement must be recognized: the trot used during general travel, and the bounding used while chasing prey. The trot is an easy gait of about 5 m.p.h. on firm footing (Mech 1970), and can be continued for hours at a time. During periods of deep snow and high penetrability, most wolf travel is on frozen waterways, roads, snowmobile trails, and animal trails, including the wolves' own pathways, which become well packed with frequent use (fig. 6, 7A, B). Such travel was observed during each of the three winters of this study.

The second type of wolf movement affected by snow is the leaping and bounding associated with chasing prey. The shallower angle of the wolf's bound (fig. 8) (compared with that of the deer) often causes the wolf to flounder in snow that presents little hinderance to deer (Mech 1970). Such was the case in January and February 1967 in our study area. During 1967-68 no observations of wolves chasing deer were made by the authors, but reports by other field workers indicated that running conditions were similar to those of 1967.

During the winter of 1968-69, wolves also bogged down a great deal in snow when chasing deer. However, after January 1969 the snow was so deep that deer were floundering even more than wolves in many cases. The fact that wolves could run in the trail broken by deer probably also gave the wolves an advantage under the conditions that severely restricted deer movements.

The above observations of snow conditions, deer movements, and wolf movements during the three winters of the study are in accord with observations made on the differences in the ability of the wolves to capture deer during the same period. Two indices support the conclusion that wolves had a much easier time catching deer during February and March 1969 than earlier in the winter and in the two previous winters: (1) the degree of utilization of wolf-killed deer, and (2) the kill rate of radiotagged wolves.

During the winters of 1966-67 and 1967-68, and in December and early January 1968-69, most wolf-killed deer found had been thoroughly eaten, and the bones--if present at all--were well chewed and scattered at each kill (fig. 9). All skin and flesh from the skull were eaten, and the mandible was usually separated from the skull. During late February and early March 1967, few fresh kills were even found, and wolves were returning several times to old kills that had been cleaned up many days before.

However, in late January 1969 a substantial change began taking place. The skeletons of most kills found were almost intact, the flesh having been eaten from around the bones (fig. 10). Appreciably more skin was usually left on the carcass, especially on the side lying on the snow, and the neck and head were generally intact. This was true even of fawns, which in the past often were almost completely consumed.

In several cases, only about half of the flesh had been eaten from the carcasses. On February 2, 1969, four deer recently killed by wolves were found along a 1-1/2-mile stretch of Birch Lake and nearby Polaris Lake (Minnesota-Ontario border). One large doe was completely uneaten and remained so for at least 24 hours after discovery from the air. Further, one fawn had only a few pounds of flesh eaten, a yearling doe was half eaten, and another fawn was about 75 percent eaten. Hazardous landing conditions during this period severely limited the number of carcasses that could be examined from the ground, but on February 6 a yearling doe was discovered that had only about 5 to 10 pounds of flesh eaten, and on February 8 an adult doe was found that was completely intact except for wounds.