FOOTNOTES:

"Feb. 7, 1969—about 11:30 a.m.—aerial and visual—1059 and 2 other wolves traveling overland about halfway between Heart L. and August L. (R10W-T61 N.Sect. 17 center). They were traveling quickly and intently along a fresh wolf trail, with a lighter reddish individual in the lead. The other 2 animals were darker colored, and one of them was smaller than the other. One of them must have been 1059."We soon found that about half a mile ahead of the pack was a dark wolf hurrying away from the three. This animal often looked back and ran whenever it encountered good running conditions. It soon became obvious that the pack of 3 was chasing this individual. Because it [the lone wolf] often broke its own trail, the pack gradually gained on this animal. The single wolf flushed a deer which ran when the wolf was about 75 feet away and floundered in the snow, but the wolf continued hurrying on by."Although the deer ran only about 50 yards and stopped, the pack of 3 also hurried on by. The single wolf flushed another deer, ignored it, and continued by, as did the pack of 3. The chase continued for 2 miles as we watched, into the N.E. corner of Sect. 18 and then into the N. Central part of Sect. 8, and the pack got to within 150 yards of the single wolf."However, at this point, the 2 darker members of the pack had fallen about 100 yards behind the lead one. The lead animal stopped and waited for them, as it had done a few times before. It then turned around and headed back to these animals. When they met, the reddish animal's tail was held vertically and there was much tail wagging by all for about 1 minute. Then all animals lay down for a minute and then went up on a knoll. There was much activity and 'playing' on the knoll. (12:10 p.m.)"The single wolf continued running and looking back for at least another mile. We left at 12:21 p.m."At 4:07 p.m. we saw a single wolf running across a small lake and looking behind it about 8 miles N.W. of these animals. The creature behaved the same as the one being chased today, and we wondered whether it could be the same animal."

"We soon found that about half a mile ahead of the pack was a dark wolf hurrying away from the three. This animal often looked back and ran whenever it encountered good running conditions. It soon became obvious that the pack of 3 was chasing this individual. Because it [the lone wolf] often broke its own trail, the pack gradually gained on this animal. The single wolf flushed a deer which ran when the wolf was about 75 feet away and floundered in the snow, but the wolf continued hurrying on by.

"Although the deer ran only about 50 yards and stopped, the pack of 3 also hurried on by. The single wolf flushed another deer, ignored it, and continued by, as did the pack of 3. The chase continued for 2 miles as we watched, into the N.E. corner of Sect. 18 and then into the N. Central part of Sect. 8, and the pack got to within 150 yards of the single wolf.

"However, at this point, the 2 darker members of the pack had fallen about 100 yards behind the lead one. The lead animal stopped and waited for them, as it had done a few times before. It then turned around and headed back to these animals. When they met, the reddish animal's tail was held vertically and there was much tail wagging by all for about 1 minute. Then all animals lay down for a minute and then went up on a knoll. There was much activity and 'playing' on the knoll. (12:10 p.m.)

"The single wolf continued running and looking back for at least another mile. We left at 12:21 p.m.

"At 4:07 p.m. we saw a single wolf running across a small lake and looking behind it about 8 miles N.W. of these animals. The creature behaved the same as the one being chased today, and we wondered whether it could be the same animal."

On February 18, 1969, Ream made a similar observation, as follows (quoted from his field notes):

"Got visual sighting on 1059 with 3 other wolves at 11:55 about a mile west of Omaday Lake and they were running along fairly fast on a trail. When we circled a second time we saw 2 wolves curled up sleeping on a knoll ahead (south) of the running pack. We then realized the running wolves were on the trail of the sleeping wolves and when the pack of 4 with "red" in the lead was about 50 yards from the knoll the 2 sleeping wolves jumped up and charged away in the opposite direction full tilt, and split and went in 2 directions. When the pack reached the knoll they started off on the trail of the wolf that headed N.E. and then changed and went after the one that headed S.W. The reddish wolf was in the lead and really picked up the pace. Although the reddish wolf seemed to gain on the chased one 3 or 4 times, the pack as a whole couldn't catch up, even though the single was breaking trail. The reddish wolf, after gaining, always stopped and waited for the others or went back to find them. They chased this wolf for 2½ to 3 miles, all the way down to Highway 1 at a point 3.0 miles from the lab [Kawishiwi Field Station, U.S. Forest Service]. There was a dense patch, 10-15 acres, of woods just before Highway 1 and we lost sight of the chased wolf for a while and also the 4 when they entered it, but shortly we found that the chased one had somehow doubled back and was heading N.E. again. The pack was apparently confused for at one point 3 of them were wandering back and forth on Highway 1, apparently looking for the trail of the chased wolf. Two of these paralleled the Highway for a couple hundred yards and then stopped on top of a hill, apparently resting. During this chase both the single wolf and the pack chased up deer from their route of travel and didn't seem topay much attention to them, even though some were really floundering in the deep snow. We finally stopped watching all of this at 1:30 p.m. and proceeded on our rounds."

On February 21 we also saw a single wolf running and looking behind several times on Ojibway Lake. Even when it saw a fisherman on the lake within 1/8 mile, it continued across to the opposite shore seeming most intent on avoiding whatever was on its trail. Presumably it had also been chased by a pack.

The cases of tolerance or indifference that we witnessed between wolves involved our lone animals. On January 27, 1051 was at a kill he had made the day before, and another wolf was sitting within 100 feet looking toward the carcass. Eventually the unidentified wolf left without approaching any closer. A lone wolf was also seen near 1053 in the general vicinity of a moose carcass, which probably both were feeding on at different times. Three such observations were made, on February 10, 15, and 18; and on February 21 another wolf was also seen near 1053 some 2.5 miles away from the moose carcass. In all cases, the two animals were 80 to 200 yards apart in open country and must have been aware of each other's presence.

The primary prey of most wolves in our study area is the white-tailed deer (fig. 29), but some moose (fig. 30) are also killed. We have examined the remains of six moose that were eaten by wolves, two of which were killed by them (fig. 31). One was found on February 25, 1967, on Gillis Lake and the other on March 7, 1969, on Twinkle Lake. These locations are within 3 miles of each other, suggesting that a wolf pack in that area may be more accustomed to preying on moose than other packs. The other four moose carcasses were found in other parts of the study area, but circumstances were such that the causes of death of those animals could not be determined. A discussion of the details of wolf-moose relations in our study area must await the collection of additional data.

Figure 29.—The main prey of wolves in northern Minnesota is the white-tailed deer. (Photo courtesy of L. D. Mech.)Figure 29.—The main prey of wolves in northern Minnesota is the white-tailed deer. (Photo courtesy of L. D. Mech.)

Figure 30.—Moose are also killed by wolves. (Photo courtesy of Allan Taylor.)Figure 30.—Moose are also killed by wolves. (Photo courtesy of Allan Taylor.)

The remains of 93 wolf-killed deer, and 49 probable wolf-kills, were examined for age, sex, and condition and were compared with a sample of 433 hunter-killed deer from the same general area. The wolf-killed deer were generally much older than the hunter-kills and had a significantly higher percentage of jaw and limb abnormalities (see Mech and Frenzel,page 35).

Figure 31.—Only a few wolf-killed moose were located during the study. (Photo courtesy of Laurence Pringle.)Figure 31.—Only a few wolf-killed moose were located during the study. (Photo courtesy of Laurence Pringle.)

Until recently the only observations of wolves hunting deer were those reported by Stenlund (1955) for northern Minnesota. He described two reports of actual observations and two reports of interpretations of tracks in the snow, all successful hunts. Since that time several descriptions of successful and unsuccessful hunts have also been published (Mech 1966b, Rutter and Pimlott 1968, Pimlottet al.1969, Mech 1970). Nevertheless, many more observations must be made before generalizations can be formed.

During the present study we were able to witness a number of hunts from the air and piece together others based on tracks. The following descriptions are quoted from the field notes of Mech:

"26 January 1967. About 3/4 mile N.E. of Alice Lake."Jack Burgess [pilot] and I were following a pack of 8 wolves, when at 4:15 they veered from their former line of travel, about 30°. They were then about 200 yards from 2 deer. They began wagging their tails when about 175 yards from the deer. One deer, on the edge of a steep bank, was lying, but one was standing about 75 yards N. of it in open hardwoods. The wolves continued toward the latter deer."This deer remained standing in the same place until the wolves approached to within about 100 feet of it. The lead wolf stopped, when that distance from the deer, and the others caught up but also stopped when within about 25 feet behind the lead wolf. By this time the deer, whose body was facing away from the wolves, had its head turned back over its shoulder toward the wolves. The wolves and the deer remained absolutely still while staring at each other, 100 feet apart, for 1-2 minutes, while we made several circles."Suddenly the deer bolted, and instantly the wolves pursued. I am fairly certain that it was the deer that bolted first, but could be mistaken. The action was almost simultaneous. The deer headed toward the other deer near the top of the high bank. This animal had been lying but had arisen when the wolves were about 150 yards away."The lead wolf followed in the deer's trail, but the others cut toward the bank. This flushed the second deer (near the edge of the bank), which ran down the bank. Meanwhile when the first deer reached the edge of the bank, it headed due W. along the top of it. Only the lead wolf pursued this animal. The other deer had headed down the bank to the S.E., and at least a few of the wolves followed it."We could not watch both deer, so we continued following the first. The deer had no trouble in snowdrifts, but the wolf was hindered by them. The wolf followed the deer for about 200 yards along the top of the bank, and then gave up after losing ground. The wolf had run a total distance of about 250-275 yards. He then lay down and rested."We noticed at least 3 wolves stopped part way down the bank in the trail of the second deer. However, we did notsee the remaining wolves or the second deer."Eventually (after about 5 minutes), these wolves joined the first, and all rested. At 4:25 p.m., one wolf started toward a third deer, which had been lying under a tree while the former chase took place. The deer was about 150 yards from where the wolves rested, and it had stood before the wolf started toward it. We could not see whether the deer or wolf bolted first, but suddenly both animals were bounding away. The wolf chased the deer about 125 yards and gave up after losing ground. The other wolves followed slowly in its trail, and all assembled and rested. The deer continued running for at least 1/4-mile.""27 February 1969. 2 miles N. of August Lake."1059's pack of 5 was heading N.E. at 4:10 p.m. when they got to within 100 yards of 2 standing deer. The deer had been standing alertly in a shallow draw, and when at least 2 wolves got to within 100 yards, they fled. The wolves began running after them."The deer were in snow up to their bellies and had to hesitate slightly at each bound. But they ran fast. We could only see one wolf very much [of the time]. It was also having a difficult time in the snow, and after a total run of about 250 yards (100 to the deer's original location and 150 after the deer), the wolf lay on the snow and rested about 10 minutes. The deer ran only about 200 yards more and stood alertly for the next 20 minutes at least. The wolves then went on."27 March 1969. About 2 miles S.E. of Central Lakes, Minnesota."At 3:00 p.m. while we were following wolf 1051 by aircraft in above location, we saw a deer running very quickly on top of the crusted snow and then stand and watch its backtrail. About 1½ minutes later we saw 1051 running along the same route. We did not see when the deer fled again, but saw it running about 100 yards from the wolf and doubling back paralleling its original route. When the wolf got near the approximate doubling-back point, he lay down and rested for about 5 minutes. The deer continued fleeing for about 350 yards, stopped, and for several minutes faced its backtrail. The wolf finally continued on in his original direction, giving up the chase."At 4:30 p.m.—1½ miles S. of Central Lakes, Minnesota—Wolf 1051 had come to within 100 yards of [four-lane] Highway 53 and was hesitant to approach it. Several cars were going by in both directions. Thus the wolf headed S. parallel with the highway about 150 yards E. of it."Suddenly two deer, which we had noticed S. of the wolf earlier, fled across the highway. The wolf soon got to the point where they crossed, hesitated about a minute and then ran across. No cars came at that time."We could not always see the deer or the wolf when W. of the road because there were several patches of evergreens. The wolf did head straight W. after crossing the road. Then about 250 yards W. of this point we saw a deer come out onto an old woods road which lay in a N.W.-S.E. axis. The deer ran N.W. on the road and then we saw the wolf where the deer had come out onto the road. While the deer ran N.W., the wolf cut into the woods to his right, N.E. We could not see it then but presumed it was running N.W. paralleling the road."After the deer had run about 50 yards up the road, it also headed N.E. into the evergreens. Within a few seconds it fled right back out and started S.E. down the road. The wolf was about 50 feet behind it and began gaining."When the deer got back to where the wolf had headed into the woods from the road before, it also headed N.E. into the woods. The wolf was then about 20 feet away and the deer was headed N. around in a circle with the wolf closing in on the outside. The wolf did not emerge from the evergreens for at least 15 minutes,nor did we see the deer, so I presume the wolf killed the deer. [But see entry for April 1.]"1 April 1969. Dan Frenzel and I searched the area described on March 27 for 1 hour and found no sign of a kill. Old wolf tracks were seen, but only a single wandering track. No concentration such as usually seen at kills. Best conclusion is that 1051 didnotkill the deer where seen from the air March 27."

"26 January 1967. About 3/4 mile N.E. of Alice Lake.

"Jack Burgess [pilot] and I were following a pack of 8 wolves, when at 4:15 they veered from their former line of travel, about 30°. They were then about 200 yards from 2 deer. They began wagging their tails when about 175 yards from the deer. One deer, on the edge of a steep bank, was lying, but one was standing about 75 yards N. of it in open hardwoods. The wolves continued toward the latter deer.

"This deer remained standing in the same place until the wolves approached to within about 100 feet of it. The lead wolf stopped, when that distance from the deer, and the others caught up but also stopped when within about 25 feet behind the lead wolf. By this time the deer, whose body was facing away from the wolves, had its head turned back over its shoulder toward the wolves. The wolves and the deer remained absolutely still while staring at each other, 100 feet apart, for 1-2 minutes, while we made several circles.

"Suddenly the deer bolted, and instantly the wolves pursued. I am fairly certain that it was the deer that bolted first, but could be mistaken. The action was almost simultaneous. The deer headed toward the other deer near the top of the high bank. This animal had been lying but had arisen when the wolves were about 150 yards away.

"The lead wolf followed in the deer's trail, but the others cut toward the bank. This flushed the second deer (near the edge of the bank), which ran down the bank. Meanwhile when the first deer reached the edge of the bank, it headed due W. along the top of it. Only the lead wolf pursued this animal. The other deer had headed down the bank to the S.E., and at least a few of the wolves followed it.

"We could not watch both deer, so we continued following the first. The deer had no trouble in snowdrifts, but the wolf was hindered by them. The wolf followed the deer for about 200 yards along the top of the bank, and then gave up after losing ground. The wolf had run a total distance of about 250-275 yards. He then lay down and rested.

"We noticed at least 3 wolves stopped part way down the bank in the trail of the second deer. However, we did notsee the remaining wolves or the second deer.

"Eventually (after about 5 minutes), these wolves joined the first, and all rested. At 4:25 p.m., one wolf started toward a third deer, which had been lying under a tree while the former chase took place. The deer was about 150 yards from where the wolves rested, and it had stood before the wolf started toward it. We could not see whether the deer or wolf bolted first, but suddenly both animals were bounding away. The wolf chased the deer about 125 yards and gave up after losing ground. The other wolves followed slowly in its trail, and all assembled and rested. The deer continued running for at least 1/4-mile."

"27 February 1969. 2 miles N. of August Lake.

"1059's pack of 5 was heading N.E. at 4:10 p.m. when they got to within 100 yards of 2 standing deer. The deer had been standing alertly in a shallow draw, and when at least 2 wolves got to within 100 yards, they fled. The wolves began running after them.

"The deer were in snow up to their bellies and had to hesitate slightly at each bound. But they ran fast. We could only see one wolf very much [of the time]. It was also having a difficult time in the snow, and after a total run of about 250 yards (100 to the deer's original location and 150 after the deer), the wolf lay on the snow and rested about 10 minutes. The deer ran only about 200 yards more and stood alertly for the next 20 minutes at least. The wolves then went on.

"27 March 1969. About 2 miles S.E. of Central Lakes, Minnesota.

"At 3:00 p.m. while we were following wolf 1051 by aircraft in above location, we saw a deer running very quickly on top of the crusted snow and then stand and watch its backtrail. About 1½ minutes later we saw 1051 running along the same route. We did not see when the deer fled again, but saw it running about 100 yards from the wolf and doubling back paralleling its original route. When the wolf got near the approximate doubling-back point, he lay down and rested for about 5 minutes. The deer continued fleeing for about 350 yards, stopped, and for several minutes faced its backtrail. The wolf finally continued on in his original direction, giving up the chase.

"At 4:30 p.m.—1½ miles S. of Central Lakes, Minnesota—Wolf 1051 had come to within 100 yards of [four-lane] Highway 53 and was hesitant to approach it. Several cars were going by in both directions. Thus the wolf headed S. parallel with the highway about 150 yards E. of it.

"Suddenly two deer, which we had noticed S. of the wolf earlier, fled across the highway. The wolf soon got to the point where they crossed, hesitated about a minute and then ran across. No cars came at that time.

"We could not always see the deer or the wolf when W. of the road because there were several patches of evergreens. The wolf did head straight W. after crossing the road. Then about 250 yards W. of this point we saw a deer come out onto an old woods road which lay in a N.W.-S.E. axis. The deer ran N.W. on the road and then we saw the wolf where the deer had come out onto the road. While the deer ran N.W., the wolf cut into the woods to his right, N.E. We could not see it then but presumed it was running N.W. paralleling the road.

"After the deer had run about 50 yards up the road, it also headed N.E. into the evergreens. Within a few seconds it fled right back out and started S.E. down the road. The wolf was about 50 feet behind it and began gaining.

"When the deer got back to where the wolf had headed into the woods from the road before, it also headed N.E. into the woods. The wolf was then about 20 feet away and the deer was headed N. around in a circle with the wolf closing in on the outside. The wolf did not emerge from the evergreens for at least 15 minutes,nor did we see the deer, so I presume the wolf killed the deer. [But see entry for April 1.]

"1 April 1969. Dan Frenzel and I searched the area described on March 27 for 1 hour and found no sign of a kill. Old wolf tracks were seen, but only a single wandering track. No concentration such as usually seen at kills. Best conclusion is that 1051 didnotkill the deer where seen from the air March 27."

We also saw 1055 and her associate actually kill a deer, on February 6, 1969, but we did not realize what was going on and it happened so fast that we only saw a wolf rushing and biting at the front end of the downed animal. The chase had to have lasted only a few seconds.

In addition to the above direct observations, we also were able to piece together from tracks in the snow the chase and successful encounter between a single wolf and a deer in two instances. In the first case, on January 25, 1967 (11:50 a.m.), we arrived at the scene (near Grub Lake, just N. of Snowbank Lake) within an hour of the encounter, and the wolf was still feeding on the deer, which had been a 2½-year-old female. Mech examined the area from the ground and made the following observations:

"The deer had come S.W. down the middle of the lake at a fast walk, turned around, backtracked a few yards and headed to the N.W. shore of the lake. Meanwhile a wolf had come at a trot along the deer's track, but it had cut to the N.W. shore about 50 yards N.E. of where the deer had. When still on the ice about 15 feet from shore, the wolf began running as evidenced by his long bounds. He continued running inland about 50 feet from shore toward the deer. The deer had walked inland from the shore and may have stood there about 25 feet from shore. Suddenly it had bounded away. The bounding wolf track was in the same trail as the deer's for about 25 yards but then it paralleled the deer's about 5 feet away on the inland side. After about 125 yards from where the deer flushed, the deer was pulled down. It wasnoton its side but rather had sunk into the snow in more-or-less of an upright position."Apparently the deer had just about reached the shore when the wolf noticed it, and it detected the wolf. At this time the wolf must have been up the shore about 50 yards where his tracks first showed he began bounding. There was no sign that the wolf had spotted the deer on the lake and had tried to cut it off from shore by running inland along the shore and then waiting for the deer to come inland. Once the wolf had begun bounding, he continued until he pulled the deer down.... Sign showed that the deer dropped within about 20 feet of where she had begun bleeding."

"Apparently the deer had just about reached the shore when the wolf noticed it, and it detected the wolf. At this time the wolf must have been up the shore about 50 yards where his tracks first showed he began bounding. There was no sign that the wolf had spotted the deer on the lake and had tried to cut it off from shore by running inland along the shore and then waiting for the deer to come inland. Once the wolf had begun bounding, he continued until he pulled the deer down.... Sign showed that the deer dropped within about 20 feet of where she had begun bleeding."

The second case involved a 5½-year-old buck, No. M-28, which had arthritis of his right hind foot and probably had defective gait (see Mech and Frenzelp. 35). The attack took place on Basswood Lake on February 2, 1967, and excerpts from field notes by Mech follow:

"A single wolf had killed this deer after chasing, following, or tracking the deer about 3.75 miles. The deer's last 350 yards was a fast walk—the tracks were one in front of the other and about 2 feet apart, and there was no leaping or bounding. Same with the wolf—a fast trot."Where the tracks came together, the deer apparently had fallen, but there was no blood. From there, the deer dragged its feet or the wolf for about 25 feet and then went down again. The wolf circled the deer, and for the next 150 feet, the 2 animals had fought or scuffled and then the deer had gone down where we found it."The 4-mile persistence of this wolf—whether tracking, following, or chasing the deer—is remarkable [compared with most chases] and makes me believe the wolf had good reason to believe it could kill the deer."

"Where the tracks came together, the deer apparently had fallen, but there was no blood. From there, the deer dragged its feet or the wolf for about 25 feet and then went down again. The wolf circled the deer, and for the next 150 feet, the 2 animals had fought or scuffled and then the deer had gone down where we found it.

"The 4-mile persistence of this wolf—whether tracking, following, or chasing the deer—is remarkable [compared with most chases] and makes me believe the wolf had good reason to believe it could kill the deer."

Our observations of wounds on fresh kills confirm the following description by Stenlund (1955, p. 31) of the location and manner of attack of wolves on deer: "No evidence of hamstringing of deer was found on freshly killed carcasses, although the possibility does exist. Usually deer are run down from behind, thewolf or wolves biting at the hind flanks and abdomen, or at the hind flanks and head region simultaneously."

On each kill, all the flesh and much of the skin and bones were eaten, at least during the winters of 1966-67 and 1967-68. This was also true during December 1968 and much of January 1969. However, during February and March 1969 when an unusual accumulation of snow had built up, most of the kills were only partly eaten (see Mechet al.,page 51). In previous years deer freshly killed by single wolves were sometimes found with only a few pounds of flesh or viscera missing. However, in each case the carcasses were almost completely cleaned up within a few days, often by packs to which the single wolves may have belonged (Mech 1970).

Usually the first parts of a carcass to be eaten are the hams and part of the viscera from the coelomic cavity. In one case where a wolf was interrupted while feeding it was apparent that the animal had been stripping the omental fat from the carcass. This may be the wolf's favorite part of a deer, for the stomach of one wolf that we examined in January 1967 contained nothing but such fat.

The average consumption and kill rate of deer by wolves has not yet been determined, but we have some information bearing on the subject. Because our data were obtained during a winter of unusually deep snow, and it was obvious that wolves were killing more deer than they could eat at the moment (see Mechet al.,page 51), our figures should be considered much higher than average. However, they should be useful in that they probably represent the maximum kill rate not only throughout the year but also throughout a period of many years.

By observing each of our radiotagged wolves whenever possible and noting whether or not it was feeding on a kill, we learned that our wolves generally remained close to their kills for periods of from 1 to 7 days, depending on how recently they had eaten (fig. 32). Thus, when a wolf was found at a new location each day, the assumption could be made that the animal did not currently have a kill.

Figure 32.—Periods spent by radiotagged wolves and their associates feeding on kills judged to be their own. This does not include periods when they were known to be feeding on carrion.Figure 32.—Periods spent by radiotagged wolves and their associates feeding on kills judged to be their own. This does not include periods when they were known to be feeding on carrion.(Click for larger image.)

Figure 33.—Radiotagged wolf (upper left) found at kill (lower right). (Photo courtesy of L. D. Frenzel.)Figure 33.—Radiotagged wolf (upper left) found at kill (lower right). (Photo courtesy of L. D. Frenzel.)

We assumed that wolves found at fresh kills (fig. 33) had made them unless there was evidence to the contrary as with 1053, the scavenger. When a wolf was found at one location for several consecutive days but could not be observed, we assumed it was feeding on a kill, since whenever wolves were observed remaining in the same location for several days they were seen feeding. Thus a range of possible number of kills per wolf was determined, with the lower limit being the known minimum and the upper limit the possible maximum. When more than one wolf fed on a kill, as with the pack, the figures were calculated on a per-wolf basis.

In this way we obtained data on a total of 468 wolf-days and found a total kill of 35 to 48 deer (table 7). This averages out to a kill rate of one deer per 10 to 14 days per wolf. The figure varied considerably among individuals—1051 had the highest rate of one kill per 6.3 to 7.2 days, and each wolf in 1059's pack had the lowest rate (except for 1053, the scavenger) of one deer per 14.0 to 18.0 days.

It is significant that the pack of five wolves had a lower kill rate per wolf than did single wolves and pairs. This is explainable because the ability of wolves to kill deer during early 1969 was much greater than usual (see Mechet al.,p. 51). Thus single wolves probably could kill deer just as easily as could packs, but they did not need to share them. This differs markedly from the situation on Isle Royale, where lone wolves usually feed only on moose remains left by packs (Mech 1966a, Jordanet al.1967).

That lone wolves had more of a food surplus than those in the pack is confirmed by the figures on the average number of days that the various wolves fed on kills (table 7). Wolf 1051 spent an average of only 2.2 to 2.4 days feeding at each of his kills, whereas 1059's pack of five spent an average of 5.8 to 7.5 wolf-days at each kill. Further confirmation is found in the fact that even when most wolves were leaving their kills partly uneaten, a pack of 8 to 10 wolves (probably that to which 1057 belonged) was seen completely devouring a kill.

Table 7.—Kill rate of deer by radiotagged wolves and their associates

WolfnumberWolvesDatesWolf-daysof dataKillsWolf-daysper kill[14]Wolf-daysfeedingWolf-daysfeedingper killNumberNumberNumberMean NumberNumberMean Number10511Nov. 26 to Apr. 310114-166.3-7.233-402.2-2.41053[15]1Dec. 14 to Mar. 27752-325.0-37.59-184.5-6.010551-2Jan. 9 to Mar. 14614-96.7-15.013-252.8-3.310571-13Jan. 24 to Feb. 28515-77.3-10.225-334.7-5.010595Jan. 25 to Mar. 1418010-1314.0-18.0755.8-7.5Summary22Nov. 27 to Apr. 346835-48[16]9.8-13.4145-181[17]3.8-4.1Before Feb. 11427-9[18]15.7-20.339-565.1-5.6After Jan. 3132628-398.4-11.6106-1253.2-3.8

FOOTNOTES:[14]Kill rate per wolf.[15]Figures for this animal are so low because she was basically a scavenger.[16]Average kill rate per wolf for all radiotagged wolves and their associates, derived by dividing total number of wolf-days by total number of kills.[17]Average number of days that each wolf spent at each kill, derived by dividing total number of wolf-days spent feeding by the total number of kills.[18]This figure probably is the closest to the actual kill rate during most winters.

[14]Kill rate per wolf.

[15]Figures for this animal are so low because she was basically a scavenger.

[16]Average kill rate per wolf for all radiotagged wolves and their associates, derived by dividing total number of wolf-days by total number of kills.

[17]Average number of days that each wolf spent at each kill, derived by dividing total number of wolf-days spent feeding by the total number of kills.

[18]This figure probably is the closest to the actual kill rate during most winters.

Therefore it is probable that the kill rate per wolf for members of the pack of five is much closer to the usual average winter kill rate. It can still be considered higher than the usual winter rate, however, because this pack also was leaving some of its kills partly uneaten.

A reasonable approximation of the average kill rate during most winters would be the rate found for our radiotagged wolves before February 1, because the relations among the wolves, the deer, and the snow during that period were not unlike those of most winters. The average kill rate per wolf before February 1 was estimated at one deer per 15.7 to 20.3 days.

After this period, the rate increased to about one deer per 8.4 to 11.6 days, and an estimated 50 percent of the available food was left uneaten (see Mechet al.,page 51). This implies that the kill rate during February and March was about twice as high as usual. On this basis, the usual kill rate would be estimated at one deer per 16.8 to 23.2 days, which checks well with the rate found before February (one deer per 15.7 to 20.3 days). Thus we feel that an estimated kill rate of about one deer per 18 days per wolf is a close approximation of the average kill rate for most winters. This is about 50 percent less than the kill rate of one deer per 4 days estimated by Stenlund (1955) for two packs of three wolves (one deer per 12 days per wolf). However, it compares favorably with the actual kill rate of one deer per wolf per 17.6 days found for a pack of eight wolves in Ontario.[19]

Once the average rate of kill is known, the average food consumption per wolf can be calculated. The average deer (considering both fawns and adults) from the Superior National Forest during winter weighs about 113 pounds (calculated from Ericksonet al.1961), and an arbitrary 13 pounds can be deducted from this for inedible portions. This leaves 100 pounds of deer per wolf per 18 days, or 5.6 pounds per wolf per day. This figure is much less than the 10 to 14 pounds estimated consumption rate for wolves feeding on moose on Isle Royale (Mech 1966a). However, much variation can be expected in an animal whose physiology must be adapted to a feast-or-famine existence.

Wolves can be maintained in captivity on 2.5 pounds of meat per day, and large active dogs (Canis familiaris) require 3.7 pounds per day, so it is likely that the minimum daily requirement for wolves in the wild is about 4.0 pounds per day (Mech 1970). This figure agrees well with the estimated consumption rate for our study area.

FOOTNOTES:[19]Kolenosky, G. B. Wolf movements, activities and predation impact on a wintering deer population in East-Central Ontario. (Manuscript in preparation for publication.)

[19]Kolenosky, G. B. Wolf movements, activities and predation impact on a wintering deer population in East-Central Ontario. (Manuscript in preparation for publication.)

Censusing wolves in a 1.5-million-acre study area is a difficult task, and we have no direct information on which to base a population estimate. However, some deductions can be made about the relative population densities in our study area between the period 1948 to 1953 and the period of the present study, 1967 to 1969.

R. A. Rausch (1967a) hypothesized that the frequency of large packs is higher when population density is high, and presented evidence supporting this idea. On this assumption, a comparison of pack-size distributions between various periods can indicate relative population densities between periods. The advantage of this method is that it eliminates the usual type of year-to-year biases in wolf censuses such as might result from differences in precise census route, type of aircraft, skill of observers, and other conditions. Only a difference that would cause a bias in thesizeof the packs seen would be of importance.

Therefore, we tested the difference in size distributions of population units between the 1948-53 study period and the present period (table 1), using a Kolmogorov-Smirnov two-sample test (Siegel 1956). The average "pack" size in the earlier years was 2.8, compared with 4.2 at present; thus pack sizes are significantly larger at present (95 percent level). This indicates that the population density from 1967 to 1969 may have been higher than from 1948 to 1953. This apparent change may be attributable to a reduction in snaring, trapping, and aerial hunting that took place between the two periods as a result of changes in State game regulations.

A similar comparison between our observations from 1967 and those from 1968-69 (table 1) shows no significant difference between theseyears, so it appears that the density of wolves in our area has remained about the same over the period of three winters. This agrees with the results of several other studies summarized by Mech (1970) in which wolf populations unaffected by man have been found to remain relatively stable from year to year.

The movements, behavior, and ecology of the wolves in our study area during winter are variable, and are influenced considerably by snow conditions. This may explain the fact that in late February 1969 wolves 1051, 1053, and 1055 suddenly extended their travels and range (fig. F-34andtable 4).

However, increased travel may have resulted from other factors. For one thing, the wolves apparently did not need to spend so much time hunting as before. Because of the deep snow, the ability of wolves to capture deer increased, and the animals had a surplus of food. Perhaps under such conditions wolves may use more of their energy for traveling than for hunting.

Figure 34.—Net weekly (straight-line) distances traveled by three radiotagged wolves.Figure 34.—Net weekly (straight-line) distances traveled by three radiotagged wolves.

In this respect it is interesting that 1051 moved right out of his area and traveled into country that presumably was unknown to him. Wolves 1053 and 1055 each ventured into an area that was almost devoid of deer and that even had few moose in it. Without sufficient fat reserves in all these animals, it would seem disadvantageous for them to have made these travels.

Evidently wolves can obtain enough food in much smaller areas than these three animals used after February. Both 1059's pack of five and 1057 lived in relatively small areas throughout the winter and seemed to survive well. Before late February, 1051, 1053, and 1055 did also. Thus some factor other than food must have influenced the movements of these three animals from late February through April.

The fact that the increased movement began during the breeding season makes one suspect a relationship between the two. One possibility is that the factors increasing the hormonal flow associated with breeding in adults stimulate a hormone output in immature or subordinate individuals that causes an increase in their movements. An alternative is that the breeding behavior of resident packs involves the beginning of, or an increase in, aggression toward neighboring nonmembers. This might force the lone animals to shift about over large areas in avoidance of such aggression.

Whatever the cause of the changes in movements of these animals, the fact that the pack used a much smaller area than any of the lone wolves may be of central importance in trying to understand the organization of the wolf population. The following pieces of information are also pertinent to such an understanding: (1) the pack, which can be presumed to include a breeding pair (Mech 1970), chased other wolves in its area; (2) the lone wolves, which apparently did not breed, were tolerant of, or indifferent to, other lone wolves in their areas; (3) the ranges of the lone wolves overlapped considerably (fig. 35); (4) the lone wolves seemed to avoid certain large areas that one might logically think would have been visited by them (fig. 35); and (5) packs of wolves were sometimes observed in these large areas (fig. 35).

Figure 35.—Locations of all radiotagged wolves and unmarked packs observed during winter 1968-69, except dispersal of 1051 out of the study area. Only selected lakes shown.Figure 35.—Locations of all radiotagged wolves and unmarked packs observed during winter 1968-69, except dispersal of 1051 out of the study area. Only selected lakes shown.(Click for larger image.)

From the above information it can be hypothesized that the wolf population consists basically of groups of breeding packs defending territories of limited size, with lone wolves and other nonbreeding population units that are tolerant of each other shifting about in much larger nonexclusive areas among these territories. The information from Isle Royale (Mech 1966a, Jordanet al.1967) is consistent with this idea, but the area of that island (210 square miles) is too small to allow untested extrapolations to be made about spacing in much larger wolf populations. Data from Algonquin Park, Ontario (Pimlottet al.1969) also strongly suggest this hypothesis. However, the packs studied there could not be identified with certainty, and little information was obtained about nonbreeding population units.

To test the proposed hypothesis with certainty, a larger number of identifiable breeding and nonbreeding population units from the same general area must be followed during at least one winter. This will be the main objective of our next study.

During the winters of 1966-67, 1967-68, and 1968-69, aerial observations of timber wolves (Canis lupus) were made in the Superior National Forest in northeastern Minnesota, where the primary prey is white-tailed deer (Odocoileus virginianus). In 480 hours of flying during the study, 77 sightings involving 323 wolves were made. In addition, during 1968-69, five radiotagged wolves and their associates were tracked via receivers in aircraft for a total of 570 "wolf-days." Visual observations were made during 65 percent of the times the wolves were located from December through April.

The average size of each population unit (including single wolves, pairs, and packs) observed was 4.2, although packs of as many as 13 wolves were sighted. Radiotagged wolves spent most of their daylight hours resting during winter, and when traveling, hunting or feeding during the day, tended to do so before 11:00 a.m. and after 3:00 p.m.

Considerable variation was discovered in the movement patterns of individual wolves, with straight line distances between consecutive daily locations ranging from 0.0 to 12.8 miles, and between weekly locations, 0.0 to 49.0 miles. A pack of five wolves used a range about 43 square miles in extent, whereas lone wolves covered areas many times this size. One animal in an apparent dispersal was tracked a straight line distance of 129 miles between extreme points.

A reddish male wolf was the leader of the pack of five and led two observed chases after alien wolves in the pack's territory. This animal was also most active during scent marking by the pack. Lone wolves were apparently indifferent to other wolves, and thus exclusive areas, or territories, were not observed among lone wolves.

Hunts involving a total of seven deer were observed and described, and two successful attacks on deer were interpreted from tracks in the snow. Wolves generally consumed all the flesh and much of the hair and bones from kills, except during February and March 1969 when extreme snow conditions increased the vulnerability of deer to an unusual degree. At that time kills were found that were partly or totally uneaten. The kill rate by radiotagged wolves and associates during the winter of 1968-69, based on 468 wolf-days of data, varied from one deer per 6.3 days to one per 37.5 days per wolf, with the average being one deer per 10 to 13 days. The rate was much lower per wolf for members of the pack of five than for lone wolves, and much lower before February 1, 1969, than after. The average rate of kill during more usual winters was estimated to be about one deer per 18 days. This is a consumption rate of about 5.6 pounds of deer per wolf per day.

Indirect evidence based on comparisons of pack-size distributions for different periods indicates that the wolf density in the study area may have increased since 1953, but that it has remained the same from 1967 to 1969.

On the basis of data presented in this paper, the following hypothesis about the organization of the wolf population studied is proposed: The wolf population consists basically of groups of breeding packs defending territories of limited size, with lone wolves and other nonbreeding population units, tolerant of each other, shifting about in much larger nonexclusive areas among these territories.

This study was supported by Macalester College, the U.S. Bureau of Sport Fisheries and Wildlife, the USDA Forest Service, the Minnesota Department of Conservation, and the New York Zoological Society. Special thanks are due the following for their help and cooperation withthis project: Mr. J. O. Wernham, former Supervisor, Mr. L. T. Magnus, Wildlife Biologist, numerous District Rangers, and other supporting personnel of the Superior National Forest, Mr. J. T. Morgan, North Central Forest Experiment Station; and Mr. S. E. Jorgensen and Mr. C. E. Faulkner, U.S. Bureau of Sport Fisheries and Wildlife.

Drs. C. T. Cushwa, L. F. Ohmann, Catherine Ream, and D. G. Schneider aided in the field work. Mr. W. W. Cochran provided advice and suggestions on the radiotracking technique, Dr. U. S. Seal furnished the drugs and the advice on their use with wolves, and Mr. R. Himes contributed significantly in the wolf trapping. Mr. L. Ringham, Ontario Department of Lands and Forests, granted permission for research personnel to radiotrack wolves crossing into Quetico Park, Canada. Numerous students from Macalester College also contributed to the field effort.

Thanks are also due pilots Robert Hodge, Pat Magie, Ken Bellos, Don Murray, Jack Burgess, and several others, who along with pilot-biologist John Winship, expertly flew the aircraft used in the study.

This report was reviewed by the following biologists: Mr. G. B. Kolenosky, Dr. P. A. Jordan, Mr. M. H. Stenlund, and Dr. D. L. Allen.

Mr. Wallace C. Dayton, Miss Elizabeth Dayton, and the Quetico-Superior Foundation, all of Minneapolis, generously contributed funds to support Mech during the preparation of the paper.

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

The selective effect of predation on prey populations is of significance in studies of evolution and population dynamics. Selective predation can be an important agent in the process of natural selection, and it influences the extent to which predators limit the numbers of their prey.

One of the predators most commonly chosen for investigating the selective effect upon prey is the wolf (Canis lupus). Because animals preyed upon by wolves generally are large, their remains can be more easily located and examined. It already has been established that in most areas wolves kill primarily young, old, and other inferior members of such prey populations as Dall sheep (Ovis dalli), moose (Alces alces), caribou (Rangifer tarandus), bison (Bison bison), and musk-oxen (Ovibos moschatus); evidence for this generalization has been summarized by Mech (1970).

However, only recently has it been shown that this generalization may extend to predation on the smallest hoofed prey of the wolf in North America, the white-tailed deer (Odocoileus virginianus). Pimlottet al.(1969) demonstrated a difference between the age structure of 331 deer killed by wolves during winter in Algonquin Park, Ontario, and 275 deer assumed to represent the actual population in the same area. Whereas only 13 percent of the deer from the population at large were estimated to be more than 5 years old, 58 percent of the wolf-kills were in this age category.

We employed a similar analysis for deer killed by wolves in northeastern Minnesota, but used a more refined aging technique and included comparisons of the age and sex structures of various subsamples of wolf-kills. Whereas the Ontario research involved a prey population unlimited by man, our work was carried out on both a hunted population and on one relatively unhunted. Further comparisons were made between deer killed during periods of normal snow conditions and those taken during unusually high snow accumulations. The incidence of various abnormalities in wolf-killed deer was also comparedwith that in hunter-killed animals.

The study was carried out in the Superior National Forest in northern St. Louis, Lake, and Cook Counties of northeastern Minnesota (fig. 1), in conjunction with other aspects of wolf research (see Mechet al.p. 1).

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