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.

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:

_No. of_

_Period_ _observations_ _Associations_

Jan. 13 1 10 other wolves

Jan. 14-17 1 None

Jan. 18 1 10 or 11 other wolves

Jan. 19-29 2 None

Jan. 30 1 1 other wolf

Jan. 31 to Feb. 2 2 None

Feb. 3-4 2 2 other wolves

Feb. 5 1 1 other wolf

Feb. 6-13 6 None

Feb. 14 1 3 other wolves

Feb. 15-22 5 None

Feb. 23 1 10 to 13 other wolves

Feb. 24 to Apr. 24 6 None

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.

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.

[Illustration: _Figure 26.--One of the radiotagged wolves was a member

of this pack of five. (Photo courtesy of L. D. Mech.)_]

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).

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:

[Illustration: _Figure 27.--Feces, urine, and scratching in a

conspicuous spot indicate a wolf "scent post." (Photo courtesy of

L. D. Mech.)_]

"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.

[Illustration: _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.]."

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.

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