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

AN EXPERIMENTAL TRANSLOCATION OF

THE EASTERN TIMBER WOLF

TABLE OF CONTENTS

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 km²). The pack of three moved
generally westward for 13 days and then explored a
1,631 square-mile (4,224 km²) region but settled
after 2 months in a 246 square-mile (637 km²)
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.

[Pg 1]

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).

Fig. 1—Original and present range of the Easter Timber Wolf

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 km²) 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.

[Pg 2]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° N Latitude; 88° 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 km²) 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.

[Pg 3]

Table 1. Density of Rural Human Populations in Four Wolf Ranges
in the Great Lakes Region

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 km²).
Deer densities of 10 to 15 per square mile (3.9 to 5.8
per km²) supported wolf densities of one wolf per 10
square miles (26 km²) in Algonquin Provincial Park,
Ontario (Pimlott 1967).

The population of deer wintering on the 14 square-mile
(36 km²) 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 km²) 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 km²) Huron Mountain Club was
estimated at 46.9, or about 1.9 beavers per square
mile (0.7 per km²) (Laundre 1975). Moose (Alces
alces) are rare on mainland Michigan.

[Pg 4]

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° N
Latitude, 93° 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.

Fig. 4.—Wolf caught in trap (Photo by Don Breneman)

Fig. 5.—The captured wolves were drugged and
carried to an enclosure in Minnesota (USFWS
Photo by L. David Mech)

[Pg 5]

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.

Fig. 6.—Before being transported to Michigan, each
wolf was weighed (USFWS Photo by Don Reilly)

Table 2. Background information on the translocated wolves

[Pg 6]

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).

Fig. 7.—Standard body measurements were also
taken (USFWS Photo by Don Reilly)

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.

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 × 7.6 m × 3.7 m) holding
pen on the Huron Mountain Club property 35 miles
(56.3 km) northwest of Marquette.

Fig. 8.—Various vitamins and vaccines were administered
to each wolf to insure their health and freedom
from common canine diseases (USFWS Photo
by Don Reilly)

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.[Pg 7]

Fig. 9.—The anesthetized wolves were placed aboard
an aircraft in International Falls, Minnesota
(USFWS Photo by Don Reilly)

Fig. 10.—The wolves were kept lightly drugged during
the flight to Michigan (USFWS Photo by L.
David Mech)

Fig. 11.—In the Huron Mountain area of Upper
Michigan the wolves were taken to another holding
pen (Photo by Don Pavloski)

Fig. 12.—Biologists checked the signal from each
radio-collar before the wolves were released into
the holding pen (Photo by Don Pavloski)

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.[Pg 8]

Fig. 13.—While in captivity, the wolves were fed primarily on road-killed deer (Photo by Don Pavloski)

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).

[Pg 9]

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.

Fig. 14.—The Minnesota wolves in their Michigan pen (Photo by Tom Weise)

[Pg 10]

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

Fig. 15.—The wolves often used woods roads for
traveling (Photo by James Havemen)

Fig. 16.—The released wolves were sometimes observed
from the aircraft feeding on deer they had
killed (Photo by Richard P. Smith)

[Pg 11]

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 km²) 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 km²) 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 1½ 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[Pg 12]
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 1¼ 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.[Pg 13]

[Pg 14]

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[Pg 15]
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 km²),
about a 30% increase. She seemed to be heading back
to the Silver Lake area when she was killed.

[Pg 16]

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 4½-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 5½-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

[Pg 17]

Fig. 21.—Each deer killed by the translocated wolves
was examined from the ground (Photo by Richard
P. Smith)

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

[Pg 18]

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

[Pg 19]

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

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[Pg 20]
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.

Fig. 22.—The remains of Wolf No. 12 were found
near a highway, and broken bones indicated he had
been hit by a vehicle (Photo by Richard P. Smith)

Wolf No. 13 was killed next. He had been located
south of Sagola in Dickinson County on July 20, the
first time he was found since June 27. He was still
there on July 27, so a ground check was made. It
revealed that the wolf had been dead for perhaps 2
or 3 weeks. His flesh had decomposed, and only hair,
bones and the transmitting collar remained (Fig. 23).
His leg bones and ribs were mostly disarticulated, his
skull was separated from the vertebral column, and
his mandible had separated. A small caliber bullet
had passed through the ramus of the left mandible
and had entered the base of the cranium. The hole
through the mandible was 0.26 inch × 0.34 inch (6.6
mm. × 8.6 mm.) and that through the cranium was
0.34 inch × 1.30 inch (8.6 mm. × 33.0 mm.). Three
small lead fragments were removed from the cranium.

Fig. 23.—Wolf No. 13 had been shot, as the hole in
the jawbone indicates (Photo by Tom Weise)

The remains of Wolf No. 13 were sent to the
Michigan Department of Natural Resources Wildlife
Research Center at Rose Lake and examined by staff
pathologists Dr. L. D. Fay and Mr. John Stuht. No
fractures or other signs were found that might indicate
that he had been trapped. However, some of the
smaller foot bones were missing and a complete examination
was not possible. Notches were found in both
shoulder blades, and one rib was broken, suggesting
that the animal had been shot twice by a small caliber
firearm in addition to the head shot. The hole in the
left scapula indicated a deep penetrating wound. The
notch in the right scapula indicated a bullet traveling
more parallel to the body.

Table 8. Details of Deaths of Translocated Wolves

[Pg 21]

Wolf No. 11 was caught the night of September 19,
1974 in a coyote trap set by a trapper from Channing.
The next morning the trapper came upon the trapped
wolf by surprise at a range of 12 feet (3.6 m). She
growled and lunged toward him, and thinking he was
in danger, the trapper shot the wolf in the head. The
.22 caliber bullet entered below the right eye and
lodged in the skull. The trapper immediately took
the animal to the Michigan Department of Natural
Resources office in Crystal Falls and reported the
incident.

The wolf weighed 56.5 lb. (25.6 kg), 1.5 lb. (0.68
kg) less than when she was brought to Michigan. Her
general condition was good, with some omental fat,
but no subcutaneous fat. She did harbor ten tapeworms
(Taenia pisiformis) about 40–50 cm long and
a few hookworms (Uncinaria stenocephala), as determined
by Mr. John Wenstrom (personal communication),
Biology Department, Northern Michigan University.
Both are common tapeworms of wolves (Mech
1970).

Wolf No. 10 was shot by a deer hunter, probably on
the morning of November 16, the second day of firearms
deer season. On November 17 her signal was
heard from near a cabin on the south shore of Van
Riper Lake. The hunters occupying the cabin later
said they had removed the collar from the wolf, which
they had found dead on the afternoon of November
16. Before we had learned this, the carcass of Wolf
No. 10 was discovered without the collar by another
hunter, about a half mile (0.8 km) south of Van
Riper Lake. It had been shot through the right leg,
shattering the radius and ulna, and through the head,
the bullet entering the left frontal bone and exiting
below the right eye. In addition the radio collar had
been shattered by a bullet and was missing, and one
ear had been cut off. We identified the wolf from
the tag in the other ear.

The wolf had gained 6 lb. (2.7 kg) since she had
been brought to Michigan, and had heavy internal
and subcutaneous fat. She had light infections of two
species of tapeworms (Echinococcus granulosus and
Taenia pisiformis), and of one species of hookworm
(Uncinaria stenocephala), as determined by John
Wenstrom. Echinococcus granulosus is not uncommon
in wolves (Mech 1970). The other two species were
discussed above.

DISCUSSION

Wolves No. 11, 12, and 13 undoubtedly were members
of the same pack. This conclusion is based on the
fact that they did not fight when placed together in
captivity, that they freely intermixed while penned,
that No. 11 and No. 12 copulated, and that all three
wolves generally traveled as a unit after their release.
No. 11 and No. 12 were always located together from
a few days after their release until the death of No.
12. Temporary splitting, as with No. 13 is a normal
occurrence in wild wolf packs (Mech 1966).

The identity of Wolf No. 10 remains unknown. She
was captured 7.5 miles (12.1 km) away from the
other three, and in captivity she behaved differently
from them, remaining more to herself but intermingling
with the others occasionally, with no signs of
aggression. The face licking of No. 10 by No. 11 could
be interpreted as a sign of patronizing intimacy as an
adult might treat a subordinate offspring. The teeth
of Wolf No. 10 had very little wear, indicating that
she probably was less than 3-years old, whereas the
teeth of No. 11 were blunt from wear. The tendency
for No. 10 to withdraw from the others and from
human beings indicated that she probably was a low-ranking
or subordinate animal, a peripheral member
of the pack (Woolpy 1968), or even a lone wolf
currently dispersing from the pack (Mech 1973).

The separation of No. 10 from the others upon
release does not necessarily mean that she was not a
member of the pack. No. 10's radio collar was replaced
just before she was released. The handling
without sedation could have frightened her enough
that she ran some distance before the others were
even released. The fact that No. 10 returned to within
a half mile (0.8 km) of the release pen on March
20 and to within less than 100 feet (30.5 m) on April
18 may indicate she was seeking the other wolves.
However, she may also just have used the release pen
as a reference point in a generally unfamiliar area, or
may have been attracted by the remains of carcasses
left there.

Effect of Captivity and Human Contact

The necessary capture, captivity, translocation and
contact of the experimental wolves with humans had
an unknown effect on the wolves. They had been
exposed to humans for over 2 months while in captivity.
No attempts were made to tame them, and
they never passed the escape stage of socialization as
described by Woolpy and Ginsburg (1967). The
dominant wolves (No. 11 and No. 12) were more
relaxed when approached than were No. 10 and No.
13, however.

The failure of female No. 11 to bear young probably
can be attributed to her captivity and handling.
The fact that two couplings were observed over a
5-day period indicates normal estrus in the female,
and a normal response in the male. Conception would[Pg 22]
have been expected from such a mating. In wild
wolves, it is known that there is only a small loss between
number of ova shed, number of embryos implanting,
and number of fetuses being carried (Rausch
1967). Thus it seems unlikely that, if No. 11 conceived,
she lost her fetuses in utero. Rather, she probably
did not conceive, or perhaps the embryos never
implanted. This wolf lost about 11% of her capture
weight during captivity, despite an adequate food
supply. This fact, plus the results of her blood tests
indicate a high degree of stress, which probably explains
why she never produced pups.

The possible interference of the drugs used can be
ruled out, for they were chosen because of their
known lack of effect on pregnancy (Seal et al. 1970).

The radio collars placed on the wolves had no
noticeable effect on the animals. Radioed wolves are
regularly accepted back into their packs in Minnesota,
where they also reproduce and function normally
(Mech and Frenzel 1971; Mech 1973, 1974).

Movements

Environmental Influences

Lake Superior was a barrier to the northward and
eastward movements of the wolves. Apparently it also
directed wolves No. 11, 12, and 13 southward around
Keweenaw Bay, and possibly it prevented their eastward
movement on April 2 when they approached
Keweenaw Bay from the western side. The Bay is
approximately 6-miles (9.6 km) wide there, and was
frozen until late April.

One to two miles (3.2 km) south of the release site,
the Huron Mountains, with an elevation of 1,500
feet (457.5 m) might have prevented the southward
movement of the wolves. Along the lakeshore, the
land is relatively flat, which may have facilitated east-west
movement. Wolves No. 11 and 13 were found at
an elevation of 1,300 feet (490 m) the day after
release but had returned to the flat shore areas (600
to 700 feet, or 200 to 230 meters above sea level) by
the next day. Topography likely had effects in other
areas but the actual travel routes, in most instances,
are unknown. The pack did travel along an abandoned
railroad grade near Gibbs City and for 2 miles
(3.2 km) on a muddy road north of Kenton. Wolf
No. 10 used a railroad bridge to cross a river in mid-March.
It is well known that wolves generally choose
the easiest routes of travel (DeVos 1950, Stenlund
1955, Mech 1966).

Possible Homing Tendencies

Some of the movements of the wolves during the
Directional Movements Phase could in part have
resulted from a tendency for the animals to home,
that is to return to their home territory. Packs have
been observed to travel 45 miles (72 km) in 24 hours
in Minnesota (Stenlund 1955), Alaska (Burkholder
1959) and on Isle Royale (Mech 1966). In Minnesota,
a radioed wolf was tracked a straight-line distance
of 129 miles (208 km) over a 2-month period
before being lost by researchers (Mech and Frenzel
1971), and annual migratory movements of over 200
miles (320 km) have been reported for Canadian
wolves (Kuyt 1972). Therefore it seems within the
capabilities of the released wolves to return the 270-mile
(434 km) straight-line distance, or the 340-mile
(547 km) travel distance around Lake Superior to
Ray, Minnesota, if the orientation ability and inclination
were present.

Homing tendencies have been reported in wolves
and other carnivores. One of five laboratory-reared
wolves returned to her Barrow, Alaska homesite within
about 4 months after a 175-mile (282 km) displacement
(Henshaw and Stephenson 1974). An
adult female red fox (Vulpes vulpes) returned to her
homesite within 12 days after being displaced 35 miles
(56.3 km) (Phillips and Mech 1970). For black bears
there are many records of apparent homing. Harger
(1970) displaced 107 adult black bears from 10.0 to
168.5 miles (16.1 to 270.3 km) with an average displacement
of 62.5 miles (100.6 km). Thirty-seven of
them homed and 11 others moved long distances
toward home. The longest distance homed was 142.5
miles (229.4 km). The return travel routes seemed
direct, with little evidence of wandering or circling.
Harger (1970) concluded that bears could navigate
by some means, as yet undetermined.

There is some indication that the pack of three
wolves may have attempted to return home to Minnesota,
although it is possible that exploration itself
also may have produced the movement pattern
observed.

If the translocated wolves were to try homing
directly toward their previous territory, they would
have had to travel west-northwestward. However,
within a few miles they would have encountered Lake
Superior. The next closest choice would have been to
head westward, and this is what the pack did (Fig.
17). The next possible barrier to their homeward
movements would have been Huron Bay, which
would have forced them southwestward, at least temporarily.
Again this is what actually happened. The
pack maintained its southwestward movement beyond
Huron Bay until reaching a point southeast of the[Pg 23]
next possible barrier, Keweenaw Bay. They then continued
westward south of Keweenaw Bay to the Prickett
Dam area, and veered northwestward to Twin
Lakes on March 25.

By this time, the wolves had traveled for 13 days
and covered a minimum distance of 59 miles (94.9
km), and they were 42 miles (67.6 km), closer to
home (16% of the straight-line distance between
home and release site). The directions of the movements
of the wolves were consistent with what they
would have to be if the wolves were to return home.

However, after March 25, the directionality in the
movements of the pack ended (Fig. 17), and the
animals began what we consider the Exploratory
Phase of their movements. If the wolves actually were
homing, perhaps the tendency diminished as they
failed to encounter familiar terrain, or perhaps they
met too many obstacles, or became confused after
encountering too much human activity. Or possibly
these factors or the need to find food and security
overcame the homing tendency. As discussed earlier
in relation to the unusual number of times the wolves
were observed, it is clear that they were not moving
normally during this period.

The lone wolf, No. 10, dispersed from the release
site in as much of an opposite direction as it could
from the pack (Fig. 20). Thus there is no evidence
that this animal was trying to home. However, it is
of interest to note that the first 32 miles (51.5 km)
of her travel was directional rather than random.
Furthermore, when the animal encountered what
probably was a psychological barrier, a high concentration
of human activity along Highway 41, she
reversed her movements but still maintained a directionality
by returning to the release area. In fact a
striking pattern of southeast-northwest movements
characterized this wolf's travels for several months
after her release, with a gradual westward drift developing
in the southeast-northwest movements (Fig.
20).

Mech and Frenzel (1971) found that a wolf dispersing
from his former home range in Minnesota
maintained a general southwestward movement for
a straight-line distance of 129 miles (207.6 km) over
a 2-month period, and Mech (unpublished) has three
additional records of dispersing wolves that maintained
directionality for distances of 48 to 130 miles
(77.2 to 209.2 km). Storm (1972) followed 12 dispersing
red foxes in Iowa, Illinois, and Minnesota
that moved directionally for distances of 12 to 110
miles (19.2 to 176.0 km).

The ability of wolves to orient and navigate even in
unfamiliar surroundings was demonstrated dramatically
by the separation of Wolf No. 13 from his two
packmates and his later rejoining of them. On May 2
he was 51 miles (82.1 km) away from them. Five
days later he and his packmates were only 6 miles
(9.6 km) apart, in an area 62 miles (99.8 km) from
where No. 13 had been on May 2, and 45 miles (72.4
km) from where his packmates were on that date
(Fig. 18 and p. 11).

Because No. 13 had taken such a divergent route
from that of No. 11 and 12 upon splitting, and then
had met them again at a point so far from (1) where
they had split and (2) where either had gone after
the split, mere backtracking would seem to be ruled
out as explanation of how they were able to rendezvous.
Possibly No. 13 backtracked to the separation
point and then followed the others by scent, although
this seems unlikely because of the amount of time
that had elapsed. Perhaps a combination of memory
of the general lay of the land, and some backtracking
and eventually howling and the crossing of each
group's fresh tracks could explain this remarkable
feat.

Distances Traveled

The average daily straight-line distances (average
of all known 24-hour moves) traveled by Wolf No.
10 was 3.6 miles (5.8 km). For Wolf No. 11 and her
associates it was 5.8 miles (9.3 km) for the period
before the settled Phase of their movements. The
daily summer straight-line movements of an immature
radioed female in Ontario ranged from 0.0 to 3.5
miles (5.6 km) per day and averaged 1.0 (1.6 km)
per day (Kolenosky and Johnston 1967). Mech and
Frenzel (1971) found that the average daily straight-line
distance traveled in Minnesota by three lone
wolves was 2.0, 1.0 and 2.9 miles (3.2, 1.6, and 4.6
km), and a pack of five averaged 2.5 miles (4.0 km)
straight-line distance per day. A pack of eight wolves
in Ontario traveled actual distances of 0.0 to 13.2
miles (21.1 km) per day during winter with an average
movement of 4.4 miles (7.1 km) per day (Kolenosky
1972).

Thus distances moved by both lone Wolf No. 10
and the pack were greater than the distances reported
for lone wolves and packs in their native range. In
Harger's (1970) study of homing in black bears, he
also found increased movement by displaced animals.

There was a general reduction in distances moved
by the pack in May and June after the wolves had
settled in Iron County (Fig. 19), compared with
their earlier exploratory movements (Fig. 24). The
movements during the Settled Phase were similar to
those reported from the studies in Ontario and
Minnesota.[Pg 24]

[Pg 25]

Home Range Size

At least in some areas, wolves are territorial (Mech
1972, 1973), and the sizes of their home ranges are
restricted somewhat by boundaries established by the
scent marks of surrounding packs (Peters and Mech
1975). The introduced wolves probably encountered
no native packs with established territories (Hendrickson
et al. 1975), so they would not be similarly restricted.
The total area that wolves No. 11, 12, and
13 explored, 2,918 square miles (7,586 km²), is larger
than any reported from the Great Lakes area and is
comparable to home ranges of "tundra wolves"
(Mech 1970). Even the area in which they settled
(May 7 to July 6) until the deaths of the males was
246 square miles (637 km²), which is larger than
most reported ranges in the Great Lakes Region.

The deaths of the two males seemed to cause an
increase in both daily distance traveled and home
range in Wolf No. 11. Essentially she began traveling
as extensively as do lone wolves in Minnesota (Mech
and Frenzel 1971).

The home range of Wolf No. 10 from March
through mid-November, 346 square miles (895.7
km²), was smaller than those of lone wolves in Minnesota
(Mech and Frenzel 1971). Apparently she was
still expanding her range when killed, however.

Selection of a Territory

The eventual settling of the pack of translocated
wolves into a territory would be expected because
such behavior is characteristic of wolves in other
areas. The translocated pack did settle into a territory
of 246 square miles (637 km²) after about 2 months
(Fig. 18). Although the region where they settled was
not as remote as the release area, it was more inaccessible
than most of the rest of the 1,631 square
mile (4,224 km²) area they explored after dispersing.
As with the rest of Upper Michigan, the pack's
adopted territory was inhabited by a moderate population
of deer and beavers. It seems significant that
this area is one of three where a few native Michigan
wolves are known to still exist (Hendrickson et al.
1975).

Vulnerability and Mortality

It could be expected that the translocated wolves
would be more vulnerable than wolves in their native
environment. Although no data are available from
any previous study of translocated wild wolves, Harger's
(1970) investigation of displaced wild black
bears showed that they were more vulnerable. In our
study, it was clear that during the Directional Movement
and Exploratory Phases Wolves No. 11, 12, and
13 were observed by local residents an unusual number
of times (Table 6). No. 10, which did not explore
such an extensive area and which spent considerable
time in a more remote area, was seen less (Table 5).

It is not clear why the wolves were not killed by
humans during these periods when they appeared so
vulnerable. Perhaps the novelty of the transplant
coupled with the awareness that frequent aerial
checks were being made of the wolves had some effect.
Furthermore, spring is not generally a season of intensive
hunting and trapping.

Whatever the explanation, the wolves did survive
what seemed to be their most vulnerable period. We
do not believe that the deaths of the wolves can be
attributed to the conditions of their translocations.
Instead, we think that the most important factor in
their demise was the accessibility of the area to
human beings and the attitudes of humans towards
wolves.

As indicated earlier, there appears to be an inverse
relationship between human density and wolf density
in the Great Lakes Region (Table 1). Wolves are
vulnerable to both accidental and deliberate mortality
from humans. For example, in winter 1947–48 at
least 14 wolves were struck by automobiles in northern
Ontario (DeVos 1949). In Michigan, a $15–$20
bounty still exists on coyotes, so these animals are
commonly shot and trapped. Because many people
cannot distinguish wolves from coyotes, and because
wolves are often caught in the same kind of trap
sets made for coyotes, wolves might be killed accidentally.

Whether the killing of the translocated wolves was
deliberate or accidental is unknown except in the case
of No. 11. No. 11 was caught accidentally in a coyote
trap, but was killed deliberately when the trapper
thought the animal might attack him. The best guess
about No. 12, which was killed by a car, is that it was
accidental. No. 10 and No. 13 were shot, but it is
possible that the hunters in each case may have mistaken
them for coyotes. On the same day that No. 10
was killed, a deer hunter shot a 76-lb. (34.5 kg)
native Michigan wolf and turned himself in to authorities,
stating that he had thought it was a coyote,
and in March 1975 there was a similar occurrence.

Some Upper Michigan residents strongly opposed
the transplant experiment, largely out of concern for
deer populations. The Northern Michigan Sportsmen's
Association passed a resolution against it, and
the Baraga County Wolf Hunters Association was
formed with the express purpose of interfering with
the transplant effort. This association offered a reward
of $100 to a person killing a wolf (Fig. 25).
Supposedly 132 memberships at $1.50 each were sold.[Pg 26]

It is unlikely that members of the Baraga County
group killed the experimental wolves, for it would be
extremely difficult for anyone to deliberately hunt
down and kill a wolf. Most wolves that are shot anywhere
just happen to be seen by a few of the hundreds
of thousands of hunters that are afield or by
local residents who keep a gun handy. Thus the more
accessible the area, and the higher the density of
human beings, the greater the chances that wolves
will encounter such people.

Of course there was also excellent public support
for the experiment. With weekly newspaper accounts
of the travels of the wolves, many people began to
develop an interest in, and sympathy for, the wolves.
Some letters in the newspapers expressed regret that
the animals had been killed.

Food Habits and Predation

The translocated wolves apparently scavenged more
in Michigan than in Minnesota, at least shortly after
their release. There were no known garbage dumps
within their native territory. The dumps in Michigan
presumably offered more readily available food during
a time when the wolves appeared preoccupied with
extensive travel.

Nevertheless, the wolves did kill at least the three
deer that we found, and no doubt took several others.
Although the sample size is small, the results of our
analysis of the condition of the deer are consistent
with those from other studies, indicating that wolves
prey primarily on debilitated deer (Pimlott et al.
1969, Mech and Frenzel 1971).

All three deer killed by the wolves were seriously
malnourished, with 6% or less fat content in the marrow
of their femurs, or thigh bones. At less than 25%
fat in the marrow, serious malnutrition has developed
(Cheatum 1949). (In comparison, the femur fat of
59 doe deer killed by automobiles in the Upper Peninsula
in March and April 1974 averaged 46%,
according to Dr. L. D. Fay, Michigan Department of
Natural Resources.) In addition, one of the animals
killed by the wolves had been wounded by a bullet
and had a broken leg; all three were does, and were
over 4 years of age, a factor that Pimlott et al. (1969)
and Mech and Frenzel (1971) have also found important
in wolf kills.

An Alternate Approach

Although the time of release for the four wolves in
this study was selected in order to maximize chances
that they would remain in their new range, possibly
a release earlier in winter would be more successful.
The failure of the adult female to conceive was
probably a result of captivity and handling, although
this needs confirmation through additional studies.
Nevertheless, an early winter release might be favored
by deep snows hindering travel. Furthermore, by
breeding season in late February the wolves might
already have settled into an area. Then the entire
breeding cycle might take place outside captivity and
stand a better chance of succeeding.

CONCLUSIONS

Three principal conclusions can be drawn from the
results of this experiment: (1) It is possible to transplant
a pack of wild wolves into a new range. That
new range, however, must be large enough to permit
some initial wandering. The animals cannot be expected
to establish a home range centered on or even
including the point of release. (2) The habitat in
Upper Michigan apparently is adequate to support
wolves, in terms of food and cover, for the carcasses
of the two experimental wolves that could be examined
intact had maintained or improved their condition
during their 6-to-8-month residence in Michigan.
(3) The reason for the failure of the experimental
wolves to re-establish themselves was direct mortality
by human beings, just as Hendrickson et al. (1975)
concluded was the case for the failure of native and
immigrant Michigan wolves to re-establish a population.
This mortality probably is related to two factors,
negative human attitudes toward wolves and
accessibility of humans to wolf range.

We are convinced that, ecologically, wolves can be
re-established in Upper Michigan. However, a successful
program of re-establishment will require the
following:

1. A survey of public attitudes in Upper Michigan
toward re-establishing wolves,

2. An intensive public relations campaign to promote
an understanding of wolf ecology and the benefits
of a wolf population,

3. Suspension or removal of the bounty on coyotes,

4. Releases of additional wolves in larger numbers
perhaps over a period of a few years, if public
attitudes appear favorable,

5. A concentrated effort to inform the public of the
penalties for killing wolves,

6. A concerted law enforcement program, and

7. Monitoring of translocated animals through radio-tracking
to determine the results.

Fig. 25.—Although the transplant experiment enjoyed
wide public support, some people opposed it
and organized the Baraga County Wolf Hunters
Association to try to prevent the re-establishment
effort

[Pg 27]

ACKNOWLEDGMENTS

This project was a cooperative effort among the
Michigan Department of Natural Resources, The
Minnesota Department of Natural Resources, the
U. S. Fish and Wildlife Service, Northern Michigan
University, the Huron Mountain Wildlife Foundation,
and the National Audubon Society.

The Michigan and Minnesota Departments of
Natural Resources provided the legal permits and
logistical support necessary for the transplant. The
Fish and Wildlife Service assisted in the planning and
fund-raising for the overall project, and provided the
technical expertise in the live-trapping, radio-tagging
and radio-tracking of the wolves. Northern Michigan
University initiated and administered the project and
conducted the Michigan aspects of the work. Financial
support was provided by the Huron Mountain
Wildlife Foundation and the National Audubon
Society.

Dr. U. S. Seal of the Veterans Administration
Hospital in Minneapolis analyzed and interpreted the
blood samples, and Dr. Ray D. Barnes, University of
Minnesota, the female reproductive tracts.

The authors wish to thank all of the people mentioned
above and the following individuals: Ralph
Bailey and Robert Rafferty, Michigan Department of
Natural Resources; Jeff Renneberg, U. S. Fish and
Wildlife Service; Fred Harrington, State University of
N. Y. at Stony Brook; Roger Peters, University of
Michigan; Tom Jernstad, Leo Maki, and Leo Wouri,
Huron Mountain Club; the late William P. Harris Jr.
and Theodore A. McGraw of the Huron Mountain
Wildlife Foundation; Edward H. Brigham III, National
Audubon Society; Dennis Diaz and William
Rowloff, pilots of Northern Airmotive; Robert Neil,
owner of the research airplane; Denis Kallery, Cynthia
Watt, and George Wilson of Marquette; and
last but certainly not least, wolf-trapper Robert Himes
of Ray, Minnesota.

LITERATURE CITED

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The Audubon Conservation Report series:

National Audubon Society, 950 Third Avenue, New York City 10022