Part 2

We had scheduled the release for mid-March for several reasons which we felt would maximize chances for success. Deer are concentrated then in the Huron Mountain area and vulnerable to predation. Pregnancy and subsequent whelping of the alpha female might increase her attachment to the new area. Furthermore, the snow is usually deepest then and hinders travel. However, a few days before the release, a freak rainstorm had settled the snow, and cold temperatures had frozen it so hard that animals could walk readily on top, making travel conditions excellent.

An observation shack 120 feet (36.6 m) from the pen was used to determine the activities and interactions of the four wolves. Weise spent three nights in the shack and also observed the wolves each day of the one-week penned period, for a total of 20 hours of observation (Fig. 14).

During preliminary air and ground checks of radio equipment, we discovered that Wolf No. 10 had a defective collar. Thus on March 12, we subdued her with a choker, restrained her with ropes, replaced her collar and released her just after sunset. We then opened the pen, and let the other wolves loose.

The subsequent locations of the wolves were then checked intermittently through aerial radio-tracking (Mech 1974), with a receiver and antenna from the AVM Instrument Co., Champaign, Illinois, used in a Cessna 172 and a Piper Colt. We made two flights each day for the first 2 days after release, one each day when weather permitted, until April 20, three per week in May, approximately two per week from June through September, and three per week in October and November. A total of 194 hours were flown, 80 per cent by Weise, and the remainder by Hook. Aerial locations were usually recorded to the nearest 40 acres (16 ha.).

We also tracked the wolves from the ground whenever interesting or significant activities were observed during flights or were reported by ground observers. Carcasses of prey animals were investigated from the ground after consumption was complete and the wolves had left. Deer eaten by the wolves were considered killed by them if the ground check revealed fresh blood or flesh, or signs of a struggle. Scats were collected along the tracks of the wolves in the snow whenever possible.

When radio signals were received from the same location for unusually long periods, ground checks were made to determine the cause.

Attempts were made to verify all sightings and track records reported by local citizens, by comparing them with the aerially-determined locations.

RESULTS

Social Structure of the Translocated Wolves

Wolves No. 11, 12, and 13 were captured in Minnesota within a mile (1.6 km) of each other, and No. 11 and 12 were taken in the same trap set 12 days apart; Wolf No. 10 was caught approximately 7.5 miles (12.1 km) southeast of the others (Table 2). All were judged to be thin but in good condition.

Females No. 10 and 11 were introduced into the same pen on January 23. No. 11 was reluctant to enter the pen containing No. 10 while several observers were around, but entered within 15 minutes after all but one had left. No. 11 went directly to No. 10 which was lying in a corner as she usually did, and pawed the fence at No. 10's back. When the pawing became more vigorous, No. 10 snapped at No. 11, moving only her head and neck. No. 11 then turned directly to No. 10, sniffed the top of her head and mane, and lay down beside No. 10 with her nose still in No. 10's mane. No. 10 remained still throughout the whole process. The trapper reported that later No. 11 licked the face of No. 10. Sniffing and licking anteriorally are usually signs of intimacy between wolves (Schenkel 1947).

The two male wolves (No. 12 and No. 13) were allowed into the pen with the two females on February 4. No. 13 remained in the original adjoining pen and did not move in with the females immediately, but No. 12 did. There were no signs of aggression among any of the four wolves. No. 11, 12, and 13 moved freely around the pen while in Minnesota, but No. 10 most often lay in one corner by herself.

Trapper Himes first observed vaginal bleeding in female 11 on February 7. He observed Wolves 11 and 12 mating (with normal coupling) on February 12 and 16.

No unusual aggressive or agonistic social interactions of consequence were observed among the wolves while penned in Michigan, from March 5 to 12. Animals 11, 12 and 13 would lie down and feed together in various combinations. No. 10 was less active than the others and often stayed inside a shelter box within the enclosure, but would come out and mix with the other wolves for brief periods when humans were not in evidence. Her actions were indicative of a low ranking, immature, distressed, or alien animal.

Male No. 12 was the only wolf that would stare directly at a person approaching the pen. He was bolder and more direct in his actions than any of the other animals. This is the wolf that mated with adult female No. 11 while penned in Minnesota, and thus can be considered the "alpha male," or pack leader.

When approached by humans, all the wolves would urinate and defecate; No. 11 and 12 would pace, No. 10 (when out of the shelter box) and No. 13 would lie in the far corner of the pen and remain motionless (Fig. 14). No. 11 limped on her right front foot throughout the penned period, but this limp did not appear to have a significant effect on her activities or movements.

Blood samples taken on March 5, 1974 were analyzed and interpreted by Dr. U. S. Seal of the Veterans Administration Hospital in Minneapolis. The assays performed included hematology, 16 blood chemistries, thryoxine, and cortisol (Seal et al., 1975), plus estrogen and progesterone. According to Seal (personal communication), all blood values for wolves No. 10, 12, and 13 were similar and indicative of good health and minimal stress, as indicated by very low levels of the enzymes LDH, CPK, and SGOT. Such levels are typical of animals in a state of good nutrition that have been in captivity for several weeks and have accepted their captive circumstances. The MCV's were normal, indicating no vitamin deficiency, and the MCHC showed full hemoglobin content in the red cells, indicating no lack of iron. The white blood cell counts were much lower than usually seen in newly trapped wolves. All the remaining chemistry values from these three wolves were in the normal range for the season.

Wolf No. 11, however, differed in that she had a much higher hemoglobin level, higher blood glucose and white cell count, and higher levels of LDH, CPK, and SGOT, indicating that she was significantly stressed. This is corroborated by a low thyroxine level of 0.6 micrograms percent, which is hypothyroid for wolves.

The fibrinogen levels of all four animals were normal, indicating that there was no acute or chronic inflammation in progress.

The wolves ate well in captivity but still lost from 11% to 20% of their capture weight (Table 2). Himes estimated that they consumed an average of 8 lb. (3.6 kg) of food per wolf per day, while penned in Minnesota. In Michigan the wolves consumed about a deer and a half, or an estimated 5.5 lb. (2.5 kg) per wolf per day. These estimates fall within the range of food consumption figures estimated for wolves in the wild (Mech and Frenzel 1971). After the wolves began feeding on the first carcass, they completely consumed it before starting a second one, even though four carcasses were available; they ate nothing from the other two carcasses.

We released the wolves at dusk on March 12, 1974. Having just restrained Wolf No. 10 without drugs, to replace her collar, we untied her and let her free; she bounded off northwestward. We then opened the pen, and No. 12, whom we had judged to be the alpha male, left in less than 5 minutes and trotted off steadily toward the west-southwest. The remaining two animals paced around the pen for about 5 minutes and then lay down. Because we felt that they might become too widely separated from the others, three of us approached the pen opposite the door to encourage the wolves to find the open gate. Five minutes later No. 13 left the pen running southwestward, and No. 11 left less than 5 minutes later. Upon exiting, No. 11 appeared to smell the track of No. 12 and slowly trotted in his direction.

Aerial Tracking

Our success in locating the translocated wolves by aerial radio-tracking was 95% (Table 3), similar to that of Mech and Frenzel (1971) working with wolves in their native range in Minnesota.

During the part of the study in which extensive snow cover was present (March 13 to April 20) wolves No. 11, 12, and 13 were observed 14 times from the aircraft. The first time they were seen, near Laws Lake, they appeared alarmed and moved into heavy cover. The next day, however, and on all subsequent observations, the aircraft appeared to have little effect on their behavior, although they sometimes looked up at it. No. 10 was seen only once by a passenger in the tracking aircraft, and she immediately hid from view. It seems likely that she avoided the aircraft. After the snow melted and leaves appeared, we no longer saw the wolves.

The activities of the three wolves during the 14 aerial observations were as follows: traveling 4 (Fig. 15), feeding and scavenging 5 (Fig. 16), resting 4, and sleeping 1.

_Table 3. Success in locating wolves by aerial tracking_

Wolf Number 10 11 12 13 --------------------------------------------------------- Number of tracking attempts 113 65 59 67

Number of times located 105 62 59 61

Percent located 93% 95% 100% 91%

Number of times observed 1 14(Pack)

Last date tracked Nov. 17 Sept. 19 July 10 July 27 ---------------------------------------------------------

Movements of the Translocated Wolves

Wolf No. 10 never joined any of the other radioed wolves after their release, whereas the others generally remained as a pack. Thus the movements of the pack will be described separately from those of lone wolf No. 10.

Four phases were seen in the movements of the pack: (1) Post-Release Phase, March 12 to 14; (2) Directional Movement Phase, March 15 to 24; (3) Exploratory Phase, March 25 to May 7, and (4) Settled Phase, May 7 to July 6.

Post-Release Phase

This first phase of the wolves' movements, including the first 2 days after release, seemed to be characterized by confusion and indecision. On March 13, the morning after the release, the three wolves were separated, but all remained within 2.0 miles (3.2 km) south to west of the release site, the general direction in which they had headed upon release (Fig. 17). No. 11 and 13 were about a half-mile (0.8 km) apart in the morning, and by late afternoon, No. 13 apparently had joined No. 11. No. 12 remained about 2 miles away from the others all day, although he did move about a half-mile during the day. By the 14th, No. 11 and 13 had moved 2 miles southwestward, but were separated by a half-mile; No. 12 had moved only a half-mile west.

Directional Movement Phase

During this phase, all three wolves left the immediate vicinity of the release point and headed southwestward. Early in this phase, wolves No. 11 and 13 rejoined (by March 15) and traveled 9 miles (14.5 km) west-southwest of their previous day's location, while No. 12 took a more northerly route. Nevertheless, by March 19, No. 12 had joined the other two wolves near Skanee, some 14 miles (22.5 km) west-southwest of the release point (Fig. 17). For the next several weeks these wolves all remained together and travelled a straight-line distance of about 40 miles (64.1 km) to a point just north of Prickett Dam about 11 miles (17.6 km) west-southwest of L'Anse, arriving there on March 24 (Fig. 17).

Exploratory Phase

In the Exploratory Phase of their movements, from March 25 to May 7, wolves No. 11, 12, and 13 covered a 1,631-square-mile (4,224 km^2) area from the town of Atlantic Mine on the Keweenaw Peninsula to the north to a point about 64 miles (103.0 km) south, near Gibbs City (Fig. 18). In the opposite dimension, they ranged from Keweenaw Bay on the east to 9 miles (14.5 km) south of Ontonagon, 42 miles (67.6 km) west of there. This phase was characterized by long movements, considerable zigzagging, and revisiting of certain general regions such as the base of the Keweenaw Peninsula and areas east and north of Kenton (Fig. 18).

An interesting social change also occurred during this phase: No. 13 split from the pack sometime after April 26 when the pack had reached its westernmost location, south of Ontonagon. Whereas No. 11 and 12 returned east-northeastward toward Otter Lake, where they had been in late March, No. 13 headed west-northwestward to the Porcupine Mountains, 18 miles (30.0 km) west of where the pack had last been located together (Fig. 18). Thus on May 2, Wolf No. 13 was 51 miles (82.0 km) west of No. 11 and No. 12. Nevertheless, 5 days later all the wolves were found near Gibbs City, 62 miles (99.8 km) southwest of the Porcupine Mountains, and 45 miles (72.4 km) south of Otter Lake; No. 13 was only 6 miles (9.7 km) from his packmates. The next time an attempt could be made to locate the wolves, on May 16, they had reunited.

Settled Phase

This last phase of the wolves' movements includes the period when the animals had settled into an area similar to the size of home ranges reported for other wolves in the Great Lakes Region (Mech 1970). From May 7 to July 6, this pack lived in a 246-square mile (637 km^2) area with its center north of Gibbs City (Fig. 18). On July 10, wolf No. 12 was found dead. Presumably he had died by July 6, for he had not moved since then.

Wolf No. 13 had again split from his associates between June 14 and 19 and begun to travel alone. On July 20, his remains were discovered 24 miles (38.6 km) southeast of where the pack had settled. These deaths will be discussed in detail later.

Movements of the Remaining Pack Member

After the death of her mate (No. 12), Wolf No. 11 left the 246-square-mile area in which the pack had settled (Fig. 19). By July 15, she had traveled 28 miles (45.0 km) northwest of this area and by the 20th, was back by Otter Lake, 40 miles (64.4 km) north. She returned south of Gibbs City on July 27, and was found about 3 miles (4.8 km) north of the Wisconsin border on August 2, near Lac Vieux Desert about a half mile (0.8 km) north of the Wisconsin border on August 6, and near Bruce Crossing on August 9.

On August 13, Wolf No. 11 was located 1-1/2 miles (2.4 km) southeast of Ewen on the western edge of her previous locations. She was not located again until August 28. By then she had moved a straight-line distance of almost 60 miles (96.5 km) to an area in Marquette County just south of Squaw Lake, in the Witch Lake area. In doing so, she probably had passed through the area previously explored just north of the Iron County region where the pack had spent so much of its time. These widespread movements are characteristic of lone wolves even in their native range (Mech and Frenzel 1971).

No. 11 was still in the Witch Lake area on September 2. Due to poor flying conditions we did not locate her again until September 19. At that time she was on the Floodwood Plains a quarter mile (0.4 km) north of the Floodwood Lakes. She was caught in a coyote trap during the night of September 19 and shot about 10 a.m. on September 20.

Movements of Wolf No. 10

The movements of female wolf No. 10 during the post-release phase were markedly different from those of the pack. In fact, this wolf apparently skipped the relatively sedentary post-release phase of movements that the pack displayed, and immediately dispersed (Fig. 20).

By the morning after release, No. 10 was 10 miles (16.0 km) southeast of the release point and by late afternoon was an additional 5.5 miles (8.8 km) southeast (March 13). On the night of March 15 this wolf crossed four-lane Highway 41, and on the 16th was found 1-1/4 miles (2.0 km) south of the Marquette County Airport, approximately 32 miles (51.5 km) from the release site; she had traveled a minimum of 36 miles (57.9 km) to get there. However by March 20 she had returned to within 4 miles (6.4 km) of the release point, and by the 24th was within a quarter mile of the site.

The other three wolves had already dispersed westward and were near Prickett Dam, some 40 miles (64.0 km) away. It is not known whether No. 10 tried to locate them. Her locations indicate that she did not, although she may not have been able to find or follow their route. From April 2 to 15, No. 10 made a second exploration southward, again returning to the Huron Mountain area. She also made a third such trip on June 14 to 22, even crossing Highway 41 again.

From the time of release until the first week in September, there seemed to be a pattern to the movements of Wolf No. 10. She made nine trips of about 40 miles (64.0 km) each, starting near Huron Mountain, extending southeasterly about 20 miles (32.0 km), and then returning northwesterly to the Huron Mountain area (Fig. 20).

During March, April, and the first week of May Wolf No. 10 made three of these trips roughly paralleling the Lake Superior shore, and she remained in the Huron Mountain area for several days between trips. From late May until mid-July she made four such trips but did not remain long anywhere. During that time she gradually moved westerly to near the Dead River Basin. In late July she made another trip to the Dead River Basin area after a stay near the Big Bay dump. These trips enlarged No. 10's range considerably.

Early in July, Wolf No. 10 moved almost directly south from the Huron Mountain area to the Silver Lake area, again expanding her range to the west. From September 5 until October 10 she remained in the Silver Lake area, and there was no apparent pattern to her movements then. After the wolf was located on September 15 near a bait that bear hunters had put out on the west edge of the Mulligan Plains, a ground check was made. No evidence of the wolf was found at the bear bait, consisting mostly of fish, and no signal was heard there. A signal was picked up in the southwest corner of the Mulligan Plains, and the wolf was flushed from her bed about 80 yards (75 m) away.

On October 10, this wolf began a westward move, and on October 22 she was found south of Herman, 25 miles (40.2 km) west of Silver Lake. On October 24 she was located 6.5 miles (10.4 km) to the northeast, near Dirkman Lake. By October 26 she had moved 12 miles (19.3 km) southeast to within a mile of the town of Michigamme. From there she gradually moved northeastward. She was shot near Van Riper Lake during deer hunting season, probably on the morning of November 16.

During the westward move, this wolf had increased the size of her range by 87 square miles (222.7 km^2), about a 30% increase. She seemed to be heading back to the Silver Lake area when she was killed.

Feeding Habits

What little information we could obtain on the wolves' feeding habits indicated considerable variation (Table 4).

In the Skanee area, which the pack of three first visited after leaving the release area, deer were abundant, and 7 to 10 were seen within a quarter mile (0.4 km) of the pack on March 20. It is possible that the wolves killed a deer there, for they remained in the area for a few days. They did scavenge deer feet and head remains on the 22nd at Laws Lake, 12 miles (19.3 km) southwest of Skanee. Deer were also sighted within a quarter mile of the wolves on March 25, April 15, April 16, May 7, June 8, and June 14.

The first confirmed deer kill was made east of Otter Lake about April 1. The deer was a 4-1/2-year-old doe with a partly healed broken left front leg (radius) and fat-depleted bone marrow (1%); a bullet was found in the skin of the right front leg.

The pack also fed on a discarded deer carcass near Nisula, and then killed a 5-1/2-year-old doe near Kenton on April 15 (Fig. 21); this animal also had bone marrow with a low fat content (6%).

The next day, lone Wolf No. 10, back in the Huron Mountain area, killed a 4-5-year-old doe with fat-depleted marrow (5.6%).

No doubt not all of the deer killed or fed upon by the translocated wolves were found, even when snow was present. However, it is clear from the observations we did make, and from the fact that all 26 scats we analyzed from this pack contained deer hair, that the wolves did adapt to killing deer in their new environment and that it was their primary food.

Near Atlantic Mine the wolves scavenged on garbage from loggers, and then near Otter Lake they spent several days also feeding on garbage. A discarded cow (_Bos taurus_) head was scavenged, and at least one red-backed vole (_Clethrionomys gapperi_) was consumed. Lone Wolf No. 10 was found near the Big Bay dump nine times, or 29% of the times she was located during tourist season (May through August).

_Table 4. Analysis of scats collected from released wolves_

No. Wolf Date Scats No. Location and items found --------------------------------------------------------------------- March 22 5 Pack Laws Lake, deer hair

March 29 1 Pack Otter Lake area, deer hair, red-backed vole hair, grass, refuse (including coffee grounds)

April 3 2 Pack Otter Lake deer kill, scats soft and dark, some deer hair

April 8 3 Pack Nisula, deer hair

April 17 5 Pack Kenton deer kill, scats soft and dark, deer hair

June 28 3 Pack Gibbs city area, summer and winter deer hair --------------------------------------------------------------------- Total (Pack) 19 --------------------------------------------------------------------- March 27 2 No. 10 Conway Lake, deer hair

April 18 2 No. 10 Pine Lake, deer hair