Part 2

No. 1057 and No. 1059 were both members of packs. No. 1057 was captured during the night after a pack of 13 wolves was seen heading toward the area; 5 days later she was seen with 10 other wolves, which no doubt represented this same pack. This wolf's association with the pack was interrupted, however, because of the foot injury sustained during capture. When 1059 was caught, tracks of two other wolves were seen in the immediate vicinity, and one of the animals was seen within a quarter mile of the trapped wolf. Three days after 1059's release, and perhaps sooner, she was back with her pack, with which she remained at least through March.

The detailed histories of the associations of the radiotagged wolves will be discussed in a later section.

Radiotagged wolves were tracked every day that weather permitted during December, January, and February; every week during March, April, and May; and once a month during June, July, and August (fig. 16). Information was obtained for a total of 570 "wolf-days"--a wolf-day being a day in which one radiotagged wolf was located; a pack of five being located for 1 day would constitute 5 wolf-days.

The last day that animals 1051 and 1057 were heard from was April 24, 1969. Both had traveled long distances during the previous week and may have moved out of range. Signals from wolf 1055 were last heard on May 30; this animal had also been ranging widely. Circles with radii of at least 50 miles around the last known locations of each wolf were searched unsuccessfully for the signals. During all subsequent tracking nights for the remaining wolves, the missing animals were also sought, but to no avail. Before the last dates that signals from these animals were heard, attempts to locate marked animals from the air had failed in only three instances.

FOOTNOTES:

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

Daytime Activity Patterns

When radiotagged wolves were located, notes were kept on the type of activity they were engaged in; the results are summarized in figure 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 more before 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.

Movements and Range

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, Pimlott _et 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 in table 5 should 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_

#: _Number_ %: _Percent_

:---------------------------- : : Net weekly : Net daily distances : distances :----------------------------------------------:---------------------------- : : : : :Mean net : : : : : : : : :distance : : : : : Days :Days no : Days :Mean net:per day : :Weeks:Mean net: Wolf: data : net :movement:distance:excluding :Range:data :distance: Range # :obtained:movement: :per day :days of no: : :per week: : : : : :movement : : : : :--------:--------:--------:--------:----------:-----:-----:--------:------- : # : # % : # % : Miles : Miles :Miles: # : Miles : Miles

1051 54 13 24 41 76 2.0 2.6 0.0-12.0 22 12.7 1.0-46.0 1053 37 20 54 17 46 1.0 2.1 0.0-5.0 23 6.3 0.0-23.6 1055 46 7 15 39 85 2.9 3.6 0.0-12.8 21 15.6 1.7-49.0 1057 29 11 38 18 62 1.0 1.5 0.0-4.5 15 4.6 0.0-31.0 1059 26 1 4 25 96 2.5 2.6 0.0-5.6 18 2.9 0.0-4.6 ----:--------:--------:--------:--------:----------:-----:-----:--------:-------

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_

+-------+-------------------------+-------------------------+ | | Mean net | Greatest net | | Wolf | daily distance | daily distance | |number +------------+------------+------------+------------+ | | Feb. 1-23 | Feb. 23-28 | Feb. 1-23 | Feb. 23-28 | +-------+------------+------------+------------+------------+ | 1051 1.1 3.5 2.3 4.8 | | 1053 0.7 2.5 3.0 5.0 | | 1055 2.7 6.2 8.0 12.8 | | 1057 1.0 1.5 4.0 4.5 | | 1059 2.2 3.1 4.0 5.6 | +-------+------------+------------+------------+------------+

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, Jordan _et 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_

+--------+----------+----------+--------------+----------------------+ | Wolf | Greatest | Greatest | Total area[9]|Area[9] of intense use| | number | length | width | | (before late Feb.) | +--------+----------+----------+--------------+----------------------+ | _Miles_ _Miles_ _Sq. miles_ _Sq. miles_ | | | |1051[10] 28.5 13.6 318 13 (Location A[11])| | 45 (Location B) | | 16 (Location C) | |1053 31.1 22.0 392 31 | |1055 55.4 24.9 997 40 | |1057 32.3 3.8 77 14 | |1059[12] 8.4 8.0 43 39 | +--------+----------+----------+--------------+----------------------+

FOOTNOTES:

[9] Minimum area method (Mohr 1947).

[10] Before dispersal.

[11] See text and figure 19.

[12] Pack of five.

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.

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 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).

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.

_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).

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, Omaday Lake, 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.

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.

Wolf Associations, Social Behavior, and Reproduction

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.

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_

+------------+--------+------------+---------+--------------- | | |Mean size of| Largest | Authority Area |Observations| Wolves | population |pack size|calculated from | | | unit | | ---------+------------+--------+------------+---------+--------------- _Number_ _Number_

Alaska 310 1,041 3.4 12 Kelly 1954 Alaska 1,268 4,823 3.8 21 R. A. Rausch[13] Lapland 118 311 2.5 12 Pulliainen 1965 E. Finland 460 984 2.1 12 Pulliainen 1965 Minnesota 112 318 2.8 12 Stenlund 1955 Minnesota 77 323 4.2 13 Present study ----------------------------------------------------------------------

FOOTNOTES:

[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.