Chapter I: Part 1

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FOREWORD

The Fish and Wildlife Service is proud to present this bulletin describing an experimental attempt to re-establish an endangered species in part of its native range. Two States, a Federal agency, a university, and two private conservation groups pooled their resources to make the project possible. This effort exemplifies the type of cooperation the Department of the Interior believes is imperative in beginning the gigantic task of trying to save and restore the threatened and endangered animals in this country today.

Our pride is bittersweet, however. The experiment was a complete success in providing the information sought: What might happen when a pack of wolves is transplanted to a new area where the native population has been all but exterminated by Man? It was the answer to this question that was disappointing. Nevertheless, experiments are for learning, no matter what the answers may be. We are convinced that the answers provided by this project will ultimately be most helpful in future attempts to restore endangered animals to parts of their native ranges where they can begin again on the road to recovery.

DIRECTOR

U. S. Fish and Wildlife Service

Additional Copies Available from

UNITED STATES DEPARTMENT OF THE INTERIOR

FISH AND WILDLIFE SERVICE

REGION 3

Federal Building

Fort Snelling

Twin Cities, Minnesota 55111

AN EXPERIMENTAL TRANSLOCATION OF
THE EASTERN TIMBER WOLF

THOMAS F. WEISE
Department of Biology
Northern Michigan University[1]

WILLIAM L. ROBINSON
Department of Biology
Northern Michigan University

RICHARD A. HOOK
Department of Biology
Northern Michigan University

L. DAVID MECH
Endangered Wildlife Research Program
U. S. Fish & Wildlife Service[2]

[1] _Marquette, Michigan 49855_

[2] _Division of Cooperative Research, Patuxent Wildlife Research Center, Laurel, Md. 20810. Mailing address: North Central Forest Experiment Station, Folwell Ave., St. Paul, MN. 55101._

TABLE OF CONTENTS

FOREWORD Inside front cover

ABSTRACT Back side of title page

INTRODUCTION 1

THE STUDY AREA 2

METHODS 4

RESULTS 8
Social Structure of the Translocated Wolves 8
Aerial Tracking 10
Movements of the Translocated Wolves 11
Post-Release Phase 11
Directional Movement Phase 11
Exploratory Phase 11
Settled Phase 11
Movements of the Remaining Pack Member 11
Movements of Wolf No. 10 12
Feeding Habits 16
Citizen Sightings 17
Habitat Use 19
Failure of Female No. 11 to Whelp 19
Demise of the Translocated Wolves 19

DISCUSSION 21
Effect of Captivity and Human Contact 21
Movements 22
Environmental Influences 22
Possible Homing Tendencies 22
Distances Traveled 23
Home Range Size 25
Selection of a Territory 25
Vulnerability and Mortality 25
Food Habits and Predation 26
An Alternate Approach 26

CONCLUSIONS 26

ACKNOWLEDGMENTS 27

LITERATURE CITED 27

ABSTRACT

Two male and two female eastern timber wolves (_Canis lupus lycaon_), live-trapped in Minnesota were released in March 1974 near Huron Mountain in the Upper Peninsula of Michigan. Their movements were monitored by aerial radio-telemetry.

The wolves separated into a group of three and a single animal after release. The single, a young female, remained in the release region in an area of 346 square miles (896 km^{2}). The pack of three moved generally westward for 13 days and then explored a 1,631 square-mile (4,224 km^{2}) region but settled after 2 months in a 246 square-mile (637 km^{2}) area about 55 miles (88 km) southwest of the release site. The adult female, which mated while captive prior to release, failed to whelp.

In early July, one male was killed by an automobile, and the other was shot. The remaining female from the pack then began to move over a much larger area again. On September 20th she was trapped by a coyote (_Canis latrans_) trapper and shot. Two months later the single female was killed by a deer (_Odocoileus virginianus_) hunter.

These results indicated that wolves can be transplanted to a new region, although they may not settle in the release area itself. The displacement of the translocated wolves in this experiment apparently caused an initial increase in their daily movements, and probably increased their vulnerability, at least during the first 2 months after release. The two females examined post-mortem were in good physical condition indicating that food supplies were adequate in Michigan.

Human-caused mortality was responsible for the failure of the wolves to establish themselves. Therefore recommendations for a more successful re-establishment effort include a stronger public-education campaign, removal of the coyote bounty, and release of a greater number of wolves.

INTRODUCTION

The eastern timber wolf (_Canis lupus lycaon_) originally occurred throughout the eastern United States and Canada but is now extinct in most of the United States. The only substantial population left inhabits northern Minnesota (Fig. 1). The estimated wolf population in the Superior National Forest of northeastern Minnesota in winter 1972-73 was about 390 (Mech 1973), and a tentative population estimate for the entire state is 500 to 1,000 (Mech and Rausch 1975). A well known population of about 15 to 30 wolves is also found in Isle Royale National Park, Lake Superior, Michigan (Mech 1966; Wolfe and Allen 1973; Peterson 1974).

In the Upper Peninsula of Michigan, Hendrickson et al. (1975) estimated the wolf population in 1973 at 6 to 10 animals, existing in three scattered areas: Iron County, Northern Marquette County, and Chippewa and Mackinac Counties (Fig. 2). Lone wolves made up 90 per cent of verified wolf observations there in recent years, and no more than two animals have been found together in at least the past 13 years.

Hendrickson et al. (1975) postulated that the current low wolf population is maintained through possible sporadic breeding and immigration from Ontario and Minnesota (via Wisconsin), but is suppressed by illegal shooting and losses incidental to coyote (_Canis latrans_) bounty trapping.

The eastern timber wolf was classed as an endangered species in the conterminous United States in 1967 under the Endangered Species Act of 1966. There then followed widespread national and international concern and support for preserving natural wolf populations. Substantial scientific and ethical arguments exist for preventing the extinction of a species or subspecies of any plant or animal. In addition, the presence of the wolf adds immeasurably to a wilderness experience; its esthetic value is incalculable.

Thus in 1970, D. W. Douglass, Chief of the Wildlife Division, Michigan Department of Natural Resources, suggested that restoration of a viable population of wolves in Michigan would be desirable, especially if such efforts could be supported by private organizations. In 1973 the Huron Mountain Wildlife Foundation and the National Audubon Society offered financial support, and we undertook this pilot project to obtain information necessary for a full-scale restoration effort.

The objectives of the research project were to determine whether (1) wild wolves could be moved to a new location, (2) such translocated wolves could remain in the new area, (3) they could learn to find and procure enough food in the new area, (4) they could tolerate and survive human activities, and (5) they would breed and help to re-establish a new population in Upper Michigan.

As background we had the results of three previous attempts to transplant wolves to new areas. In 1952, one male and three female zoo wolves were released on Isle Royale (Mech 1966). They were attracted to humans, became nuisances, and had to be disposed of. Two were shot, one was captured and returned to the mainland, and the male escaped; his fate is unknown.

The second transplant effort took place on uninhabited, 36-square-mile (92 km^{2}) Coronation Island in southeastern Alaska (Merriam 1964; Mech 1970). In 1960, two male and two female, 19-month-old captive wolves, were released there. They learned to prey on black-tailed deer (_Odocoileus hemionus columbianus_), and multiplied to about 11 members by 1964.

In the third case, two male and three female laboratory wolves from Barrow, Alaska were released near Umiat in August 1972, 175 miles (282 km) southeast of Barrow (Henshaw and Stephenson 1974). Eventually, all moved toward centers of human habitation and three were shot within 7 months. A fourth returned to the pens where she was reared, and was recaptured, while the fate of the fifth wolf remains unknown. Three of the five had taken the correct homing direction.

Because results of the earlier attempts at translocating wolves suggested that pen-reared wolves did not fare well in the wild, we decided to use wild wolves that were accustomed to fending for themselves and avoiding people. They would have to be released in the most inaccessible area we could find and encouraged to stay there. To maximize their chances of breeding, we would have to try to obtain animals with already established social ties, that is, members of the same pack. Approval was obtained from the Minnesota Department of Natural Resources to live-trap up to five wolves in Minnesota, and a permit was granted by the Michigan Department of Natural Resources to release up to five in Upper Michigan.

This bulletin describes the results of the experimental translocation.

THE STUDY AREA

The area selected for the release of the translocated wolves was the Huron Mountain area (Fig. 2) in northern Marquette County in the Upper Peninsula of Michigan (47 deg. N Latitude; 88 deg. W Longitude). This is one of the largest roadless tracts in Michigan, and has one of the lowest year-around densities of resident humans. Much of the area is owned by the Huron Mountain Club, on which accessibility is restricted.

The Upper Peninsula is 16,491 square miles (42,693 km^{2}) in area, bounded by Lake Superior to the north, and by Lakes Huron and Michigan to the east and south. The Wisconsin border along the western portion of the Upper Peninsula forms no distinctive ecological boundary. The Upper Peninsula is in the Canadian biotic province (Dice 1952), characterized by a northern hardwoods climax, interspersed with spruce-fir and pine subclimaxes. The northwestern portion of the Upper Peninsula, including Marquette, Baraga, Houghton, Ontonagon, and Iron Counties, contains rugged highlands and rock outcroppings which rise to elevations approaching 2,000 feet (610 m) in several locations.

The human population of the Upper Peninsula is 303,342, with a rural density of about 9.0 persons per square mile or 3.5 persons per square kilometer (Table 1). The population of the Upper Peninsula has remained at about 300,000 for the past 50 years, and the rural human populations of local areas have generally declined or remained stable. During those 50 years, the wolf population has declined from several hundred animals to near extinction, with the population estimated by Hendrickson et al. (1975) at 6 to 10 remaining wolves. These authors concluded that the bounty on wolves between 1935 and 1960 was largely responsible for the demise of the species in the Upper Peninsula. The bounty was removed in 1960, after only one wolf was taken in 1959. Legal protection was granted by Michigan in 1965. The Endangered Species Act of 1973 added federal protection in 1974.

_Table 1. Density of Rural Human Populations in Four Wolf Ranges in the Great Lakes Region_

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Rural[4]
Area in Population
Square Miles Density Per
(Square Percent Rural Square Mile
Location Kilometers) Urban[3] Population (Square Kilometer)
----------------------------------------------------------------------
Ontario[5] 412,582 3.3
(1,068,125) 80.4 1,364,33 (1.27)

Northern[6] 12,627 6.4
Minnesota (32,690) 68.0 81,246 (2.5)

Upper 16,491 9.0
Michigan[7] (42,693) 51.4 147,841 (3.5)

Iron and Oneida
Co.[8] 1,859 12.3
Wisconsin (4,812) 26.0 22,899 (4.7)
----------------------------------------------------------------------

[3] Towns or cities of more than 2,500 people

[4] Including towns with a population less than 2,500

[5] 1966 Census, 1970-71 Canada Yearbook

[6] Cook, Koochiching, Lake and St. Louis Counties

[7] All 15 Upper Peninsula counties

[8] Last described wolf range in Wisconsin (Thompson 1952)

The white-tailed deer (_Odocoileus virginianus_) would be the major prey for wolves in Michigan, and there appear to be sufficient numbers to support wolves. The Michigan Department of Natural Resources pellet count estimates for the spring deer population in the Upper Peninsula in 1973 was 10 +/- 21.9% deer per square mile (3.9 +/- 21.9% per km^{2}). Deer densities of 10 to 15 per square mile (3.9 to 5.8 per km^{2}) supported wolf densities of one wolf per 10 square miles (26 km^{2}) in Algonquin Provincial Park, Ontario (Pimlott 1967).

The population of deer wintering on the 14 square-mile (36 km^{2}) Huron Mountain deer yard in winter 1973-74 was estimated at 73.3 +/- 49.5% deer per square mile (28.3 +/- 49.5% deer per km^{2}) by the pellet count method (Laundre 1975). Thus total wintering population on the Huron Mountain Club, the wolf release area, would be about 1,000 deer.

The utilization of available browse by deer in the Huron Mountain deer yard reached 95% by March 8, 1969 and 92% by March 5, 1970 (Westover 1971). The minimum winter deer loss (actual number found) in 1969 was 40 animals, of which at least 12 had starved, and it was estimated that perhaps up to 33% of the deer starved in the Huron Mountain Yard in 1968-69 (Westover 1971). The Huron Mountain yard continues to be overbrowsed, with high deer mortality expected in severe winters. Many other northern deer yards of the Upper Peninsula are also overbrowsed and are dwindling in area. Thus we expected that numbers of vulnerable deer (Pimlott et al. 1969; Mech and Frenzel 1971) would be available to wolves.

Beavers (_Castor canadensis_) are an important food source for wolves in many areas during summer (Mech 1970), and they are common throughout the Upper Peninsula. The beaver population on the 26 square-mile (67 km^{2}) Huron Mountain Club was estimated at 46.9, or about 1.9 beavers per square mile (0.7 per km^{2}) (Laundre 1975). Moose (_Alces alces_) are rare on mainland Michigan.

METHODS

The general procedure for this study was to attempt to capture an intact pack of wolves in Minnesota, fit each animal with a radio-collar (Cochran & Lord 1963), release them in northern Michigan, and follow their fate through aerial and ground radio-tracking (Kolenosky and Johnston 1967).

A pack was selected from an area near Ray, Minnesota (Fig. 3), south of International Falls (48 deg. N Latitude, 93 deg. W Longitude), where wolf hunting and trapping were legal. Two male and two female wolves were captured by professional trapper Robert Himes, under contract for the project, between December 24, 1973 and January 21, 1974 (Table 2). Three of the wolves were trapped (Fig. 4) in No. 4 or 14 steel traps (Mech 1974), and one (No. 13) was live-snared (Nellis 1968). If these animals had not been solicited for this study, they would have been killed and their pelts sold, as part of the trapper's livelihood, before the Endangered Species Act of 1973 took effect.

At capture each wolf was immobilized with a combination of phencyclidine hydrochloride (Sernylan) and promazine hydrochloride (Sparine) intramuscularly (Mech 1974), with dosage recommendations from Seal et al. (1970). They were then carried out of the woods (Fig. 5), held in pens in Minnesota, and fed road-killed white-tailed deer, supplemented with beef scraps.

There is no certain way of ascertaining that wolves are related or that they belong to the same pack. Thus to maximize chances that members of the same pack would be captured, the trapper set traps where he suspected only one pack ranged. To try to determine whether the individual wolves he caught were socially related, we instructed the trapper to hold the wolves in individual pens until we could observe their introductions to each other. Wolves No. 10 and 11 were placed together on January 23, 1974, and No. 13 and 14 were released into the pen with No. 10 and 11 on February 4.

_Table 2. Background information on the translocated wolves_

----------------------------------------------------------------------
Wolf Number 10 11 12 13
----------------------------------------------------------------------
Sex F F M M

Estimated age[9] 1-2 years 6-7 years 2-3 years 2-3 years

Capture date 12-24-73 1-5-74 1-19-74 1-21-74

Capture Method Trapped Trapped Trapped Live-snared

Capture foot Left front Right front Right front

Capture-related Two nails Three nails None None
damage lost lost

Weight at 55 lb. 65 lb. 74 lb. 75 lb.
capture (24.9 kg) (29.4 kg) (33.5 kg) (33.9 kg)

Weight, March 5 46 lb. 58 lb. 66 lb. 60 lb.
(20.8 kg.) (26.3 kg) (29.9 kg) (27.2 kg)

% weight loss 16% 11% 11% 20%

Canine length, 0.83" 0.25-0.50" 0.93" 0.87"
upper (21 mm) (6-13 mm) (24 mm) (22 mm)

Canine length, 0.75" very worn 0.82" 0.85"
lower (19 mm) (21 mm) (21 mm)

Testes[10] ---- ---- 0.5 x 1.0" 0.5 x 0.75"
(13x25 mm) (13x19 mm)

Teats Tiny, not Dark, ---- ----
apparent evident
----------------------------------------------------------------------

[9] Gross subjective estimates based on tooth wear

[10] Estimated

On March 5, 1974, the wolves were again immobilized for pre-release processing in Minnesota. An initial dose, and several supplemental doses of phencyclidine and promazine were administered intramuscularly and intraperitoneally between 9:00 a.m. and 4:30 p.m. CDT. The wolves were restrained with muzzles and their legs were bound together during processing and transport. Two of the wolves were blindfolded because they were too active otherwise.

The wolves were ear-tagged with both Minnesota and Michigan Department of Natural Resources tags, and weights and body measurements were taken (Fig. 6, 7). Their teeth were inspected and canines were measured to try to obtain an indication of age. Each animal was fitted with a radio transmitter (AVM Instrument Co., Champaign, Illinois[11]) molded into an acrylic collar (Mech, 1974).

Each wolf was injected with 1,200,000 units of Bicillin (Wyeth), 2 cc of distemper-hepatitus-leptospirosis vaccine (BioCeutic Laboratories D-Vac HL), 0.5 cc of vitamins A, D, E, (Hoffman-LaRoche[11] Injacom 100), 1 cc of vitamin C-fortified vitamin B complex (Eli-Lilly, Betalin Complex FC), and 2 cc anti rabies vaccine (Fromms Raboid). These injections (Fig. 8) were given to insure that the wolves would be as healthy as possible upon release, and would not contract or introduce diseases in the release area.

[11] _Mention of trade names does not constitute endorsement by the U. S. Government._

Some 30 to 60 cc of blood were drawn from each wolf for analysis of its physical condition (Seal et al. 1975).

The processing of the wolves took from 8:45 a.m. to 2:00 p.m. CDT on March 5, 1974. The animals were then transported by truck to International Falls, loaded on an airplane (Fig. 9), and flown for 2 hours (Fig. 10) to the Marquette County Airport, Michigan. They were turned on different sides each half hour while drugged during their processing and transport to prevent lung congestion. At the Marquette Airport they were transferred by van to a 25 foot by 25 foot by 12 foot (7.6 m x 7.6 m x 3.7 m) holding pen on the Huron Mountain Club property 35 miles (56.3 km) northwest of Marquette.

The wolves were released individually into the holding pen while each was still partly under sedation (Fig. 11). The transmitting frequency of each wolf's collar was rechecked on the receiver as each wolf was released into the pen (Fig. 12). All wolves were in the pen by 10:00 p.m. EDT, and were held there until March 12.

Four road-killed deer carcasses, provided by the Michigan Department of Natural Resources, had been placed inside the pen for food (Fig. 13), and a tub of drinking water was provided. Carcasses of five road-killed deer and a black bear (_Ursus americanus_) were placed within a half-mile (0.8 km) of the release pen as food for the wolves after their release.

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

June 1 1 No. 10 Huron Mountain Club, fawn deer hair and hoof
---------------------------------------------------------------------
Total No. 10 5
---------------------------------------------------------------------
Sept. 20 1 No. 11 Floodwood Plains 3.1 miles (5.0 km) south of
Witch Lake, deer hair, and ruffed grouse
(_Bonasa umbellus_) bones and nails

July 1 1 No. 12 Collected from under dead No. 12, 1.9 miles
(3.0 km) north of Amasa, deer hair
---------------------------------------------------------------------
Total 26 All

The three wolves were located near beaver lodges or dams on April 10, April 15, May 7, June 8, and June 12. No beavers were known to have been killed by them, however, and no beaver remains were found in their scats (Table 4).

Citizen Sightings

The wolves were seen by many citizens early after their release (Table 5 and 6), no doubt because of the wolves' confusion, their extensive movements, and their lack of familiarity with the region. They often traveled near populated areas and probably moved more during the day than they would have in their native territory. They were known to have made 14 daytime moves (from citizen reports) in addition to those observed from the aircraft. In at least five of the citizen reports, the wolves were observed sitting alongside the road, or otherwise making little attempt to move away immediately. However, after April 13 the group of three wolves was reported by citizens only twice, and Wolf No. 10, three times.

_Table 5. Significant events in history of Wolf No. 10_

Date Event

March 12 Wolves released in Huron Mountain area
(T52N-R28W-Sec 20)

March 13 No. 10 separated from the other three wolves and
never reunited

March 15 Sighted from tracking car crossing County Road 492
south of Marquette County Airport, 6:35 p.m. (EDT)
(T47N-R26W-Sec 33)

March 15 Crossed a four-lane highway between Marquette and
Negaunee about 4:00 p.m. (EDT) (T49N-R26W-Sec 29)

March 24 Located from the air less than 0.5 miles (0.8 km)
from release pen (T52N-R28W-Sec 20)

March 27 Reported seen by Huron Mountain Club guard on edge
of First Pine Lake, 6:30 p.m. (EDT)
(T52N-R28W-Sec 29)

April 18 Visited bear carcass 100 feet (30.5 m) from
release pen, had also visited 3 nearby deer
carcasses (T52N-R28N-Sec 20)

April 18 Confirmed deer kill by No. 10 near Pine Lake,
Huron Mountain Club (T52N-R28W-Sec 20)

June 6 Reported seen by gate guard, Huron Mountain Club
(T51N-R27W-Sec 6)

June 3 Reported seen north of Saux Head Lake on Lake
Superior beach (T50N-R26W-Sec 17)

June 20 Reported seen crossing four-lane highway headed
north about 5 miles (8.0 km) west of Marquette
(T50N-R26W-Sec 24)

May 22 }
May 23 }
June 5 } Located near Big Bay dump, probably scavenging.
July 15} Bears are baited at the dump by local citizens and
July 20} tourists (T51N-R27W-Sec 16)
July 31}
Aug. 6 }
Aug. 13}

Aug. 16 Back in Huron Mountain area between Conway and
Ives Lakes. 5:35 p.m. (EDT) (T52N-R28W-Sec 35)

Aug. 27 Returned to Big Bay dump, 11:10 a.m. (EDT)
(T51N-R27W-Sec 16)

Aug. 30 Huron Mountain area, 8:45 a.m. (EDT)
(T49N-R28W-Sec 9)

Sept. 2 Left Huron Mountain area for last time. Located on
Yellow Dog Plains, 8:45 a.m. (EDT)
(T50N-R28W-Sec 13)

Sept. 5 Near Silver Lake, 8:45 a.m. (EDT). Begins rambling
move westward out of established range
(T49N-R28W-Sec 17)

Sept. 15 Tracked on ground on Mulligan Plains, 4:45 p.m.
(EDT) (T49N-R28W-Sec 9)

Oct. 22 Farthest west, 22 miles (35.4 km) west of Silver
Lake. Begins rambling return east.

Nov. 16 Killed 1/2 mile (0.8 km) south of Van Riper Lake,
5.4 miles (8.4 km) north of Champion
(T49N-R30W-Sec 36)

_Table 6. Significant events in history of Wolves
No. 11, 12 and 13_

Date Event

March 12 Wolves released in Huron Mountain area
(T52N-R28W-Sec 20)

March 18 Two wolves reported seen near Ravine River, Skanee
area, the smaller one limping (T51N-R31W-Sec 2)

March 19 First aerial fix of the three wolves in the same
location (T52N-R31W-Sec 36)

March 20 Wolves reported howling about 2 miles (3.2 km)
east of Arvon Tower, 10 miles (16 km) south of
Skanee (T50N-R31W-Sec 4)

March 22 Wolves dug up five discarded doe and
fawn heads and 27 deer legs near Laws
Lake (T50N-R32W-Sec 18)

March 22 Wolves reported crossing highway north of Herman,
4 miles (6.4 km) southeast of L'Anse, 8:30 a.m.
(EDT) (T50N-R33W)

March 25 Wolves reported in Pelkie area 6 miles (9.6 km)
east of Baraga by DNR officer, 8:30 a.m. (EDT)
(T51N-R34W-Sec 27SW)

March 25 Wolves crossed road 2.5 miles (4 km) north of
Pelkie near Otter River 11:00 a.m. (EDT) 5 miles
(8 km) southwest of Otter Lake (T51N-R34W-Sec 5)

March 25 Wolves reported seen crossing Highway M26, 2 miles
(3.2 km) north of Twin Lakes 7:30 a.m. (EDT)
(T52N-R38W-Sec 12)

March 26 Wolves reported seen by logger during most of
morning 9:00-11:00 a.m. (EDT), 4 miles (6.4 km)
south of Houghton, (T54N-R35W-Sec 14)

March 26 Wolves crossed Highway M26 south of Atlantic, 4:30
p.m. (EDT), (T54N-R34W-Sec 16)

March 26 Wolves sighted from aircraft, eating garbage from
cutting crew, 4:20 p.m. (EDT) (T54N-R34W-Sec 9NE)

March 29 Wolves reported being chased away from house by
dog, had been feeding on discarded cow head 150
feet (45.7 m) from house near Otter Lake
(T52N-R33W-Sec 5)

March 31 Wolves sighted in Otter Lake area (T52N-R33W-Sec 5)

April 2 First confirmed wolf-killed deer, Arnheim area
about 10 miles (16 km) north of Baraga
(T52N-R33W-Sec 11)

April 5 Wolves reported seen at 9:00 a.m. (EDT) on county
road 5 miles (8 km) southwest of Otter Lake, small
wolf reported as appearing fat (T53N-R35W-Sec 36)

April 8 Wolves dug up old deer carcass about 150
feet (45.7 m) from house near Nisula
(T50N-R36W-Sec 4)

April 10 Wolves reported seen by logger in Nisula area
(T50N-R36W-Sec 5)

April 13 One wolf sighted crossing Highway M28 in morning
between Kenton and Sidnaw

April 15 Wolves killed deer near Kenton (T47N-R36W-Sec 8)

April 18 Observed the three wolves from the tracking
aircraft swim the East Branch of Ontonagon River,
southeast of Kenton (T47N-R37W-Sec 7)

May 2 No. 13 split from other two wolves; found in
northwest Ontonagon County (T51N-R32W-Sec 21)

May 7 All wolves back in Iron County for the second
time, not known to leave until July 15

May 7 Forest service crew reported seeing the wolves and
tracking aircraft north of Gibbs City near old
deer carcass (T45N-R35W-Sec 26)

May 15 Loggers reported six wolves (one with collar)
(T54N-R37W-Sec 33)--Probably saw the collared
wolves twice

May 16 Confirmation from aerial location that the three
wolves had reunited south of Mallard Lake after
May 2 split

June 19 No. 13 again separated from No. 11 and 12

July 11 Wolf No. 12 found dead, killed by automobile just
before July 6, north of Amasa (T45N-R33W-Sec 17)

July 15 Wolf No. 11 moved out of Iron County for the first
time since May 7, found north of Kenton
(T49N-R38W-Sec 31)

July 20 Wolf No. 13 found dead from gunshot, south of
Sagola, last previous location (June 27) at same
location where No. 12 killed by automobile
(T52N-R30W-Sec 5)

Aug. 6 Wolf No. 11 located near Wisconsin border, 3/4
miles (1.2 km) east of Lac Vieux Desert, 10:15
a.m. (EDT) (T43N-R38W-Sec 9)

Aug. 13 Wolf No. 11 located 1.5 miles (2.4 km) southeast
of Ewen 25 miles (40.5 km) north of Lac Vieux
Desert, 10:10 a.m. (EDT) (T46N-R40W-Sec 36)

Aug. 28 No locations since Aug. 13. Wolf No. 11 back in
Marquette County .25 miles (0.4 km) south of Squaw
Lake, a 60-mile (96.5 km) move eastward
(T45N-R30W-Sec 21)

Sept. 20 No. 11 trapped and shot on Floodwood Plaine 3.1
miles (5.0 km) south of Witch Lake
(T44N-R24W-Sec 11)

Habitat Use

The relative percentages of various habitats in which the translocated wolves were found during aerial locations (Table 7) did not indicate a preference for any particular habitat type. Evidently the animals chose their travel routes and ranges on some basis other than forest habitat, or at least habitat was not of any overriding importance in their movements.

_Table 7. Habitat types in which the released wolves were located_

--------------------------------------------------------
No. of Percent Percent
Habitat Locations of Total Available[12]
--------------------------------------------------------
Northern Hardwoods 43 48.3 40.9

Northern Hardwoods-
Coniferous[13] (57) ...[13] ...[13]

Spruce-fir 19 21.3 17.0

Aspen-hardwoods 11 12.4 20.5

Elm-ash-maple 1 1.1 4.5

Pine 2 2.2 5.5

Oak 0 0.0 1.4

Non-commercial
forests 0 0.0 2.6

--------------------------------------------------------
Other (near towns,
farms, dumps) 13 14.6(8.9)[13] 7.6
__ ______ _____

Totals 89(146) 100.00 100.0
--------------------------------------------------------

[12] Spencer and Pfeifer 1966.

[13] This forest type was not distinguished separately by Spencer and Pfeifer (1966), so they did not provide availability figures for it. Thus in this comparison, we did not include the 57 wolf locations that fell in the type. However in calculating percentage figures for non-forest areas (towns, farms, dumps), these 57 fixes could validly be used as representing forest locations.

Failure of Female No. 11 to Whelp

There was no sign that adult female No. 11 whelped or attempted to locate or construct a den. The usual gestation period for wolves is about 63 days (Brown 1936). Because No. 11 was seen coupled in copulation on February 12 and 16, she should have whelped between April 13 and April 21, if she had conceived. Probably she would have moved little during the preceding 2 or 3 weeks (Mech 1970). However no such changes in this animal's movements were noticed. The three wolves stayed near Kenton between April 15 and April 18 but also killed a deer during that time. They moved extensively from April 19 to May 7. The only indirect evidence that the female may have been pregnant was an observation made by a local citizen on April 5 (Table 6) who saw the three wolves and stated that the small wolf looked "fat." This would probably have been No. 11, but a full stomach could easily have been mistaken for pregnancy.

Unfortunately, neither the reproductive tract collected from No. 11 in September nor the blood sample taken in early March shed any light on the cause for the wolf's failure to produce pups. The ovaries did contain _corpora albicantia_, indicating that at some time the wolf had ovulated, but it could not be stated with certainty just when (R. D. Barnes, personal communication). The blood progesterone levels were more helpful. No. 11 had 3,560 picograms of progesterone per milliliter, compared to 56 picograms per milliliter for Wolf No. 10, whose reproductive tract appeared immature. This high progesterone level of No. 11 indicated that the animal had recently ovulated, but it was impossible to tell whether she was carrying any fetuses at the time the sample was taken (U. S. Seal, personal communication).

Demise of the Translocated Wolves

All four translocated wolves were killed by humans (Table 8). The alpha male (No. 12) was the first victim. He was found from the air in the same location on July 6 and 10. A ground check on July 11 showed him already decomposed. He lay about 60 feet (18.3 m) from paved highway US 141 north of Amasa (Fig. 22). The articular processes on the right side of his fifth and sixth cervical vertebrae were broken and inverted. Part of the process of the sixth cervical vertebra was lodged in the neural canal between the fifth and sixth cervical vertebrae and would have exerted pressure on his spinal cord. His acrylic radio collar was also cracked on the right side in three places. We concluded that he had been struck and killed by an automobile. A scat found beneath the remains contained deer hair, so apparently the animal had been feeding not long before his death.