FOOTNOTES:

FOOTNOTES:[8]David W. Kuehn, Personal correspondence to L. D. Mech, 1970.

[8]David W. Kuehn, Personal correspondence to L. D. Mech, 1970.

When radiotagged wolves were located, notes were kept on the type of activity they were engaged in; the results are summarized infigure 17. In a total of 171 observations made between 9:00 a.m. and 6:00 p.m., the wolves were resting 62 percent of the time, traveling 28 percent and feeding 10 percent. They tended to travel morebefore 11:00 a.m. and after 3:00 p.m., although resting still composed at least 45 percent of the activity during every hour (fig. 18).

These results generally agree with the statement by Mech (1966a) that wolves on nearby Isle Royale tend to rest about 11:00 a.m. and begin traveling again about 4:00 p.m. However, it does appear that the Minnesota wolves spend much more of the day resting than do the Isle Royale animals. The difference may be caused by the difference in pack sizes studied. The Isle Royale pack of 15 to 16 may have had to travel more to find enough food to feed all its members than did the lone wolves and pack of five in the present study.

Figure 17.—Percentage of time spent by radiotagged wolves in various types of activity throughout the day, from December through April.Figure 17.—Percentage of time spent by radiotagged wolves in various types of activity throughout the day, from December through April.

Figure 18.—Generally the wolves rested during most of the day. (Photo courtesy of L. D. Mech.)Figure 18.—Generally the wolves rested during most of the day. (Photo courtesy of L. D. Mech.)

Wolf movement is greatly hindered by deep, soft snow, so during winter travel, wolves frequently use areas where they sink into the snow the least. In our study area, frozen waterways are used extensively where possible, just as reported by Stenlund (1955). Where few lakes or rivers exist, wolves follow railroad beds and logging roads, often soon after a plow or other vehicle has driven on them. In cutting cross country through deep snow, wolves travel single file and tend to stick to windblown ridges and to trails of deer and moose. Wolves that have ranges small enough to cover in a few days form a network of their own trails, which they can maintain merely by traveling regularly over them. Packs on Isle Royale depended a great deal on such a system of trails (Mech 1966a), and so did Pack No. 1059 in our study area.

Wolf packs can travel up to 45 miles in a day but it is usually larger packs that do so (Stenlund 1955, Burkholder 1959, Mech 1966a, Pimlottet al.1969). In our study area we sometimes saw evidence of long moves by large packs along strings of lakes and waterways. However, most of our movement data pertain to lone wolves and a pack of five. The daily travel of these animals was usually much less than that reported for large packs.

Our radiotracking data provide an index to the extent of travel for each wolf rather than the actual amount of travel, for it is based on straight line distances between consecutive points at which an animal was found. This measure will be referred to as the "net daily distance."

Much variation was found in the net daily distances of wolves, with the longest ranging from 4.5 miles for 1057 to 12.8 for 1055 (table 3). The mean net daily distance for each animal, excluding days with no net movement, varied from 1.5 to 3.6 miles. The movements of these wolves may have been affected by the snow depth and penetrability, for mean and maximum net daily distances suddenly increased for all animals between February 23 and 28, when snow penetrability had decreased to a point where walking wolves would be expected to sink in only about 6 inches (table 4). Other possible explanations for the wolves' sudden increase in movements will be discussed below.

The straight line distances traveled between consecutive weekly locations (called the "net weekly distances") showed a similar variation (table 3). The maximum net weekly distance for each wolf varied from 4.6 miles for 1059 to 49.0 for 1055, with means ranging from 2.9 to 15.6 miles for the same wolves. No doubt 1059's net weekly distances were relatively short because her total range and that of her pack were much smaller than those of the other wolves.

It is difficult to obtain comparable measures of the extent of the ranges covered by each of the radiotagged wolves because their patterns of travel varied so much. Thus the figures given intable 5should be regarded only as gross indicators of the minimum range of each animal. The area figures are especially deceiving in the case of 1055, for she had a horseshoe-shaped range, much of which apparently was not used.

Table 3.—Straight line distances (miles) between consecutive locations of radiotagged wolves

WolfnumberNet daily distancesNet weekly distancesDaysdataobtainedDays nonetmovementDaysmovementMean netdistanceper dayMean netdistanceper dayexcludingdays of nomovementRangeWeeksdataMean netdistanceper weekRangeNumberNumberPercentNumberPercentMilesMilesMilesNumberMilesMiles105154132441762.02.60.0-12.02212.71.0-46.0105337205417461.02.10.0-05.0236.30.0-23.610554671539852.93.60.0-12.82115.61.7-49.0105729113818621.01.50.0-04.5154.60.0-31.01059261425962.52.60.0-05.6182.90.0-04.6

Nevertheless, one major piece of information is obvious from the figures: 1059's pack of five wolves had a much smaller range than any of the other uninjured animals—approximately 43 square miles when figured by the minimum-area method (Mohr 1947). The next smallest range was that of 1051 (excluding the area of his later dispersal—see below), which was some seven times the size of the pack's range.

Table 4.—Straight line distances (miles) traveled between consecutive days ("net daily distance") by radiotagged wolves in northeastern Minnesota during February 1969

WolfnumberMean net daily distanceGreatest net daily distanceFeb. 1-23Feb. 23-28Feb. 1-23Feb. 23-2810511.13.52.34.810530.72.53.05.010552.76.28.012.810571.01.54.04.510592.23.14.05.6

There is little published information on the movements and ranges of lone wolves with which to compare our data. Mech (1970) summarized information regarding ranges of packs. Reported ranges varied from 36 square miles for a pack of two wolves in Minnesota (Stenlund 1955) to 5,000 square miles for a pack of 10 in Alaska (Burkholder 1959). Considering only data based on intensive study in the same general region (Minnesota, Isle Royale, and Ontario) as our study area, the largest range reported was 210 square miles for a pack of 15 to 21 wolves on Isle Royale (Mech 1966, Jordanet al.1967). On a per-wolf basis, the ranges in this region varied from 6 to 28 square miles per wolf. Our pack of five with its range of 43 square miles would have about 9 square miles per wolf.

A more accurate assessment of the ranges of the radiotagged wolves requires an individual discussion for each.

No. 1051.—The range of 1051 was composed basically of three distinct areas (fig. 19). Within 10 days after being released, the wolf left the general area of his capture (Area A near Isabella Lake) and traveled to Area B along Highway 1, some 17 miles to the southwest. From December 9 to January 4 wolf 1051 remained in Area B, which covers about 45 square miles. Between January 4 and 6 he returned to Area A and stayed in 13 square miles until February 3. Between February 3 and 5 he shifted to Area C east of Snowbank Lake, 11 miles northwest of Area A. He remained in that 16-square-mile area until February 25, then suddenly left and headed 8 miles to the northeast.

Table 5.—Extent of ranges used by radiotagged wolves

WolfnumberGreatest lengthGreatest widthTotal area[9]Area[9]of intense use(before late Feb.)MilesMilesSq. milesSq. miles1051[10]28.513.631813 (Location A[11])45 (Location B)16 (Location C)105331.122.039231105555.424.999740105732.33.877141059[12]8.48.04339

FOOTNOTES:[9]Minimum area method (Mohr 1947).[10]Before dispersal.[11]See text and figure19.[12]Pack of five.

[9]Minimum area method (Mohr 1947).

[10]Before dispersal.

[11]See text and figure19.

[12]Pack of five.

Figure 19.—Locations and range of wolf 1051. Lines are NOT travel routes; rather they merely indicate sequence of locations. Only selected lakes are shown.Figure 19.—Locations and range of wolf 1051. Lines are NOT travel routes; rather they merely indicate sequence of locations. Only selected lakes are shown.

From February 26 until April 24 the movements of 1051 were strongly indicative of dispersal (fig. 20). His average weekly straight line move during that period was 25 miles (compared with 6 miles per week before this period), and until March 14 he maintained an almost straight south-southwest heading to a location west of the town of Castle Danger. After that the animal traveled a series of northwest-southwest alternations that on April 3 took him east of Big Sandy Lake to a point 129 miles southwest of where he had begun the dispersal. There he remained for about 2 weeks, but between April 17 and 24 he traveled 26 miles northwest. We last saw him at 3:30 p.m. on April 24 heading northwest through a swamp 15 miles southeast of Grand Rapids, approximately 122 miles from where he had started. The total of straight line distances between 16 consecutive pairs of locations taken at intervals of from 1 to 8 days was 226 miles, which is the minimum distance the wolf traveled during his dispersal.

Figure 20.—Dispersal of wolf 1051. Lines merely indicate sequence of locations. Only selected lakes are shown.Figure 20.—Dispersal of wolf 1051. Lines merely indicate sequence of locations. Only selected lakes are shown.

We observed 1051 for distances of up to 5 miles during these travels; he maintained a steady trot that seemed faster than usual, and he appeared intent on heading in a straight line. He did chase deer during his travels, and twice was seen feeding on carcasses. In the area where he remained for about 2 weeks, he was twice seen closely associated with another wolf. This relationship will be discussed later.

An extensive search was made for 1051's signals on May 2 in an area of at least 50 miles radius from his last known location, but it was unsuccessful. On each subsequent tracking flight, the wolf's frequency was also monitored with no success. Possible explanations for the loss of the signal from this wolf include the following: (1) premature expiration of the transmitter, (2) capture of the wolf and breakage of the transmitter, (3) loss of the exposed antenna and consequent reduction of range, and (4) travel of the wolf out of range of the tracking aircraft.

During 1051's travels a number of interesting events took place:

Nov. 27, 1968—Captured and radiotaggedDec. 4, 1968—Crossed road in front of tracking truckDec. 8, 1968—Moved to Area BDec. 9, 1968—Surprised on the ground at distance of 35 feetDec. 18, 1968—Chased by loggers with axesDec. 25, 1968—Almost shot by trapper who saw collar and withheld fireJan. 5, 1969—Returned to Area AJan. 13, 1969—"Bumped" twice on logging road by loggers in auto but no apparent injuryFeb. 4, 1969—Moved to Area CFeb. 26, 1969—Began long-range southwest movement considered to be dispersalMar. 14, 1969—Seen feeding on old carcass within 200 yards of houses, dogs, and a man walkingMar. 27, 1969—Chased two deer across 4-lane State highway 53Apr. 3, 1969—Found with another wolf at point farthest south in his rangeApr. 24, 1969—Last contact with this animal; was seen traveling NW

Nov. 27, 1968—Captured and radiotagged

Dec. 4, 1968—Crossed road in front of tracking truck

Dec. 8, 1968—Moved to Area B

Dec. 9, 1968—Surprised on the ground at distance of 35 feet

Dec. 18, 1968—Chased by loggers with axes

Dec. 25, 1968—Almost shot by trapper who saw collar and withheld fire

Jan. 5, 1969—Returned to Area A

Jan. 13, 1969—"Bumped" twice on logging road by loggers in auto but no apparent injury

Feb. 4, 1969—Moved to Area C

Feb. 26, 1969—Began long-range southwest movement considered to be dispersal

Mar. 14, 1969—Seen feeding on old carcass within 200 yards of houses, dogs, and a man walking

Mar. 27, 1969—Chased two deer across 4-lane State highway 53

Apr. 3, 1969—Found with another wolf at point farthest south in his range

Apr. 24, 1969—Last contact with this animal; was seen traveling NW

Wolf 1053.—This wolf was basically a scavenger who subsisted for long periods on the remains of old carcasses. She was known to have visited the remains of at least four deer and three moose, and she stayed near one moose carcass from February 8 to 20, at least during the day. Between her date of capture, December 10, and February 28, 1053 traveled about in an area of 31 square miles in the Arrow Lake-Maniwaki Lake region (fig. 21).

Figure 21.—Locations and range of wolf 1053. Only selected lakes are shown.Figure 21.—Locations and range of wolf 1053. Only selected lakes are shown.

Between February 28 and March 6 she suddenly moved 13 miles to the east-southeast near the Sawbill Trail, and during the next week she traveled a straight line distance of 24 miles southwest to a point southeast of the town of Isabella. Her subsequent travels eventually took her over a much larger area. Before February 28, 1053's average weekly straight line distance was 2 miles, but after that date it increased to 11 miles.

Wolf 1055.—The range of this animal from January 5, when she was captured, to February 23 covered about 40 square miles near Stony Lake, Slate Lake, and the Jack Pine Lookout Tower (fig. 22), and her mean weekly distance was 4 miles. Between February 23 and 24, however, she traveled 13 miles northeastward, the beginning of a series of long moves. By March 5, 1055 had reached Crescent Lake, a point 39 miles east-northeast of her previous area of intensive use. She then gradually headed back toward the west and south during the next 10 days and within the next month repeated this pattern. When her signal was heard last on May 30, 1055 was near Martin Landing in the center of her range. Her mean net weekly distance after February 23 had increased to 22 miles.

Figure 22.—Locations and range of wolf 1055. Only selected lakes are shown.Figure 22.—Locations and range of wolf 1055. Only selected lakes are shown.

Wolf 1057.—The movements of 1057 cannot be considered normal because freezing of a front foot prevented her accompanying the pack of which she was a member. Nevertheless, even data from an abnormal animal can provide some information. On January 13, 5 days after capture and release on Red Rock Lake, 1057 was located 4 miles from the capture point with a pack of 10 other wolves. She was limping and fell behind when they moved. Five days later she was again seen with the pack 12 miles away between Knife Lake and Kekekabic Lake. She then remained in about 14 square miles of that general area through April 17 (fig. 23).

Figure 23.—Locations and range of wolf 1057. Only selected lakes are shown.Figure 23.—Locations and range of wolf 1057. Only selected lakes are shown.

Suddenly on April 24, 1057 was found in Ontario some 31 miles northeast of her location of the previous week. That was the last time we heard her signal even though on May 2 we scanned an area with a radius of 35 miles from her last known location and listened for her signal during every subsequent flight.

Wolf 1059.—This animal was a member of a pack of three to five wolves (see next section). The movements of the group varied little and were concentrated in the August Lake, OmadayLake, and Keeley Creek area in about 43 square miles (fig. 24). Contrary to animals 1051, 1053, and 1055, this pack did not suddenly begin a series of longer weekly movements in late February. Both before and after February 28, the average weekly straight line movement of the pack was just less than 3 miles.

Figure 24.—Location and range of wolf 1059 and pack. Only selected lakes are shown.Figure 24.—Location and range of wolf 1059 and pack. Only selected lakes are shown.

Probably these animals did begin traveling more in late February, for their net daily distances did increase at that time along with those of the other wolves (table 4). However, the increased travel took place within the restricted area of the pack's usual range rather than in new areas as occurred with the other wolves.

Because 1059 was later found to have bred and carried five fetuses, her movements during whelping season (late April and early May) are of interest. Her locations on both April 24 and May 2 were within 250 yards of each other, which might indicate that she was denning. On May 9, however, she was 2.5 miles east of these locations, on the 17th and 21st was 2 miles west of them, and on the 30th was 3 miles north of them.

In early January 1970, Wolf 1059 was killed by a trapper in the southeast corner of her pack's 1969 range.

Summer locations.—Signals from only 1053 and 1059 were heard during summer, and then tracking attempts were made only on June 28, July 29, and August 29. Locations for 1053 on those occasions were near Kelly Landing and Isabella Lake, within her previous range. Wolf 1059 was found each time within 2 miles outside of the southwest corner of the pack's winter and spring range.

In our study area, population units of wolves exist as both single animals (lone wolves) and packs. In a total of 77 observations, lone wolves constituted 32 percent of the sightings (fig. 25), with packs of from 2 to 13 members making up the remainder (Table 1). On the basis of the number of wolves seen, rather than the number of observations, lone wolves accounted for only 25 (8 percent) out of 323.

Figure 25.—Only 8 percent of the wolves observed were lone wolves. (Photo courtesy of L. D. Frenzel.)Figure 25.—Only 8 percent of the wolves observed were lone wolves. (Photo courtesy of L. D. Frenzel.)

These figures compare favorably with reports in the literature as summarized by Mech (1970). In five areas studied, lone wolves made up from 24 to 60 percent of the observations of population units, and from 8 to 28 percent of the wolves seen. In our study area during 1948 to 1953, lone wolves constituted 43 percent of the observations and 15 percent of the wolves (Stenlund 1955).

The average size of the population units observed during our study (total number of wolves seen divided by the number of observations) was 4.2, which is significantly larger (95 percent level) than the average seen in this area (2.8) from 1948 to 1953. This is also larger than that reported from any other area of comparable size (table 6).

Table 6.—Mean sizes of wolf population units reported from various areas

AreaObservationsWolvesMean size ofpopulation unitLargestpack sizeAuthoritycalculated fromNumberNumberAlaska3101,0413.412Kelly 1954Alaska1,2684,8233.821R. A. Rausch[13]Lapland1183112.512Pulliainen 1965E. Finland4609842.112Pulliainen 1965Minnesota1123182.812Stenlund 1955Minnesota773234.213Present study

FOOTNOTES:[13]R. A. Rausch. Personal correspondence to L. D. Mech, 1967.

[13]R. A. Rausch. Personal correspondence to L. D. Mech, 1967.

The largest pack seen in our study area included 13 members, and there apparently were at least two such packs. Although larger packs than this have been reported, any group containing more than 8 to 10 members is unusually large (Mech 1970).

Wolf sociology is a complex subject and is still not well understood, so the following detailed observations of the associations between our radiotagged wolves and others are given. Associations are defined as relationships in which two or more wolves relate in a close, positive manner.

As mentioned earlier, 1051 may or may not have been associated with other wolves when he was captured. However, although this animal was observed 55 times throughout winter and spring, only twice was he seen associating with another wolf. Probably the same individual was involved each time, because the location was about the same (the vicinity of the juncture of Aitkin, Carlton, and St. Louis Counties).

The first occasion was on April 3. Wolf 1051 in the previous week had moved 46 miles straight line distance from the northeast. He was then observed lying peacefully within 15 feet of another wolf near a freshly killed deer. The very proximity of the two animals implied a positive relationship. On April 7, 10 and 14, 1051 was seen 1 mile, 10 miles, and 8 miles from the kill and was alone each time.

However, on April 17, 1051 was back in the general vicinity of the kill, and he and another wolf were resting on an open hillside about 100 feet from each other. As we descended for a closer look, the smaller animal arose and headed to the larger, presumably 1051 because he had not been disturbed by the aircraft. The larger wolf did not arise for several seconds, but eventually followed the other into the woods. No tail raising or other expressive posturing was seen in either wolf. One week later 1051 was 26 miles northwest of the kill traveling alone.

Wolf 1053 was never seen less than 80 yards from another wolf, and there was no evidence that she ever associated with a conspecific. Even when she was seen 80 yards from the other wolf, both were resting, and when the strange wolf left, 1053 made no attempt to accompany or follow it.

No. 1055 apparently had been traveling with another wolf when caught on January 5, and tracks showed that the individual had remained near her until we arrived to handle her. Tracks found on January 7 and 10 suggested that 1055 was with another animal, but that animal was not seen during any of the six times 1055 was observed through February 1. However, from February 5 to 19, 1055 was with another wolf on eight of the 12 times she was seen. The two animals were observed resting, traveling, hunting, and feeding together. On February 20, and thereafter, 1055 was alone all 14 times she was seen.

It is possible that 1055's associate was killed between February 19 and 20. About March 6, a 63-pound male wolf pup was found dead (by Mr. Charles Wick, USDA Forest Service) within about 50 feet of a highway and less than a mile from where 1055 and her associate were seen on February 19. Because of the snow conditions, it was judged that the wolf had been killed (probably by an automobile) sometime in February.

Wolf 1057, whose foot froze during capture, was a member of a pack of 10 to 13 wolves, and was seen with the pack on January 13 and 18. After that she was usually found alone, although on at least five occasions she was with one or more wolves:

PeriodNo. ofobservationsAssociationsJan. 13110 other wolvesJan. 14-171NoneJan. 18110 or 11 other wolvesJan. 19-292NoneJan. 3011 other wolfJan. 31 to Feb. 22NoneFeb. 3-422 other wolvesFeb. 511 other wolfFeb. 6-136NoneFeb. 1413 other wolvesFeb. 15-225NoneFeb. 23110 to 13 other wolvesFeb. 24 to Apr. 246None

February 23 she was with the pack at a kill in her usual area, and although the pack left that night, 1057 remained near the kill the next day. Presumably this animal would have traveled with pack if she could have.

Figure 26.—One of the radiotagged wolves was a member of this pack of five. (Photo courtesy of L. D. Mech.)Figure 26.—One of the radiotagged wolves was a member of this pack of five. (Photo courtesy of L. D. Mech.)

No. 1059 was part of a pack that included three to five members (fig. 26). From January 25, the first time she was observed after release, through April 2, the animal was seen 19 times with two other wolves, eight times with at least three others, and eight times with four others. She was never seen alone until April 17; both times after this when she was seen, May 9 and 21, 1059 was also alone.

Some insight into the fluctuating size of this pack was obtained on February 27 when the five animals were followed for 2 hours. During that time two members (one of which was larger than the other) often lagged behind the other three by as much as a mile. These two romped and played considerably, with one carrying a stick or a bone part of the time. Eventually they caught up again to the other three. The behavior of the two lagging wolves would be consistent with the hypothesis that they were either pups or a courting pair of adults. In either case, they seemed to be an actual part of the pack even though they temporarily traveled separately.

The fact that 1059 was observed traveling alone three times from April 17 to May 21 may be further evidence that the pack had a den in the area at that time. The presence of a den allows individual pack members to venture off singly and return each day to a known social center, as Murie (1944) observed, so they do not need to travel with each other to maintain social bonds. Wolves in our area breed during the latter half of February (see below), and the young should be born in the latter half of April. Since dens are prepared a few weeks in advance (Young 1944), pack members might be expected to begin traveling singly in mid-April.

Some information on social relations within our radiotagged pack of five was also obtained.One of the members could often be distinguished from the others by its reddish cast and this individual appeared to be the pack leader or alpha male (Schenkel 1947). In urinating, this animal lifted his leg, a position seen almost exclusively in males. Except for only two temporary occasions, this animal always headed the pack, which usually traveled single file. The second wolf in line generally was noticeably small, possible a female, and the third wolf was twice identified as 1059 on the basis of sightings of her collar.

The leader often gained a lead on the other wolves, especially during a chase (see below), much as reported for a lead wolf on Isle Royale (Mech 1966a). Upon returning to the lagging members of the pack, this animal usually held his tail vertically, an expression of social dominance (Schenkel 1947). On two occasions he led chases against strange wolves and demonstrated the highest motivation (see below).

Figure 27.—Feces, urine, and scratching in a conspicuous spot indicate a wolf "scent post." (Photo courtesy of L. D. Mech.)Figure 27.—Feces, urine, and scratching in a conspicuous spot indicate a wolf "scent post." (Photo courtesy of L. D. Mech.)

The leader was also the most active in his reactions when scent posts were encountered. Because the function of scent-marking behavior is still unknown, it is important that detailed descriptions of the natural behavior of free-ranging wolves around scent posts be made available (fig. 27). Thus the following excerpt from field notes by Mech dated February 27, 1969, is presented:

"When they [the three wolves] came to a small frozen pond, where the wolf trail [which they had been following] branched and there were some packed down areas, they became quite excited [fig. 28]. This was especially true of the reddish wolf. He nosed several spots, and scratched around them. Usually his tail was vertical. He defecated at one spot, and right afterwards another wolf did. After about 2 minutes that pack went on.Figure 28.—A pack of wolves investigating a scent post. The raised tails indicate their excitement. (Photo courtesy of L. D. Mech.)Figure 28.—A pack of wolves investigating a scent post. The raised tails indicate their excitement. (Photo courtesy of L. D. Mech.)"About 15 minutes later the 2 'satellite' wolves arrived at this spot, hesitated, nosed around but continued on after less than a minute."The three wolves meanwhile came to a junction of 2 logging roads. There they nosed around, scratched, and acted much as described above. Again the reddish wolf was most active and had its tail up."When the last 2 wolves came to this spot, they nosed around, ran back and forth, and 1 defecated. They then headed on a different branch of the trail than the first 3 had gone on just 10 minutes before."The first 3 wolves meanwhile were running along a logging road but eventually they circled and one other than the reddish one headed across a swamp toward the last 2. Then the reddish one and the other followed this one, and they met the last 2 on a ridge. There was the usual tail wagging, then all headed off together in a new direction. They passed the first scent post again and there was some nosing by the reddish wolf but little hesitation."When they traveled, one wolf lagged behind by 150 yards. The wolf just ahead of it had its tail vertical part of the time, as did the reddish leader.

Figure 28.—A pack of wolves investigating a scent post. The raised tails indicate their excitement. (Photo courtesy of L. D. Mech.)Figure 28.—A pack of wolves investigating a scent post. The raised tails indicate their excitement. (Photo courtesy of L. D. Mech.)

"About 15 minutes later the 2 'satellite' wolves arrived at this spot, hesitated, nosed around but continued on after less than a minute.

"The three wolves meanwhile came to a junction of 2 logging roads. There they nosed around, scratched, and acted much as described above. Again the reddish wolf was most active and had its tail up.

"When the last 2 wolves came to this spot, they nosed around, ran back and forth, and 1 defecated. They then headed on a different branch of the trail than the first 3 had gone on just 10 minutes before.

"The first 3 wolves meanwhile were running along a logging road but eventually they circled and one other than the reddish one headed across a swamp toward the last 2. Then the reddish one and the other followed this one, and they met the last 2 on a ridge. There was the usual tail wagging, then all headed off together in a new direction. They passed the first scent post again and there was some nosing by the reddish wolf but little hesitation.

"When they traveled, one wolf lagged behind by 150 yards. The wolf just ahead of it had its tail vertical part of the time, as did the reddish leader.

"Soon the pack came upon another area packed with wolf tracks on a pond. There they followed every little trail, nose to the ground, wagged tails, grouped together often, chased each other, rolled over, etc. for 6 minutes. The reddish animal had tail up most of the time."The wolves continued on, and we left them about 1 mile S.W. of the S.W. arm of Bald Eagle Lake [at 6:05 p.m.]."

"The wolves continued on, and we left them about 1 mile S.W. of the S.W. arm of Bald Eagle Lake [at 6:05 p.m.]."

Unfortunately it was not known whether the trails that the wolves were following were their own or those made by other wolves.

Significant aspects of the above observation are (1) the spirited initiative of the leader, (2) the amount of time spent in scratching, urinating, and defecating, (3) the decision of the last two wolves to take a different route from that of the first three even though their goal seemed to be to catch up to the first three, and (4) the fact that the scent posts were located at trail junctions. In the last regard, we often noted from the ground that wolves urinated at the junction of newly formed human trails heading perpendicularly from roads they were following.

Copulation in wolves was only observed once during our study, on February 19, 1969. Two members of a group of four were seen coupled for 2 minutes on Kekekabic Lake. On Isle Royale, which is at the same latitude, copulations were witnessed on February 21, 24 and 27 (Mech 1966a).

On April 17, a den west of Big Moose Lake known to have been used at least intermittently for 13 years was seen from the air to have fresh activity of some kind in the snow in front of it, and on April 24 we saw a wolf at the mound. A few days later, two local human residents unaware of our interests approached this den and looked in. An adult wolf, presumably the bitch, leaped over their heads and fled the area. The men then dug up the den and removed six pups whose eyes had not yet opened.

Instances of chasing or attack by a pack of wolves on conspecifics not a part of their group have been described by Murie (1944) and Mech (1966a). Observations of such behavior are important in trying to determine conclusively whether or not wolves are territorial. Pimlottet al.(1969, p. 75) wrote "It still is not clear, however, whether or not their use of range should be defined as territorial." Mech (1970) summarized the available evidence for territoriality in wolves and postulated that it may be spatiotemporal such that packs might avoid each other at any particular point in time but over a long period might cover the same area at different times. A number of our observations are pertinent to this question, for we have evidence of both, tolerance and intolerance between population units of wolves.

Two direct cases of intolerance were observed, both involving the radiotagged pack and other wolves within the usual range of the pack. Following is a direct quote from the field notes of Mech:

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