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Title: An Experimental Translocation of the Eastern Timber Wolf
Author: Weise, Thomas F., Robinson, William L., Hook, Richard A., Mech, L. David
Language: English
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Copyright Status: Not copyrighted in the United States. If you live elsewhere check the laws of your country before downloading this ebook. See comments about copyright issues at end of book.

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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.

                                       [Illustration: Lynn A. Greenwalt]

                                       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 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.



                              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).

[Illustration: _Fig. 1.--Original and present range of the Eastern
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.

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.

[Illustration: _Fig. 2.--Range of the wolf in Upper Michigan in 1973,
and the release point (from Hendrickson et al. 1975)_]

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

  ----------------------------------------------------------------------
                                                         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 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.



                                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.

[Illustration: _Fig. 3.--Capture and release points of the translocated
wolves_]

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.

[Illustration: _Fig. 4.--Wolf caught in trap (Photo by Don Breneman)_]

[Illustration: _Fig. 5.--The captured wolves were drugged and carried to
an enclosure in Minnesota (USFWS Photo by L. David Mech)_]

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.

[Illustration: _Fig. 6.--Before being transported to Michigan, each wolf
was weighed (USFWS Photo by Don Reilly)_]

_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 × 1.0"   0.5 × 0.75"
                                                (13×25 mm)   (13×19 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).

[Illustration: _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.

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

[Illustration: _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.

[Illustration: _Fig. 9.--The anesthetized wolves were placed aboard an
aircraft in International Falls, Minnesota (USFWS Photo by Don Reilly)_]

[Illustration: _Fig. 10.--The wolves were kept lightly drugged during
the flight to Michigan (USFWS Photo by L. David Mech)_]

[Illustration: _Fig. 11.--In the Huron Mountain area of Upper Michigan
the wolves were taken to another holding pen (Photo by Don Pavloski)_]

[Illustration: _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.

[Illustration: _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).

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.

[Illustration: _Fig. 14.--The Minnesota wolves in their Michigan pen
(Photo by Tom Weise)_]

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
  ---------------------------------------------------------

[Illustration: _Fig. 15.--The wolves often used woods roads for
traveling (Photo by James Havemen)_]

[Illustration: _Fig. 16.--The released wolves were sometimes observed
from the aircraft feeding on deer they had killed (Photo by Richard P.
Smith)_]


                  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.

[Illustration: _Fig. 17.--Movements during the Post-release and
Directional Movement Phases of Wolves No. 11, 12, and 13_]

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 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.

[Illustration: _Fig. 18.--Exploratory and Settled Phases in the
movements of Wolves No. 11, 12, and 13_]

[Illustration: _Fig. 19.--Movements of No. 11 after the death of No. 12
and 13_]

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).

[Illustration: _Fig. 20.--Movements of Wolf No. 10_]

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 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_

  ---------------------------------------------------------------------
               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

[Illustration: _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_

  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 ½ 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, ¾
            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.

[Illustration: _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.

[Illustration: _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_

  ----------------------------------------------------------------------
  Wolf No.     10              11            12              13
  ----------------------------------------------------------------------
  Sex        Female          Female         Male            Male

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

  Date       Nov. 16[14]     Sept. 20       June 28 to      Early
  killed                                    July 4          July[14]

  Date       Nov. 18         Sept. 20       July 11         July 28
  found

  Manner     Gunshot in      Gunshot in,    Struck by       Gunshot in
  of death   head and        head, after    automobile      head and
             right foreleg   being trapped                  chest

  Location   Van Riper Lake  Floodwood      1.9 miles (3.0  2 miles (3.2
  of death   5.4 miles (8.7  Plain 3.1      km) north of    km) south of
             km) north of    miles (5.0     Amasa (T45N-    Sagola (T42N
             Champion (T49N  km) south of   R33W-Sec 17)    -R30W-Sec 5)
             -R30W-Sec 36)   Witch Lake
                             (T44N-R24W-
                             Sec 11)

  Weight     52 lb.          56.5 lb.
             (23.6 kg)       (25.6 kg)      Unknown[15]     Unknown[15]

  Condition  Excellent       Good           Unknown[15]     Unknown[15]
  ----------------------------------------------------------------------

[14] Estimate

[15] Decomposed

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 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 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.

[Illustration: _Fig. 24.--Straight-line distances between consecutive
locations for (A) Wolves No. 11, 12, and 13, (B) Wolf No. 10. (Gaps
between data points represent periods when no data were obtained.
Because these periods varied, and because distance traveled is partly a
function of duration between locations, it is only valid to grossly
compare distances from one period to the next.)_]


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 km^2), 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 km^2), 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 km^2), 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 km^2) 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 km^2) 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.

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.

[Illustration:]

   ___________________________________
  |                                   |
  |  =F. E. Noble, Sr., President=    |
  |                                   |
  |        =BARAGA COUNTY=            |
  |  =WOLF HUNTERS ASSOCIATION=       |
  |                                   |
  |=Preserve Our Deer  "Shoot a Wolf"=|
  |                                   |
  |    =$100.00 Reward For Any Wolf=  |
  |                                   |
  |        =$1.50 Membership Fee=     |
  |___________________________________|

_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_]



                             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


  Brown, C. E. 1936. Rearing wild animals in captivity, and
    gestation periods. J. Mammal. 17:10-13.

  Burkholder, B. L. 1959. Movements and behavior of a wolf
    pack in Alaska J. Wildl. Manage. 23:1-11.

  Cheatum, E. L. 1949. Bone marrow as an index of
    malnutrition in deer. N. Y. State Conservationist
    3(5):19-22.

  Cochran, W. W., and R. D. Lord. 1963. A radio-tracking
    system for wild animals. J. Wildl. Manage. 27:9-24.

  DeVos, A. 1949. Timber wolves (_Canis lupus lycaon_)
    killed by cars on Ontario highways. J. Mammal. 30:197.

  DeVos, A. 1950. Timber wolf movements on Sibley Peninsula,
    Ontario. J. Mammal. 31:169-175.

  Dice, L. R. 1952. Natural communities. Univ. of Mich.
    Press, Ann Arbor. 547 p.

  Douglass, D. W. 1970. History and status of the wolf in
    Michigan. p. 6-8 _In_ Jorgensen, S. E., C. E. Faulkner,
    and L. D. Mech (_Ed._) Proc. Symp. on Wolf Management in
    Selected Areas of North America. U. S. Fish & Wildl.
    Serv., Twin Cities, Mn. 50 p.

  Harger, E. M. 1970. A study of homing behavior of black
    bears. Unpubl. Master's Thesis. North. Mich. Univ.,
    Marquette. 81 p.

  Hendrickson, J., W. L. Robinson, and L. D. Mech. 1975. The
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    press).

  Henshaw, R. E. and R. O. Stephenson. 1974. Homing in the
    gray wolf (_Canis lupus_). J. Mammal. 55:234-237.

  Kolenosky, G. B. 1972. Wolf predation on wintering deer in
    east-central Ontario. J. Wildl. Manage. 36:357-369.

  Kolenosky, G. B., and D. H. Johnston. 1967. Radio-tracking
    timber wolves in Ontario. Amer. Zool. 7:289-303.

  Kuyt, E. 1972. Food habits of wolves on barren-ground
    caribou range. Can. Wildl. Serv. Rep. Series No. 21. 36
    p.

  Laundre, J. 1975. An ecological survey of the mammals of
    the Huron Mountain Area. Occ. Pap. Huron Mt. Wildl.
    Found. No. 2.

  Mech, L. D. 1966. The wolves of Isle Royale. U. S. Nat.
    Park Serv. Fauna Ser. 7. 210 p.

  Mech, L. D. 1970. The wolf: the ecology and behavior of an
    endangered species. The Nat. Hist. Press. Garden City,
    New York. 384 p.

  Mech, L. D. 1972. Spacing and possible mechanisms of
    population regulation in wolves. Am. Zool. 12(4): 9
    (abstract).

  Mech, L. D. 1973. Wolf numbers in the Superior National
    Forest of Minnesota. North Cent. For. Exp. Stn., St.
    Paul, Minn. USDA For. Serv. Res. Pap. NC-97. 10 p.

  Mech, L. D. 1974. Current techniques in the study of
    elusive wilderness carnivores. Proc. XI Int. Cong. Game
    Biol., Stockholm, p. 315-322.

  Mech, L. D., and L. D. Frenzel, Jr., eds. 1971. Ecological
    studies of the timber wolf in northeastern Minnesota.
    North Cent. For. Exp. Stn., St. Paul, Minn. USDA For.
    Serv. Res. Pap. NC-52. 62 p.

  Mech, L. D., and R. A. Rausch. 1975. Status of the wolf in
    the United States, 1973. Proc. of First Meeting of
    IUCN-SSC Wolf Specialist Group, Stockholm (In press).

  Merriam, H. R. 1964. The wolves of Coronation Island.
    Proc. Alaska Sci. Conf. 15:27-32.

  Nellis, C. H. 1968. Some methods for capturing coyotes
    alive. J. Wildl. Manage. 32:402-405.

  Peters, R. R., and L. D. Mech. 1975. Scent-marking in
    wolves: a field study. American Scientist 63(4) (In
    press).

  Peterson, R. O. 1974. Wolf ecology and prey populations on
    Isle Royale. Unpubl. Ph.D. Dissertation, Purdue Univ.,
    Lafayette, Ind. 368 p.

  Phillips, R. L., and L. D. Mech. 1970. Homing behavior in
    a red fox. J. Mammal. 51:621.

  Pimlott, D. H. 1967. Wolf predation and ungulate
    populations. Amer. Zool. 7:267-278.

  Pimlott, D. H., J. A. Shannon, and G. B. Kolenosky. 1969.
    The ecology of the timber wolf in Algonquin Park. Ont.
    Dept. Lands and Forests. Res. Rep. (Wildlife) No. 87. 92
    p.

  Rausch, R. A. 1967. Some aspects of the population ecology
    of wolves, Alaska. Am. Zool. 7:253-265.

  Schenkel, R. 1947. Expression studies of wolves. Behaviour
    1:81-129. (Translation from German by Agnes Klasson).

  Seal, U. S., A. W. Erickson, J. G. Mayo. 1970. Drug
    immobilization of the Carnivora. Int. Zoo Yearbook
    10:157-170.

  Seal, U. S., L. D. Mech, and V. Van Ballenberghe. 1975.
    Blood analyses of wolf pups and their ecological and
    metabolic interpretation. J. Mammal. 56:64-75.

  Spencer, J. S., Jr., and R. E. Pfeifer. 1966. The growing
    timber resource of Michigan--1966. Unit 2--the Western
    Upper Peninsula, Mich. Dept. Nat. Res., Lansing.

  Stenlund, M. H. 1955. A field study of the timber wolf
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    Minnesota. Minn. Dept. Cons. Tech. Bull. No. 4. 55 p.

  Storm, G. L. 1972. Population dynamics of red foxes in
    northcentral United States. Ph.D. Thesis, Univ. of
    Minn., Mpls., 227 p.

  Thompson, D. Q. 1952. Travel, range, and food habits of
    Timber Wolves in Wisconsin. J. Mammal. 25:37-43.

  U. S. Bureau of Census. U. S. Census of Population: 1970.
    No. of inhabitants. Final Rept. PC(1).

  Westover, A. L. 1971. The use of a hemlock-hardwood winter
    yard by white-tailed deer in northern Michigan. Occ.
    Pap. Huron Mt. Wildl. Found. No. 1. 59 p.

  Wolfe, M. L., and D. L. Allen. 1973. Continued studies of
    the status, socialization, and relationships of Isle
    Royale wolves, 1967 to 1970. J. Mammal. 54:611-635.

  Woolpy, J. H. 1968. The social organization of wolves.
    Nat. Hist. 77(5):46-55.

  Woolpy, J. H., and B. E. Ginsburg. 1967. Wolf
    socialization: a study of temperament in a wild species.
    Am. Zool. 7:357-363.


_The Audubon Conservation Report series_:

 No. 1 THE GOLDEN EAGLE IN THE TRANS-PECOS AND EDWARDS
        PLATEAU OF TEXAS by Walter R. Spofford. 1964.

 No. 2 THE SUBURBAN WOODLAND/Trees and Insects in the Human
        Environment by Roland C. Clement and Ian C. T.
        Nisbet. 1972.

 No. 3 SOME ENVIRONMENTAL AND ECONOMIC IMPLICATIONS OF THE
        NATIONAL WATER COMMISSION'S 1972 DRAFT REPORT by
        Roland C. Clement and Robert K. Davis. 1973.

 No. 4 PROCEEDINGS OF A CONFERENCE ON PEREGRINE FALCON
        RECOVERY Edited by Roland C. Clement. 1974.

 No. 5 AN EXPERIMENTAL TRANSLOCATION OF THE EASTERN TIMBER
        WOLF by Thomas F. Weise, William L. Robinson,
        Richard A. Hook, and L. David Mech. 1975.

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





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