Ecological Studies of the Timber Wolf in Northeastern Minnesota L. David Mech_ Louis Daniel Frenzel_ Robert R. Ream_ John W. Winship_ and P. D. Karns 793 downloads.pdf

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Title: Ecological Studies of the Timber Wolf in Northeastern Minnesota

Author: L. David Mech
Louis Daniel Frenzel
P. D. Karns
Robert R. Ream
John W. Winship

Release date: October 2, 2011 [eBook #37595]
Most recently updated: January 8, 2021

Language: English

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*** START OF THE PROJECT GUTENBERG EBOOK ECOLOGICAL STUDIES OF THE
TIMBER WOLF IN NORTHEASTERN MINNESOTA ***

USDA FOREST SERVICE
RESEARCH PAPER NC-52
1971

Ecological
Studies of the
TIMBER
WOLF
in
Northeastern
Minnesota

NORTH CENTRAL FOREST EXPERIMENT STATION
FOREST SERVICE
U. S. DEPARTMENT OF AGRICULTURE

FOREWORD
The largest population of timber wolves remaining in the United States (excluding Alaska) lives in
northern Minnesota. Many of these wolves inhabit the Superior National Forest, so protecting the
habitat of this endangered species is largely a Forest Service responsibility.
As the "Age of Ecology" broadens into the 1970's, wolves and wolf habitat will become a subject of
concerted research. Forest land managers will have to know more about how the timber wolf fits into
a forest system. Building on nearly 50 years of research in northern forests, we at the North Central
Station intend to expand our studies of wildlife habitat. We are happy to publish the enclosed papers as
one step in this direction.
D. B. King, Director

North Central Forest Experiment Station
D. B. King, Director
Forest Service—U.S. Department of Agriculture
Folwell Avenue
St. Paul, Minnesota 55101

ECOLOGICAL STUDIES OF THE TIMBER
WOLF
IN NORTHEASTERN MINNESOTA

L. David Mech and L. D. Frenzel, Jr. (Editors)

CONTENTS

Movements, Behavior, and Ecology of Timber Wolves in
Northeastern Minnesota 1
L. David Mech, L. D. Frenzel, Jr., Robert R. Ream,
and John W. Winship
An Analysis of the Age, Sex, and Condition of Deer
35
Killed by Wolves in Northeastern Minnesota
L. David Mech and L. D. Frenzel, Jr.
The Effect of Snow Conditions on the Vulnerability of
51
White-Tailed Deer to Wolf Predation
L. David Mech, L. D. Frenzel, Jr., and P. D. Karns
The Possible Occurrence of the Great Plains Wolf in
60
Northeastern Minnesota
L. David Mech and L. D. Frenzel, Jr.

THE AUTHORS

Dr. Mech, formerly with the Department of Biology,
Macalester College, St. Paul, Minnesota, is now employed
by the U.S. Bureau of Sport Fisheries and Wildlife, Twin
Cities, Minnesota.
Dr. Frenzel, formerly with the Department of Biology,
Macalester College, St. Paul, Minnesota, is now employed
by the Department of Entomology, Fisheries, and Wildlife,
University of Minnesota, St. Paul, Minnesota.
Dr. Ream, formerly with the North Central Forest
Experiment Station (maintained in cooperation with the
University of Minnesota), Forest Service, U.S. Department
of Agriculture, is now employed by the School of Forestry,
University of Montana, Missoula, Montana.
Mr. Winship is with the U.S. Bureau of Sport Fisheries and
Wildlife, Twin Cities, Minnesota.
Mr. Karns is with the Minnesota Department of
Conservation, Forest Lake, Minnesota.

For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402

MOVEMENTS, BEHAVIOR, AND ECOLOGY OF TIMBER
WOLVES
IN NORTHEASTERN MINNESOTA
L. David Mech, L. D. Frenzel, Jr.,
Robert R. Ream, and John W. Winship
The largest population of wolves (Canis lupus) remaining today in the continental United States
outside of Alaska is in northern Minnesota. As of mid-1970 this population was not legally protected,
and the species, which once ranged over almost all of North America, is now considered by the U.S.
Department of the Interior to be in danger of extinction in the contiguous 48 States. Until the present
research, the only field studies of Minnesota wolves were those of Olson (1938 a, b) and Stenlund
(1955). Those investigations provided much useful general information about Minnesota wolves and
gave the present authors an excellent background with which to begin more detailed investigations.
This paper reports on the basic aspects of a series of studies that began in 1964, and concentrates
primarily on wolf movements and activity, social behavior, hunting behavior, and population
organization. Most of the data were collected during January, February, and March 1967; February,
November, and December 1968; and January through August 1969. A total of 192 days was spent in
the field.
According to a distribution map of wolf subspecies (Goldman 1944), the race of wolves in our study
area is Canis lupus lycaon. However, evidence presented by Mech and Frenzel (see page 60) suggests
that there may be strong influence by C. l. nubilus, a more western race of wolf formerly thought to be
extinct (Goldman 1944).
Between 1965 and the present, wolves in the study area were neither protected nor bountied, and the
influence of trapping and hunting is thought to have been negligible.

THE STUDY AREA

This study was conducted in the Superior National Forest (fig. 1) in northern St. Louis, Lake, and
Cook Counties of northeastern Minnesota (92° west longitude, 48° north latitude), an area well
described by Stenlund (1955). Most of the data were collected from within and immediately south of
the Boundary Waters Canoe Area, a special wilderness region in which travel by motorized vehicles is
restricted. The total study area encompasses approximately 1.5 million acres, and numerous lakes and
rivers comprise about 15 percent of this area (fig. 2). The topography varies from large stretches of
swamps to rocky ridges, with altitudes ranging from 1,000 to 2,300 feet above sea level (fig. 3).
Winter temperatures lower than -30° F. are not unusual, and snow depths generally range from 20 to
30 inches on the level. However, an important exception occurred in early 1969 when depths of 45
inches and more accumulated in much of the area. Further details on snow conditions in the study area
during the period of this investigation are given by Mech et al. (see page 51). Conifers predominate in
the forest overstory, with the following species present: jack pine (Pinus banksiana Lamb.), white pine
(P. strobus L.), red pine (P. resinosa Ait.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (P.
glauca (Moench) Voss), balsam fir (Abies balsamea (L.) Mill.), white cedar (Thuja occidentalis L.),
and tamarack (Larix laricina (DuRoi) K. Koch). However, as a result of extensive cutting and fires
much of the conifer cover is interspersed with large stands of white birch (Betula papyrifera Marsh.)

and aspen (Populus tremuloides Michx.). Detailed descriptions of the forest vegetation were presented
by Ohmann and Ream (1969).

Figure 1 (left half)—Map of the study area.
(Click for larger image.)

Figure 1 (right half)—Map of the study area.
BOUNDARY WATERS CANOE AREA SUPERIOR NATIONAL
FOREST
(Click for larger image.)

Figure 2.—Lakes are common throughout most of the study area. (Photo courtesy of L. D.
Mech.)

Figure 3.—Ridges, islands, swamps, and bays are part of the variable topography in the Superior
National Forest. (Photo courtesy of L. D. Mech.)

METHODS

The observations discussed in this paper were all made from aircraft, the method of flying being that
reported by Burkholder (1959) and Mech (1966a). The following aircraft were used (in order of size):
Aeronca Champ,[1] Supercub, Cessna 172, Cessna 180, and Cessna 206. The smaller aircraft were
excellent for holding in tight circles during observations but had the disadvantage of being slow and
cold; the larger planes could cover the study area much more quickly and were more comfortable, but
were not as maneuverable during observations. For radiotracking, to be discussed below, the best
compromise seemed to be a Cessna 172.
To make observations of wolves, we flew over frozen waterways until tracks were found, and then
followed the tracks until we lost them or saw the wolves (fig. 4). Several times we located wolves
directly just by scanning the lakes. However, because there seemed to be a number of packs in the
area, and because most wolves were the same color (with the exception of a few black or white
individuals) (see Mech and Frenzel, page 60), it usually was not possible to follow packs from one day
to the next and be certain of identification. Moreover, it was impossible to locate any pack at will
because most wolves also spent much time inland.

Figure 4.—An important technique used in the study involved aerial tracking and observing of
wolf packs. (Photo courtesy of L. D. Frenzel.)

Therefore, to facilitate our observations and to obtain data on wolf movements and extent of range, we
began a radiotracking program in 1968-69. A professional trapper, Robert Himes, was employed to
capture the wolves. Using Newhouse No. 4 and 14 steel traps at scent-post sets, he caught two wolves,
and captured another with a live-snare similar to that used by Nellis (1968); the senior author trapped
two additional wolves (fig. 5).
The four wolves held in steel traps were restrained by a choker (fig. 6), and then anesthetized by
intramuscular injections (fig. 7A, B) of a combination of 30 mg. of phencyclidine hydrochloride
(Sernylan, Parke-Davis Co.) and 25 mg. promazine hydrochloride (Sparine, Wyeth Laboratories) as
prescribed by Seal and Erickson (1969); these drugs proved most satisfactory.

The fifth wolf (a
female), which was
captured around the
chest by the live-
snare, was handled
without drugs. A
forked stick was used
to hold down her head
(Kolenosky and
Johnston 1967), and
she offered no
resistance (fig. 8).
Evidently she went
into shock or some
other
psychophysiological
Figure 5.—A wolf caught in a trap. state of
(Photo courtesy of D. L. unconsciousness, for
after her release she Figure 6.—A choker was used to
Breneman.) restrain wolves caught in traps.
remained on her side
and did not move for (Photo courtesy of D. L.
1.5 hours, despite our Breneman.)
prodding during the first few minutes (fig. 9). Then suddenly
she leaped up and ran off.
Each wolf was examined, outfitted with a radio transmitter collar 15 inches inside circumference (fig.
10) and tagged with identification numbers in both ears (fig. 11). Each transmitter was of a different
frequency in the 150 MHz range, emitted a pulsed signal ranging from 75 to 350 pulses per minute,
and had a calculated life of at least 300 days (fig. 12). Two types of 12-inch whip antennas were used
on the transmitters: one type extended up the side of the collar and then stuck out above for 6 inches;
the other was fully attached inside the collar and extended up one side, around the top, and partly
down the other side. The transmitter, batteries, and antenna were molded into a collar of acrylic
weighing 11 ounces (Mech et al. 1965).[2] All radio equipment functioned flawlessly for at least 5
months, and one transmitter continued operating for at least 9 months.
For tracking radio-equipped wolves, a directional yagi antenna (fig. 13) was attached to each of the
wing struts of an aircraft and connected inside to a portable receiver. The usual tracking technique was
to fly at 1,500 to 3,000 feet elevation to the last known location of the wolf being sought (fig. 14). If a
signal was not obtained at that point, the aircraft spiraled upward until the signal was found or until
10,000 feet altitude had been reached. If the signal still was not heard, a search pattern was flown at
10,000 feet. The range of the signal from this altitude was 15 to 35 miles; at 3,000 feet it was 10 to 15
miles. Collars with antennas molded fully inside gave only about two-thirds the range of those
protruding partly, but could be expected to last longer because the antennas could not break off. It is
unknown whether any protruding antennas did break during the study, but on January 5, 1970, one
wolf was recaptured, and its antenna had broken.
When a signal was received, the aircraft was headed in the approximate direction of the source until
the signal strength reached a peak; a 90° turn was then made in the direction the signal seemed the
strongest. A series of these maneuvers soon narrowed the area to the point where visual search was
possible. After practice and experience with this technique, we could locate the approximate source of
the signal within 10 to 30 minutes after first receiving it.

Even though the
radiotagged wolves
spent most of their
time inland, often in
stands of conifers,
they were frequently
observed from the
aircraft. The technique
was to circle at 300 to
800 feet altitude
around a radius of a
quarter mile from the
point where the
strongest signal
emanated. From
December through
April, 65 percent of
the wolves located by
radio were sighted;
the rate was much Figure 8.—Once pinned by the
higher for more forked stick, the wolf ceased
experienced struggling. (Photo courtesy of
personnel. A pack of Richard Bend.)
Figure 7.—A. A small hypodermic five wolves that was
syringe is loaded with drugs. B. The tracked was seen 31 times out of 33 attempts during February
loaded syringe is used on the end of and March.
a pole. (Photos courtesy of D. L.
Breneman.) Whenever wolves were located, radiotagged or not,
observations were made from an altitude that did not disturb
them. Packs varied in the concern shown the aircraft, but only
one or two ran
from it. The
radiotagged
wolves, and a pack
of 10 to 13
animals, were
habituated to the
aircraft and usually
could be observed
from altitudes of
500 feet and less
Figure 9.—After release, the wolf lay without
still for 1½ hours before jumping up disturbance (fig.
and running off. (Photo courtesy of L. 15).
D. Mech.)
Almost all the Figure 10.—A radio transmitter
radiotracking was collar was placed around the neck
done from aircraft, but when inclement weather prevented of each trapped wolf. (Photo
flying, some attempts from the ground succeeded when courtesy of D. L. Breneman.)
wolves were close enough to roads. The usual range on the
ground was 0.75 to 1.50 miles. One wolf was approached to within 35 feet through radiotracking.

Figure 11.—Each ear of the wolf
was tagged with identifying Figure 12.—Each radio collar had
numbers. (Photo courtesy of a different frequency tuned to
Richard Bend.) special receivers, which allowed
each wolf to be identified. (Photo
courtesy of D. L. Breneman.)

Figure 14.—The tracking aircraft
Figure 13.—Directional yagi was usually flown at altitudes of
antennas fastened to the wing struts 1,500 to 3,000 feet. (Photo courtesy
of the aircraft were necessary to of Dick Shank.)
"home in" on the wolves. (Photo
courtesy of U.S. Bureau of Sport
Fisheries and Wildlife.)

Figure 15.—The wolves studied soon became accustomed to the aircraft
and could then be observed during their natural activity. (Photo courtesy
of L. D. Mech.)

FOOTNOTES:
[1] Mention of trade names does not constitute endorsement by the USDA Forest Service.
[2] The acrylic collar was fashioned by the Davidson Co., Minneapolis, Minnesota, which also produced
some of the transmitters. Other transmitters and two radio receivers were manufactured by the AVM
Instrument Co., Champaign, Illinois.

RESULTS AND OBSERVATIONS

Aerial observations made during this study involved 490 hours distributed as follows: January,
February, March 1967—124 hours; February 1968—10 hours; December 1968 through August 1969
—356 hours. Seventy-seven observations involving a total of 323 wolves were made (table 1),
excluding animals located through radiotracking.
One male and four female wolves were radiotagged, and they and their associates were followed
intermittently for periods of 5 to 8 months (table 2). All except one initially suffered some injury to a
foot. Three of these animals were seen limping, but only in one case was the limp judged extreme
enough to have significantly affected the movements or behavior of the animal. In that one case, the
wolf (No. 1057) was caught in a steel trap on an extremely cold night, and her foot froze. After that
she was often seen hopping on three legs. She was not able to keep up with her pack, which consisted
of 10 to 13 members, and her movements were much restricted compared with those of other wolves.
However, she was frequently observed feeding on fresh kills, and may even have made them herself.

Table 1.—Sizes of wolf population units observed in northeastern Minnesota

Wolf observations
Population unit[3]
Winter Winter Winters[4]
(number of wolves) Total
1966-67 1968-69 1948-53
Number Percent Number Percent Number Percent Number Percent
1 8 31 17 33 25 32 48 43
2 3 12 6 12 9 12 24 22
3 3 12 2 4 5 6 7 6
4 — — 7 14 7 9 7 6
5 2 7 4 8 6 8 8 7
6 3 12 4 8 7 9 7 6
7 2 7 1 2 3 4 4 4
8 3 12 3 6 6 8 3 3

9 — — 2 4 2 3 3 3
10 2 7 2 4 4 5 — —
11 — — — — — — — —
12 — — 1 2 1 1 1 1
13 — — 2 4 2 3 — —
Total number
109 — 214 — 323 — 318 —
of wolves
Total number
26 — 51 — 77 — 112 —
of observations
Mean population
4.2 — 4.2 — 4.2 — 2.8 —
unit size

FOOTNOTES:
[3] Because wolf packs sometimes split temporarily, these figures may not strictly represent actual pack
sizes; nevertheless they should provide reasonably accurate approximations.
[4] From Stenlund (1955).

Table 2.—Background information on five radiotagged wolves studied in northeastern Minnesota

Wolf Estimated
Usual Location Date Last date Days General
weight[5] associations captured captured located located condition
Number Sex (pounds)
Number
Good, but
two toes
frozen in
T62N- trap;
1051 M 75 [6] R7W- Nov. 27/68 Apr. 24/69 84 animal
None
S18 limped
lightly
for 5-6
wks.
1053 F 60 None T62N- Dec. 10/68 Aug. 29/69 72 Thin; top
R8W- of foot
S13 cut in
trap but
no
broken
bones or
frozen
toes;
limped
for at

least 10
wks.
Thin; two
toes
T61N- lightly
Another wolf
1055 F 60 R10W- Jan. 5/69 May 30/69 65 frozen;
intermittently
S26 no limp
ever
noticed.
Thin;
front foot
frozen in
T66N- trap; lost
1057 F 60 [7] R5W- Jan. 8/69 Apr. 24/69 47 use of
Pack of 13
S33 foot and
could not
stay with
pack.
Good but
thin;
T62N- captured
1059 F 65 Pack of 5 R11W- Jan. 22/69 Aug. 29/69 51 in snare;
S26 no
apparent
injury.

FOOTNOTES:
[5] Wolf 1059, when killed by a trapper on January 10, 1970, appeared to be of the same size and condition
as when radiotagged; she only weighed 53 pounds, however, indicating that probably all the weights are
overestimated.
[6] Tracks of a pack of at least two other wolves came by trap where 1051 was caught; however, there was
never any other indication that 1051 may have been a member of a pack.
[7] A frozen foot prevented 1057 from staying with her pack; but she did associate with other wolves
intermittently and with the whole pack when it came by her restricted area.

The precise ages of the radiotagged wolves were unknown. All individuals, however, had sharp
unworn teeth, indicating that they were all relatively young. No. 1051, the only male studied, had
testes 2.0 cm. long and 1.5 cm. wide; their volume therefore would be less than 4.5 cc. The small size
of these testes, compared with the 7 to 28 cc. reported by Fuller and Novakowski (1955) as the
volume of the testes from wolves taken during fall, would indicate that 1051 had not yet matured.
Since the animal's testes and canine lengths were considerably greater than those of pups caught in a
later study, we presume 1051 was 18 or 30 months old.
Two of the females, No. 1055 and No. 1059, both captured in January, had vulvas that seemed to be
beginning to swell. No. 1059 was killed by a trapper about a year later, on January 10, 1970, and an

examination revealed that she had bred in 1969 and carried five fetuses. Sectioning her incisors and
reading the apparent annulations indicated that she probably was 3+ or 4+ years old.[8]
Three of the wolves were basically lone individuals. One of these, No. 1051, was captured on a night
when tracks of at least two other wolves came by the trap, and this could mean that he had been part
of a pack. However, it is also possible that these were merely tracks of non-associated wolves that
were also traveling through the area. In any case, 1051 was not seen associating with any other wolf
until 4 months after he was caught, and even then the association seemed to be temporary and casual.
It could be argued that capture, handling by humans, or wearing a collar prevented him from regaining
old associations or making new ones. However, the wolves radiotagged by Kolenosky and Johnston
(1967) were quickly accepted back into their packs, and so were two of ours. Thus we conclude that
1051 probably was a lone wolf when captured.
When 1053 was trapped, her tracks were the only ones in the area, and she was never seen closely
associating with another wolf. No. 1055 probably was with another wolf when captured, as evidenced
by tracks. About a month after she was radiotagged she associated with another wolf intermittently for
about 2 weeks, after which she was only seen alone.
No. 1057 and No. 1059 were both members of packs. No. 1057 was captured during the night after a
pack of 13 wolves was seen heading toward the area; 5 days later she was seen with 10 other wolves,
which no doubt represented this same pack. This wolf's association with the pack was interrupted,
however, because of the foot injury sustained during capture. When 1059 was caught, tracks of two
other wolves were seen in the immediate vicinity, and one of the animals was seen within a quarter
mile of the trapped wolf. Three days after 1059's release, and perhaps sooner, she was back with her
pack, with which she remained at least through March.
The detailed histories of the associations of the radiotagged wolves will be discussed in a later section.
Radiotagged wolves were tracked every day that weather permitted during December, January, and
February; every week during March, April, and May; and once a month during June, July, and August
(fig. 16). Information was obtained for a total of 570 "wolf-days"—a wolf-day being a day in which
one radiotagged wolf was located; a pack of five being located for 1 day would constitute 5 wolf-days.

Figure 16.—Distribution of the days on which data were obtained for
each of the radiotagged wolves. Because tracking success was 99

percent, this also represents the distribution of effort. During June, July,
and August, wolves 1053 and 1059 were located 1 day each month.

The last day that animals 1051 and 1057 were heard from was April 24, 1969. Both had traveled long
distances during the previous week and may have moved out of range. Signals from wolf 1055 were
last heard on May 30; this animal had also been ranging widely. Circles with radii of at least 50 miles
around the last known locations of each wolf were searched unsuccessfully for the signals. During all
subsequent tracking nights for the remaining wolves, the missing animals were also sought, but to no
avail. Before the last dates that signals from these animals were heard, attempts to locate marked
animals from the air had failed in only three instances.

FOOTNOTES:
[8] David W. Kuehn, Personal correspondence to L. D. Mech, 1970.

Daytime Activity Patterns

When radiotagged wolves were located, notes were kept on the type of activity they were engaged in;
the results are summarized in figure 17. In a total of 171 observations made between 9:00 a.m. and
6:00 p.m., the wolves were resting 62 percent of the time, traveling 28 percent and feeding 10 percent.
They tended to travel more before 11:00 a.m. and after 3:00 p.m., although resting still composed at
least 45 percent of the activity during every hour (fig. 18).
These results generally agree with the statement by Mech (1966a) that wolves on nearby Isle Royale
tend to rest about 11:00 a.m. and begin traveling again about 4:00 p.m. However, it does appear that
the Minnesota wolves spend much more of the day resting than do the Isle Royale animals. The
difference may be caused by the difference in pack sizes studied. The Isle Royale pack of 15 to 16
may have had to travel more to find enough food to feed all its members than did the lone wolves and
pack of five in the present study.

Figure 17.—Percentage of time spent by
radiotagged wolves in various types of activity

throughout the day, from December through
April.

Figure 18.—Generally the wolves rested during most of the
day. (Photo courtesy of L. D. Mech.)

Movements and Range

Wolf movement is greatly hindered by deep, soft snow, so during winter travel, wolves frequently use
areas where they sink into the snow the least. In our study area, frozen waterways are used extensively
where possible, just as reported by Stenlund (1955). Where few lakes or rivers exist, wolves follow
railroad beds and logging roads, often soon after a plow or other vehicle has driven on them. In cutting
cross country through deep snow, wolves travel single file and tend to stick to windblown ridges and
to trails of deer and moose. Wolves that have ranges small enough to cover in a few days form a
network of their own trails, which they can maintain merely by traveling regularly over them. Packs
on Isle Royale depended a great deal on such a system of trails (Mech 1966a), and so did Pack No.
1059 in our study area.
Wolf packs can travel up to 45 miles in a day but it is usually larger packs that do so (Stenlund 1955,
Burkholder 1959, Mech 1966a, Pimlott et al. 1969). In our study area we sometimes saw evidence of
long moves by large packs along strings of lakes and waterways. However, most of our movement
data pertain to lone wolves and a pack of five. The daily travel of these animals was usually much less
than that reported for large packs.
Our radiotracking data provide an index to the extent of travel for each wolf rather than the actual
amount of travel, for it is based on straight line distances between consecutive points at which an
animal was found. This measure will be referred to as the "net daily distance."
Much variation was found in the net daily distances of wolves, with the longest ranging from 4.5 miles
for 1057 to 12.8 for 1055 (table 3). The mean net daily distance for each animal, excluding days with
no net movement, varied from 1.5 to 3.6 miles. The movements of these wolves may have been
affected by the snow depth and penetrability, for mean and maximum net daily distances suddenly
increased for all animals between February 23 and 28, when snow penetrability had decreased to a
point where walking wolves would be expected to sink in only about 6 inches (table 4). Other possible
explanations for the wolves' sudden increase in movements will be discussed below.

The straight line distances traveled between consecutive weekly locations (called the "net weekly
distances") showed a similar variation (table 3). The maximum net weekly distance for each wolf
varied from 4.6 miles for 1059 to 49.0 for 1055, with means ranging from 2.9 to 15.6 miles for the
same wolves. No doubt 1059's net weekly distances were relatively short because her total range and
that of her pack were much smaller than those of the other wolves.
It is difficult to obtain comparable measures of the extent of the ranges covered by each of the
radiotagged wolves because their patterns of travel varied so much. Thus the figures given in table 5
should be regarded only as gross indicators of the minimum range of each animal. The area figures are
especially deceiving in the case of 1055, for she had a horseshoe-shaped range, much of which
apparently was not used.

Table 3.—Straight line distances (miles) between consecutive locations of radiotagged wolves

Net daily distances Net weekly distances
Mean net
Mean
Mean distance
Wolf Days Days no net
Days net per day Weeks
number data net Range distance Range
movement distance excluding data
obtained movement per
per day days of no
week
movement
Number Number Percent Number Percent Miles Miles Miles Number Miles Miles
0.0- 1.0-
1051 54 13 24 41 76 2.0 2.6 22 12.7
12.0 46.0
0.0- 0.0-
1053 37 20 54 17 46 1.0 2.1 23 6.3
05.0 23.6
0.0- 1.7-
1055 46 7 15 39 85 2.9 3.6 21 15.6
12.8 49.0
0.0- 0.0-
1057 29 11 38 18 62 1.0 1.5 15 4.6
04.5 31.0
0.0- 0.0-
1059 26 1 4 25 96 2.5 2.6 18 2.9
05.6 04.6

Nevertheless, one major piece of information is obvious from the figures: 1059's pack of five wolves
had a much smaller range than any of the other uninjured animals—approximately 43 square miles
when figured by the minimum-area method (Mohr 1947). The next smallest range was that of 1051
(excluding the area of his later dispersal—see below), which was some seven times the size of the
pack's range.

Table 4.—Straight line distances (miles) traveled between consecutive days ("net daily distance") by
radiotagged wolves in northeastern Minnesota during February 1969

Wolf Mean net daily distance Greatest net daily distance
number Feb. 1-23 Feb. 23-28 Feb. 1-23 Feb. 23-28
1051 1.1 3.5 2.3 4.8

1053 0.7 2.5 3.0 5.0
1055 2.7 6.2 8.0 12.8
1057 1.0 1.5 4.0 4.5
1059 2.2 3.1 4.0 5.6

There is little published information on the movements and ranges of lone wolves with which to
compare our data. Mech (1970) summarized information regarding ranges of packs. Reported ranges
varied from 36 square miles for a pack of two wolves in Minnesota (Stenlund 1955) to 5,000 square
miles for a pack of 10 in Alaska (Burkholder 1959). Considering only data based on intensive study in
the same general region (Minnesota, Isle Royale, and Ontario) as our study area, the largest range
reported was 210 square miles for a pack of 15 to 21 wolves on Isle Royale (Mech 1966, Jordan et al.
1967). On a per-wolf basis, the ranges in this region varied from 6 to 28 square miles per wolf. Our
pack of five with its range of 43 square miles would have about 9 square miles per wolf.
A more accurate assessment of the ranges of the radiotagged wolves requires an individual discussion
for each.
No. 1051.—The range of 1051 was composed basically of three distinct areas (fig. 19). Within 10 days
after being released, the wolf left the general area of his capture (Area A near Isabella Lake) and
traveled to Area B along Highway 1, some 17 miles to the southwest. From December 9 to January 4
wolf 1051 remained in Area B, which covers about 45 square miles. Between January 4 and 6 he
returned to Area A and stayed in 13 square miles until February 3. Between February 3 and 5 he
shifted to Area C east of Snowbank Lake, 11 miles northwest of Area A. He remained in that 16-
square-mile area until February 25, then suddenly left and headed 8 miles to the northeast.

Table 5.—Extent of ranges used by radiotagged wolves

Area[9] of
Wolf Greatest Greatest Total intense use
number length width area[9] (before late
Feb.)
Miles Miles Sq. miles Sq. miles
13 (Location
A[11])
1051[10] 28.5 13.6 318
45 (Location B)
16 (Location C)
1053 31.1 22.0 392 31
1055 55.4 24.9 997 40
1057 32.3 3.8 77 14
1059[12] 8.4 8.0 43 39

FOOTNOTES:
[9] Minimum area method (Mohr 1947).
[10] Before dispersal.

[11] See text and figure 19.
[12] Pack of five.

Figure 19.—Locations and range of wolf 1051. Lines
are NOT travel routes; rather they merely indicate
sequence of locations. Only selected lakes are shown.

From February 26 until April 24 the movements of 1051 were strongly indicative of dispersal (fig.
20). His average weekly straight line move during that period was 25 miles (compared with 6 miles
per week before this period), and until March 14 he maintained an almost straight south-southwest
heading to a location west of the town of Castle Danger. After that the animal traveled a series of
northwest-southwest alternations that on April 3 took him east of Big Sandy Lake to a point 129 miles
southwest of where he had begun the dispersal. There he remained for about 2 weeks, but between
April 17 and 24 he traveled 26 miles northwest. We last saw him at 3:30 p.m. on April 24 heading
northwest through a swamp 15 miles southeast of Grand Rapids, approximately 122 miles from where
he had started. The total of straight line distances between 16 consecutive pairs of locations taken at
intervals of from 1 to 8 days was 226 miles, which is the minimum distance the wolf traveled during
his dispersal.

Figure 20.—Dispersal of wolf 1051. Lines merely indicate sequence of
locations. Only selected lakes are shown.

We observed 1051 for distances of up to 5 miles during these travels; he maintained a steady trot that
seemed faster than usual, and he appeared intent on heading in a straight line. He did chase deer
during his travels, and twice was seen feeding on carcasses. In the area where he remained for about 2
weeks, he was twice seen closely associated with another wolf. This relationship will be discussed
later.
An extensive search was made for 1051's signals on May 2 in an area of at least 50 miles radius from
his last known location, but it was unsuccessful. On each subsequent tracking flight, the wolf's
frequency was also monitored with no success. Possible explanations for the loss of the signal from
this wolf include the following: (1) premature expiration of the transmitter, (2) capture of the wolf and
breakage of the transmitter, (3) loss of the exposed antenna and consequent reduction of range, and (4)
travel of the wolf out of range of the tracking aircraft.
During 1051's travels a number of interesting events took place:
Nov. 27, 1968—Captured and radiotagged
Dec. 4, 1968—Crossed road in front of tracking truck
Dec. 8, 1968—Moved to Area B
Dec. 9, 1968—Surprised on the ground at distance of 35 feet

Dec. 18, 1968—Chased by loggers with axes
Dec. 25, 1968—Almost shot by trapper who saw collar and withheld fire
Jan. 5, 1969—Returned to Area A
Jan. 13, 1969—"Bumped" twice on logging road by loggers in auto but no apparent
injury
Feb. 4, 1969—Moved to Area C
Feb. 26, 1969—Began long-range southwest movement considered to be dispersal
Mar. 14, 1969—Seen feeding on old carcass within 200 yards of houses, dogs, and a
man walking
Mar. 27, 1969—Chased two deer across 4-lane State highway 53
Apr. 3, 1969—Found with another wolf at point farthest south in his range
Apr. 24, 1969—Last contact with this animal; was seen traveling NW
Wolf 1053.—This wolf was basically a scavenger who subsisted for long periods on the remains of old
carcasses. She was known to have visited the remains of at least four deer and three moose, and she
stayed near one moose carcass from February 8 to 20, at least during the day. Between her date of
capture, December 10, and February 28, 1053 traveled about in an area of 31 square miles in the
Arrow Lake-Maniwaki Lake region (fig. 21).

Figure 21.—Locations and range of wolf 1053. Only selected lakes are
shown.

Between February 28 and March 6 she suddenly moved 13 miles to the east-southeast near the Sawbill
Trail, and during the next week she traveled a straight line distance of 24 miles southwest to a point
southeast of the town of Isabella. Her subsequent travels eventually took her over a much larger area.
Before February 28, 1053's average weekly straight line distance was 2 miles, but after that date it
increased to 11 miles.
Wolf 1055.—The range of this animal from January 5, when she was captured, to February 23 covered
about 40 square miles near Stony Lake, Slate Lake, and the Jack Pine Lookout Tower (fig. 22), and
her mean weekly distance was 4 miles. Between February 23 and 24, however, she traveled 13 miles
northeastward, the beginning of a series of long moves. By March 5, 1055 had reached Crescent Lake,
a point 39 miles east-northeast of her previous area of intensive use. She then gradually headed back
toward the west and south during the next 10 days and within the next month repeated this pattern.
When her signal was heard last on May 30, 1055 was near Martin Landing in the center of her range.
Her mean net weekly distance after February 23 had increased to 22 miles.

Figure 22.—Locations and range of wolf 1055. Only selected lakes are
shown.

Wolf 1057.—The movements of 1057 cannot be considered normal because freezing of a front foot
prevented her accompanying the pack of which she was a member. Nevertheless, even data from an
abnormal animal can provide some information. On January 13, 5 days after capture and release on
Red Rock Lake, 1057 was located 4 miles from the capture point with a pack of 10 other wolves. She
was limping and fell behind when they moved. Five days later she was again seen with the pack 12
miles away between Knife Lake and Kekekabic Lake. She then remained in about 14 square miles of
that general area through April 17 (fig. 23).

Figure 23.—Locations and range of wolf 1057. Only selected lakes are
shown.

Suddenly on April 24, 1057 was found in Ontario some 31 miles northeast of her location of the
previous week. That was the last time we heard her signal even though on May 2 we scanned an area
with a radius of 35 miles from her last known location and listened for her signal during every
subsequent flight.
Wolf 1059.—This animal was a member of a pack of three to five wolves (see next section). The
movements of the group varied little and were concentrated in the August Lake, Omaday Lake, and
Keeley Creek area in about 43 square miles (fig. 24). Contrary to animals 1051, 1053, and 1055, this
pack did not suddenly begin a series of longer weekly movements in late February. Both before and
after February 28, the average weekly straight line movement of the pack was just less than 3 miles.

Figure 24.—Location and range of wolf 1059 and pack. Only selected
lakes are shown.

Probably these animals did begin traveling more in late February, for their net daily distances did
increase at that time along with those of the other wolves (table 4). However, the increased travel took
place within the restricted area of the pack's usual range rather than in new areas as occurred with the
other wolves.
Because 1059 was later found to have bred and carried five fetuses, her movements during whelping
season (late April and early May) are of interest. Her locations on both April 24 and May 2 were
within 250 yards of each other, which might indicate that she was denning. On May 9, however, she
was 2.5 miles east of these locations, on the 17th and 21st was 2 miles west of them, and on the 30th
was 3 miles north of them.
In early January 1970, Wolf 1059 was killed by a trapper in the southeast corner of her pack's 1969
range.
Summer locations.—Signals from only 1053 and 1059 were heard during summer, and then tracking
attempts were made only on June 28, July 29, and August 29. Locations for 1053 on those occasions
were near Kelly Landing and Isabella Lake, within her previous range. Wolf 1059 was found each
time within 2 miles outside of the southwest corner of the pack's winter and spring range.

Wolf Associations, Social Behavior, and Reproduction

In our study area, population units of wolves exist as both single animals (lone wolves) and packs. In a
total of 77 observations, lone wolves constituted 32 percent of the sightings (fig. 25), with packs of

from 2 to 13 members making up the remainder (Table 1). On the basis of the number of wolves seen,
rather than the number of observations, lone wolves accounted for only 25 (8 percent) out of 323.

Figure 25.—Only 8 percent of the wolves observed were lone wolves.
(Photo courtesy of L. D. Frenzel.)

These figures compare favorably with reports in the literature as summarized by Mech (1970). In five
areas studied, lone wolves made up from 24 to 60 percent of the observations of population units, and
from 8 to 28 percent of the wolves seen. In our study area during 1948 to 1953, lone wolves
constituted 43 percent of the observations and 15 percent of the wolves (Stenlund 1955).
The average size of the population units observed during our study (total number of wolves seen
divided by the number of observations) was 4.2, which is significantly larger (95 percent level) than
the average seen in this area (2.8) from 1948 to 1953. This is also larger than that reported from any
other area of comparable size (table 6).

Table 6.—Mean sizes of wolf population units reported from various areas

Mean size
Largest Authority
of
Area Observations Wolves pack calculated
population
size from
unit
Number Number
Kelly
Alaska 310 1,041 3.4 12
1954

R. A.
Alaska 1,268 4,823 3.8 21
Rausch[13]
Pulliainen
Lapland 118 311 2.5 12
1965
Pulliainen
E. Finland 460 984 2.1 12
1965
Stenlund
Minnesota 112 318 2.8 12
1955
Present
Minnesota 77 323 4.2 13
study

FOOTNOTES:
[13] R. A. Rausch. Personal correspondence to L. D. Mech, 1967.

The largest pack seen in our study area included 13 members, and there apparently were at least two
such packs. Although larger packs than this have been reported, any group containing more than 8 to
10 members is unusually large (Mech 1970).
Wolf sociology is a complex subject and is still not well understood, so the following detailed
observations of the associations between our radiotagged wolves and others are given. Associations
are defined as relationships in which two or more wolves relate in a close, positive manner.
As mentioned earlier, 1051 may or may not have been associated with other wolves when he was
captured. However, although this animal was observed 55 times throughout winter and spring, only
twice was he seen associating with another wolf. Probably the same individual was involved each
time, because the location was about the same (the vicinity of the juncture of Aitkin, Carlton, and St.
Louis Counties).
The first occasion was on April 3. Wolf 1051 in the previous week had moved 46 miles straight line
distance from the northeast. He was then observed lying peacefully within 15 feet of another wolf near
a freshly killed deer. The very proximity of the two animals implied a positive relationship. On April
7, 10 and 14, 1051 was seen 1 mile, 10 miles, and 8 miles from the kill and was alone each time.
However, on April 17, 1051 was back in the general vicinity of the kill, and he and another wolf were
resting on an open hillside about 100 feet from each other. As we descended for a closer look, the
smaller animal arose and headed to the larger, presumably 1051 because he had not been disturbed by
the aircraft. The larger wolf did not arise for several seconds, but eventually followed the other into
the woods. No tail raising or other expressive posturing was seen in either wolf. One week later 1051
was 26 miles northwest of the kill traveling alone.
Wolf 1053 was never seen less than 80 yards from another wolf, and there was no evidence that she
ever associated with a conspecific. Even when she was seen 80 yards from the other wolf, both were
resting, and when the strange wolf left, 1053 made no attempt to accompany or follow it.
No. 1055 apparently had been traveling with another wolf when caught on January 5, and tracks
showed that the individual had remained near her until we arrived to handle her. Tracks found on
January 7 and 10 suggested that 1055 was with another animal, but that animal was not seen during

any of the six times 1055 was observed through February 1. However, from February 5 to 19, 1055
was with another wolf on eight of the 12 times she was seen. The two animals were observed resting,
traveling, hunting, and feeding together. On February 20, and thereafter, 1055 was alone all 14 times
she was seen.
It is possible that 1055's associate was killed between February 19 and 20. About March 6, a 63-pound
male wolf pup was found dead (by Mr. Charles Wick, USDA Forest Service) within about 50 feet of a
highway and less than a mile from where 1055 and her associate were seen on February 19. Because
of the snow conditions, it was judged that the wolf had been killed (probably by an automobile)
sometime in February.
Wolf 1057, whose foot froze during capture, was a member of a pack of 10 to 13 wolves, and was
seen with the pack on January 13 and 18. After that she was usually found alone, although on at least
five occasions she was with one or more wolves:

No. of
Period Associations
observations
10 other
Jan. 13 1
wolves
Jan. 14-
1 None
17
10 or 11 other
Jan. 18 1
wolves
Jan. 19-
2 None
29
Jan. 30 1 1 other wolf
Jan. 31 to
2 None
Feb. 2
Feb. 3-4 2 2 other wolves
Feb. 5 1 1 other wolf
Feb. 6-13 6 None
Feb. 14 1 3 other wolves
Feb. 15-
5 None
22
10 to 13 other
Feb. 23 1
wolves
Feb. 24
6 None
to Apr. 24

February 23 she was with the pack at a kill in her usual area, and although the pack left that night,
1057 remained near the kill the next day. Presumably this animal would have traveled with pack if she
could have.
No. 1059 was part of a pack that included three to five members (fig. 26). From January 25, the first
time she was observed after release, through April 2, the animal was seen 19 times with two other
wolves, eight times with at least three others, and eight times with four others. She was never seen
alone until April 17; both times after this when she was seen, May 9 and 21, 1059 was also alone.
Some insight into the fluctuating size of this pack was obtained on February 27 when the five animals
were followed for 2 hours. During that time two members (one of which was larger than the other)

often lagged behind the other three by as much as
a mile. These two romped and played
considerably, with one carrying a stick or a bone
part of the time. Eventually they caught up again
to the other three. The behavior of the two lagging
wolves would be consistent with the hypothesis
that they were either pups or a courting pair of
adults. In either case, they seemed to be an actual
part of the pack even though they temporarily
traveled separately.
The fact that 1059 was observed traveling alone
three times from April 17 to May 21 may be
further evidence that the pack had a den in the area
at that time. The presence of a den allows
individual pack members to venture off singly and
return each day to a known social center, as Murie
(1944) observed, so they do not need to travel with
each other to maintain social bonds. Wolves in our
area breed during the latter half of February (see
below), and the young should be born in the latter
half of April. Since dens are prepared a few weeks
in advance (Young 1944), pack members might be
expected to begin traveling singly in mid-April.
Some information on social relations within our
radiotagged pack of five was also obtained. One of
the members could often be distinguished from the
others by its reddish cast and this individual Figure 26.—One of the radiotagged wolves was
appeared to be the pack leader or alpha male a member of this pack of five. (Photo courtesy
(Schenkel 1947). In urinating, this animal lifted of L. D. Mech.)
his leg, a position seen almost exclusively in
males. Except for only two temporary occasions, this animal always headed the pack, which usually
traveled single file. The second wolf in line generally was noticeably small, possible a female, and the
third wolf was twice identified as 1059 on the basis of sightings of her collar.
The leader often gained a lead on the other wolves, especially during a chase (see below), much as
reported for a lead wolf on Isle Royale (Mech 1966a). Upon returning to the lagging members of the
pack, this animal usually held his tail vertically, an expression of social dominance (Schenkel 1947).
On two occasions he led chases against strange wolves and demonstrated the highest motivation (see
below).
The leader was also the most active in his reactions when scent posts were encountered. Because the
function of scent-marking behavior is still unknown, it is important that detailed descriptions of the
natural behavior of free-ranging wolves around scent posts be made available (fig. 27). Thus the
following excerpt from field notes by Mech dated February 27, 1969, is presented:
"When they [the three wolves] came to a small frozen pond, where the wolf trail
[which they had been following] branched and there were some packed down areas,
they became quite excited [fig. 28]. This was especially true of the reddish wolf. He
nosed several spots, and scratched around them. Usually his tail was vertical. He

defecated at one spot, and right afterwards another
wolf did. After about 2 minutes that pack went on.
"About
15
minutes
later the
2
'satellite'
wolves
arrived at
this spot,
hesitated,
nosed
around
but
continue
d on after
less than
a minute.
Figure 27.—Feces, urine, and
scratching in a conspicuous spot "The
indicate a wolf "scent post." (Photo three Figure 28.—A pack of wolves
courtesy of L. D. Mech.) wolves investigating a scent post. The raised
meanwhi tails indicate their excitement. (Photo
le came courtesy of L. D. Mech.)
to a junction of 2 logging roads. There they
nosed around, scratched, and acted much as described above. Again the reddish wolf
was most active and had its tail up.
"When the last 2 wolves came to this spot, they nosed around, ran back and forth, and
1 defecated. They then headed on a different branch of the trail than the first 3 had
gone on just 10 minutes before.
"The first 3 wolves meanwhile were running along a logging road but eventually they
circled and one other than the reddish one headed across a swamp toward the last 2.
Then the reddish one and the other followed this one, and they met the last 2 on a
ridge. There was the usual tail wagging, then all headed off together in a new direction.
They passed the first scent post again and there was some nosing by the reddish wolf
but little hesitation.
"When they traveled, one wolf lagged behind by 150 yards. The wolf just ahead of it
had its tail vertical part of the time, as did the reddish leader.
"Soon the pack came upon another area packed with wolf tracks on a pond. There they
followed every little trail, nose to the ground, wagged tails, grouped together often,
chased each other, rolled over, etc. for 6 minutes. The reddish animal had tail up most
of the time.
"The wolves continued on, and we left them about 1 mile S.W. of the S.W. arm of Bald
Eagle Lake [at 6:05 p.m.]."

Unfortunately it was not known whether the trails that the wolves were following were their own or
those made by other wolves.
Significant aspects of the above observation are (1) the spirited initiative of the leader, (2) the amount
of time spent in scratching, urinating, and defecating, (3) the decision of the last two wolves to take a
different route from that of the first three even though their goal seemed to be to catch up to the first
three, and (4) the fact that the scent posts were located at trail junctions. In the last regard, we often
noted from the ground that wolves urinated at the junction of newly formed human trails heading
perpendicularly from roads they were following.
Copulation in wolves was only observed once during our study, on February 19, 1969. Two members
of a group of four were seen coupled for 2 minutes on Kekekabic Lake. On Isle Royale, which is at
the same latitude, copulations were witnessed on February 21, 24 and 27 (Mech 1966a).
On April 17, a den west of Big Moose Lake known to have been used at least intermittently for 13
years was seen from the air to have fresh activity of some kind in the snow in front of it, and on April
24 we saw a wolf at the mound. A few days later, two local human residents unaware of our interests
approached this den and looked in. An adult wolf, presumably the bitch, leaped over their heads and
fled the area. The men then dug up the den and removed six pups whose eyes had not yet opened.

Intraspecific Intolerance and Indifference

Instances of chasing or attack by a pack of wolves on conspecifics not a part of their group have been
described by Murie (1944) and Mech (1966a). Observations of such behavior are important in trying
to determine conclusively whether or not wolves are territorial. Pimlott et al. (1969, p. 75) wrote "It
still is not clear, however, whether or not their use of range should be defined as territorial." Mech
(1970) summarized the available evidence for territoriality in wolves and postulated that it may be
spatiotemporal such that packs might avoid each other at any particular point in time but over a long
period might cover the same area at different times. A number of our observations are pertinent to this
question, for we have evidence of both, tolerance and intolerance between population units of wolves.
Two direct cases of intolerance were observed, both involving the radiotagged pack and other wolves
within the usual range of the pack. Following is a direct quote from the field notes of Mech:
"Feb. 7, 1969—about 11:30 a.m.—aerial and visual—1059 and 2 other wolves
traveling overland about halfway between Heart L. and August L. (R10W-T61 N. Sect.
17 center). They were traveling quickly and intently along a fresh wolf trail, with a
lighter reddish individual in the lead. The other 2 animals were darker colored, and one
of them was smaller than the other. One of them must have been 1059.
"We soon found that about half a mile ahead of the pack was a dark wolf hurrying
away from the three. This animal often looked back and ran whenever it encountered
good running conditions. It soon became obvious that the pack of 3 was chasing this
individual. Because it [the lone wolf] often broke its own trail, the pack gradually
gained on this animal. The single wolf flushed a deer which ran when the wolf was
about 75 feet away and floundered in the snow, but the wolf continued hurrying on by.
"Although the deer ran only about 50 yards and stopped, the pack of 3 also hurried on
by. The single wolf flushed another deer, ignored it, and continued by, as did the pack
of 3. The chase continued for 2 miles as we watched, into the N.E. corner of Sect. 18
and then into the N. Central part of Sect. 8, and the pack got to within 150 yards of the
single wolf.

"However, at this point, the 2 darker members of the pack had fallen about 100 yards
behind the lead one. The lead animal stopped and waited for them, as it had done a few
times before. It then turned around and headed back to these animals. When they met,
the reddish animal's tail was held vertically and there was much tail wagging by all for
about 1 minute. Then all animals lay down for a minute and then went up on a knoll.
There was much activity and 'playing' on the knoll. (12:10 p.m.)
"The single wolf continued running and looking back for at least another mile. We left
at 12:21 p.m.
"At 4:07 p.m. we saw a single wolf running across a small lake and looking behind it
about 8 miles N.W. of these animals. The creature behaved the same as the one being
chased today, and we wondered whether it could be the same animal."
On February 18, 1969, Ream made a similar observation, as follows (quoted from his field notes):
"Got visual sighting on 1059 with 3 other wolves at 11:55 about a mile west of
Omaday Lake and they were running along fairly fast on a trail. When we circled a
second time we saw 2 wolves curled up sleeping on a knoll ahead (south) of the
running pack. We then realized the running wolves were on the trail of the sleeping
wolves and when the pack of 4 with "red" in the lead was about 50 yards from the
knoll the 2 sleeping wolves jumped up and charged away in the opposite direction full
tilt, and split and went in 2 directions. When the pack reached the knoll they started off
on the trail of the wolf that headed N.E. and then changed and went after the one that
headed S.W. The reddish wolf was in the lead and really picked up the pace. Although
the reddish wolf seemed to gain on the chased one 3 or 4 times, the pack as a whole
couldn't catch up, even though the single was breaking trail. The reddish wolf, after
gaining, always stopped and waited for the others or went back to find them. They
chased this wolf for 2½ to 3 miles, all the way down to Highway 1 at a point 3.0 miles
from the lab [Kawishiwi Field Station, U.S. Forest Service]. There was a dense patch,
10-15 acres, of woods just before Highway 1 and we lost sight of the chased wolf for a
while and also the 4 when they entered it, but shortly we found that the chased one had
somehow doubled back and was heading N.E. again. The pack was apparently
confused for at one point 3 of them were wandering back and forth on Highway 1,
apparently looking for the trail of the chased wolf. Two of these paralleled the
Highway for a couple hundred yards and then stopped on top of a hill, apparently
resting. During this chase both the single wolf and the pack chased up deer from their
route of travel and didn't seem to pay much attention to them, even though some were
really floundering in the deep snow. We finally stopped watching all of this at 1:30
p.m. and proceeded on our rounds."
On February 21 we also saw a single wolf running and looking behind several times on Ojibway Lake.
Even when it saw a fisherman on the lake within 1/8 mile, it continued across to the opposite shore
seeming most intent on avoiding whatever was on its trail. Presumably it had also been chased by a
pack.
The cases of tolerance or indifference that we witnessed between wolves involved our lone animals.
On January 27, 1051 was at a kill he had made the day before, and another wolf was sitting within 100
feet looking toward the carcass. Eventually the unidentified wolf left without approaching any closer.
A lone wolf was also seen near 1053 in the general vicinity of a moose carcass, which probably both
were feeding on at different times. Three such observations were made, on February 10, 15, and 18;
and on February 21 another wolf was also seen near 1053 some 2.5 miles away from the moose

carcass. In all cases, the two animals were 80 to 200 yards apart in open country and must have been
aware of each other's presence.

Hunting, Killing, and Feeding Behavior

The primary prey of most wolves in our study area is the white-tailed deer (fig. 29), but some moose
(fig. 30) are also killed. We have examined the remains of six moose that were eaten by wolves, two
of which were killed by them (fig. 31). One was found on February 25, 1967, on Gillis Lake and the
other on March 7, 1969, on Twinkle Lake. These locations are within 3 miles of each other, suggesting
that a wolf pack in that area may be more accustomed to preying on moose than other packs. The other
four moose carcasses were found in other parts of the study area, but circumstances were such that the
causes of death of those animals could not be determined. A discussion of the details of wolf-moose
relations in our study area must await the collection of additional data.
The remains of 93 wolf-killed deer, and 49 probable wolf-
kills, were examined for age, sex, and condition and were
compared with a sample of 433 hunter-killed deer from the
same general area. The wolf-killed deer were generally much
older than the hunter-kills and had a significantly higher
percentage of jaw and limb abnormalities (see Mech and
Frenzel, page 35).
Until recently the only observations of wolves hunting deer
were those reported by Stenlund (1955) for northern
Minnesota. He described two reports of actual observations
and two reports of interpretations of tracks in the snow, all
successful hunts. Since that time several descriptions of
successful and unsuccessful hunts have also been published
(Mech 1966b, Rutter and Pimlott 1968, Pimlott et al. 1969,
Mech 1970). Nevertheless, many more observations must be
made before generalizations can be formed.
During the present study we were able to witness a number of
hunts from the air and piece together others based on tracks.
Figure 29.—The main prey of The following descriptions are quoted from the field notes of
wolves in northern Minnesota is the Mech:
white-tailed deer. (Photo courtesy of
L. D. Mech.) "26 January 1967. About 3/4 mile N.E. of Alice
Lake.
"Jack Burgess [pilot] and I were following a pack of
8 wolves, when at 4:15 they veered from their former line of travel, about 30°. They
were then about 200 yards from 2 deer. They began wagging their tails when about 175
yards from the deer. One deer, on the edge of a steep bank, was lying, but one was
standing about 75 yards N. of it in open hardwoods. The wolves continued toward the
latter deer.
"This deer remained standing in the same place until the wolves approached to within
about 100 feet of it. The lead wolf stopped, when that distance from the deer, and the
others caught up but also stopped when within about 25 feet behind the lead wolf. By
this time the deer, whose body was facing away from the wolves, had its head turned
back over its shoulder toward the wolves. The wolves and the deer remained

absolutely still while staring at
each other, 100 feet apart, for 1-
2 minutes, while we made
several circles.
"Suddenly the deer bolted, and
instantly the wolves pursued. I
am fairly certain that it was the
deer that bolted first, but could
be mistaken. The action was
almost simultaneous. The deer
headed toward the other deer
near the top of the high bank.
This animal had been lying but
had arisen when the wolves
were about 150 yards away.
"The lead wolf followed in the
deer's trail, but the others cut
toward the bank. This flushed
the second deer (near the edge
of the bank), which ran down
the bank. Meanwhile when the Figure 30.—Moose are also killed by wolves. (Photo
first deer reached the edge of courtesy of Allan Taylor.)

the bank, it headed due W.
along the top of it. Only
the lead wolf pursued this
animal. The other deer had
headed down the bank to
the S.E., and at least a few
of the wolves followed it.
"We could not watch both
deer, so we continued
following the first. The
deer had no trouble in
snowdrifts, but the wolf
was hindered by them. The
wolf followed the deer for
about 200 yards along the
top of the bank, and then
gave up after losing
ground. The wolf had run a
total distance of about 250-
275 yards. He then lay
down and rested.
Figure 31.—Only a few wolf-killed moose were located
during the study. (Photo courtesy of Laurence Pringle.) "We noticed at least 3
wolves stopped part way
down the bank in the trail
of the second deer. However, we did not see the remaining wolves or the second deer.

"Eventually (after about 5 minutes), these wolves joined the first, and all rested. At
4:25 p.m., one wolf started toward a third deer, which had been lying under a tree
while the former chase took place. The deer was about 150 yards from where the
wolves rested, and it had stood before the wolf started toward it. We could not see
whether the deer or wolf bolted first, but suddenly both animals were bounding away.
The wolf chased the deer about 125 yards and gave up after losing ground. The other
wolves followed slowly in its trail, and all assembled and rested. The deer continued
running for at least 1/4-mile."
"27 February 1969. 2 miles N. of August Lake.
"1059's pack of 5 was heading N.E. at 4:10 p.m. when they got to within 100 yards of
2 standing deer. The deer had been standing alertly in a shallow draw, and when at
least 2 wolves got to within 100 yards, they fled. The wolves began running after them.
"The deer were in snow up to their bellies and had to hesitate slightly at each bound.
But they ran fast. We could only see one wolf very much [of the time]. It was also
having a difficult time in the snow, and after a total run of about 250 yards (100 to the
deer's original location and 150 after the deer), the wolf lay on the snow and rested
about 10 minutes. The deer ran only about 200 yards more and stood alertly for the
next 20 minutes at least. The wolves then went on.
"27 March 1969. About 2 miles S.E. of Central Lakes, Minnesota.
"At 3:00 p.m. while we were following wolf 1051 by aircraft in above location, we
saw a deer running very quickly on top of the crusted snow and then stand and watch
its backtrail. About 1½ minutes later we saw 1051 running along the same route. We
did not see when the deer fled again, but saw it running about 100 yards from the wolf
and doubling back paralleling its original route. When the wolf got near the
approximate doubling-back point, he lay down and rested for about 5 minutes. The
deer continued fleeing for about 350 yards, stopped, and for several minutes faced its
backtrail. The wolf finally continued on in his original direction, giving up the chase.
"At 4:30 p.m.—1½ miles S. of Central Lakes, Minnesota—Wolf 1051 had come to
within 100 yards of [four-lane] Highway 53 and was hesitant to approach it. Several
cars were going by in both directions. Thus the wolf headed S. parallel with the
highway about 150 yards E. of it.
"Suddenly two deer, which we had noticed S. of the wolf earlier, fled across the
highway. The wolf soon got to the point where they crossed, hesitated about a minute
and then ran across. No cars came at that time.
"We could not always see the deer or the wolf when W. of the road because there were
several patches of evergreens. The wolf did head straight W. after crossing the road.
Then about 250 yards W. of this point we saw a deer come out onto an old woods road
which lay in a N.W.-S.E. axis. The deer ran N.W. on the road and then we saw the wolf
where the deer had come out onto the road. While the deer ran N.W., the wolf cut into
the woods to his right, N.E. We could not see it then but presumed it was running N.W.
paralleling the road.
"After the deer had run about 50 yards up the road, it also headed N.E. into the
evergreens. Within a few seconds it fled right back out and started S.E. down the road.
The wolf was about 50 feet behind it and began gaining.

"When the deer got back to where the wolf had headed into the woods from the road
before, it also headed N.E. into the woods. The wolf was then about 20 feet away and
the deer was headed N. around in a circle with the wolf closing in on the outside. The
wolf did not emerge from the evergreens for at least 15 minutes, nor did we see the
deer, so I presume the wolf killed the deer. [But see entry for April 1.]
"1 April 1969. Dan Frenzel and I searched the area described on March 27 for 1 hour
and found no sign of a kill. Old wolf tracks were seen, but only a single wandering
track. No concentration such as usually seen at kills. Best conclusion is that 1051 did
not kill the deer where seen from the air March 27."
We also saw 1055 and her associate actually kill a deer, on February 6, 1969, but we did not realize
what was going on and it happened so fast that we only saw a wolf rushing and biting at the front end
of the downed animal. The chase had to have lasted only a few seconds.
In addition to the above direct observations, we also were able to piece together from tracks in the
snow the chase and successful encounter between a single wolf and a deer in two instances. In the first
case, on January 25, 1967 (11:50 a.m.), we arrived at the scene (near Grub Lake, just N. of Snowbank
Lake) within an hour of the encounter, and the wolf was still feeding on the deer, which had been a
2½-year-old female. Mech examined the area from the ground and made the following observations:
"The deer had come S.W. down the middle of the lake at a fast walk, turned around,
backtracked a few yards and headed to the N.W. shore of the lake. Meanwhile a wolf
had come at a trot along the deer's track, but it had cut to the N.W. shore about 50
yards N.E. of where the deer had. When still on the ice about 15 feet from shore, the
wolf began running as evidenced by his long bounds. He continued running inland
about 50 feet from shore toward the deer. The deer had walked inland from the shore
and may have stood there about 25 feet from shore. Suddenly it had bounded away.
The bounding wolf track was in the same trail as the deer's for about 25 yards but then
it paralleled the deer's about 5 feet away on the inland side. After about 125 yards from
where the deer flushed, the deer was pulled down. It was not on its side but rather had
sunk into the snow in more-or-less of an upright position.
"Apparently the deer had just about reached the shore when the wolf noticed it, and it
detected the wolf. At this time the wolf must have been up the shore about 50 yards
where his tracks first showed he began bounding. There was no sign that the wolf had
spotted the deer on the lake and had tried to cut it off from shore by running inland
along the shore and then waiting for the deer to come inland. Once the wolf had begun
bounding, he continued until he pulled the deer down.... Sign showed that the deer
dropped within about 20 feet of where she had begun bleeding."
The second case involved a 5½-year-old buck, No. M-28, which had arthritis of his right hind foot and
probably had defective gait (see Mech and Frenzel p. 35). The attack took place on Basswood Lake on
February 2, 1967, and excerpts from field notes by Mech follow:
"A single wolf had killed this deer after chasing, following, or tracking the deer about
3.75 miles. The deer's last 350 yards was a fast walk—the tracks were one in front of
the other and about 2 feet apart, and there was no leaping or bounding. Same with the
wolf—a fast trot.
"Where the tracks came together, the deer apparently had fallen, but there was no
blood. From there, the deer dragged its feet or the wolf for about 25 feet and then went

down again. The wolf circled the deer, and for the next 150 feet, the 2 animals had
fought or scuffled and then the deer had gone down where we found it.
"The 4-mile persistence of this wolf—whether tracking, following, or chasing the deer
—is remarkable [compared with most chases] and makes me believe the wolf had good
reason to believe it could kill the deer."
Our observations of wounds on fresh kills confirm the following description by Stenlund (1955, p. 31)
of the location and manner of attack of wolves on deer: "No evidence of hamstringing of deer was
found on freshly killed carcasses, although the possibility does exist. Usually deer are run down from
behind, the wolf or wolves biting at the hind flanks and abdomen, or at the hind flanks and head
region simultaneously."
On each kill, all the flesh and much of the skin and bones were eaten, at least during the winters of
1966-67 and 1967-68. This was also true during December 1968 and much of January 1969. However,
during February and March 1969 when an unusual accumulation of snow had built up, most of the
kills were only partly eaten (see Mech et al., page 51). In previous years deer freshly killed by single
wolves were sometimes found with only a few pounds of flesh or viscera missing. However, in each
case the carcasses were almost completely cleaned up within a few days, often by packs to which the
single wolves may have belonged (Mech 1970).
Usually the first parts of a carcass to be eaten are the hams and part of the viscera from the coelomic
cavity. In one case where a wolf was interrupted while feeding it was apparent that the animal had
been stripping the omental fat from the carcass. This may be the wolf's favorite part of a deer, for the
stomach of one wolf that we examined in January 1967 contained nothing but such fat.
The average consumption and kill rate of deer by wolves has not yet been determined, but we have
some information bearing on the subject. Because our data were obtained during a winter of unusually
deep snow, and it was obvious that wolves were killing more deer than they could eat at the moment
(see Mech et al., page 51), our figures should be considered much higher than average. However, they
should be useful in that they probably represent the maximum kill rate not only throughout the year
but also throughout a period of many years.
By observing each of our radiotagged wolves whenever possible and noting whether or not it was
feeding on a kill, we learned that our wolves generally remained close to their kills for periods of from
1 to 7 days, depending on how recently they had eaten (fig. 32). Thus, when a wolf was found at a
new location each day, the assumption could be made that the animal did not currently have a kill.
We assumed that wolves
found at fresh kills (fig.
33) had made them
unless there was
evidence to the contrary
as with 1053, the
scavenger. When a wolf
was found at one
location for several
consecutive days but
could not be observed,
Figure 32.—Periods spent by radiotagged wolves and their associates we assumed it was
feeding on kills judged to be their own. This does not include periods feeding on a kill, since
whenever wolves were

when they were known to be feeding on carrion. observed remaining in
(Click for larger image.) the same location for
several days they were

seen feeding. Thus a range of possible number of kills per
wolf was determined, with the lower limit being the known
minimum and the upper limit the possible maximum. When
more than one wolf fed on a kill, as with the pack, the figures
were calculated on a per-wolf basis.
In this way we obtained data on a total of 468 wolf-days and
found a total kill of 35 to 48 deer (table 7). This averages out
to a kill rate of one deer per 10 to 14 days per wolf. The figure
varied considerably among individuals—1051 had the highest
rate of one kill per 6.3 to 7.2 days, and each wolf in 1059's
pack had the lowest rate (except for 1053, the scavenger) of
one deer per 14.0 to 18.0 days.
It is significant that the pack of five wolves had a lower kill
rate per wolf than did single wolves and pairs. This is
explainable because the ability of wolves to kill deer during
early 1969 was much greater than usual (see Mech et al., p.
51). Thus single wolves probably could kill deer just as easily Figure 33.—Radiotagged wolf
as could packs, but they did not need to share them. This (upper left) found at kill (lower
differs markedly from the situation on Isle Royale, where lone right). (Photo courtesy of L. D.
wolves usually feed only on moose remains left by packs Frenzel.)
(Mech 1966a, Jordan et al. 1967).
That lone wolves had more of a food surplus than those in the pack is confirmed by the figures on the
average number of days that the various wolves fed on kills (table 7). Wolf 1051 spent an average of
only 2.2 to 2.4 days feeding at each of his kills, whereas 1059's pack of five spent an average of 5.8 to
7.5 wolf-days at each kill. Further confirmation is found in the fact that even when most wolves were
leaving their kills partly uneaten, a pack of 8 to 10 wolves (probably that to which 1057 belonged) was
seen completely devouring a kill.

Table 7.—Kill rate of deer by radiotagged wolves and their associates

Wolf Wolf-days Wolf-days Wolf-days Wolf-days
Wolves Dates Kills feeding
number of data per kill[14] feeding
per kill
Number Number Number Mean Number Number Mean Number
1051 1 Nov. 26 to Apr. 3 101 14-16 6.3-7.2 33-40 2.2-2.4
1053[15] 1 Dec. 14 to Mar. 27 75 2-3 25.0-37.5 9-18 4.5-6.0
1055 1-2 Jan. 9 to Mar. 14 61 4-9 6.7-15.0 13-25 2.8-3.3
1057 1-13 Jan. 24 to Feb. 28 51 5-7 7.3-10.2 25-33 4.7-5.0
1059 5 Jan. 25 to Mar. 14 180 10-13 14.0-18.0 75 5.8-7.5

Summary 22 Nov. 27 to Apr. 3 468 35-48 [16] 145-181 [17]
9.8-13.4 3.8-4.1

Before Feb. 1 142 7-9 [18] 39-56 5.1-5.6
15.7-20.3
After Jan. 31 326 28-39 8.4-11.6 106-125 3.2-3.8

FOOTNOTES:
[14] Kill rate per wolf.
[15] Figures for this animal are so low because she was basically a scavenger.
[16] Average kill rate per wolf for all radiotagged wolves and their associates, derived by
dividing total number of wolf-days by total number of kills.
[17] Average number of days that each wolf spent at each kill, derived by dividing total
number of wolf-days spent feeding by the total number of kills.
[18] This figure probably is the closest to the actual kill rate during most winters.

Therefore it is probable that the kill rate per wolf for members of the pack of five is
much closer to the usual average winter kill rate. It can still be considered higher than
the usual winter rate, however, because this pack also was leaving some of its kills
partly uneaten.
A reasonable approximation of the average kill rate during most winters would be the
rate found for our radiotagged wolves before February 1, because the relations among
the wolves, the deer, and the snow during that period were not unlike those of most
winters. The average kill rate per wolf before February 1 was estimated at one deer per
15.7 to 20.3 days.
After this period, the rate increased to about one deer per 8.4 to 11.6 days, and an
estimated 50 percent of the available food was left uneaten (see Mech et al., page 51).
This implies that the kill rate during February and March was about twice as high as
usual. On this basis, the usual kill rate would be estimated at one deer per 16.8 to 23.2
days, which checks well with the rate found before February (one deer per 15.7 to 20.3
days). Thus we feel that an estimated kill rate of about one deer per 18 days per wolf is a
close approximation of the average kill rate for most winters. This is about 50 percent
less than the kill rate of one deer per 4 days estimated by Stenlund (1955) for two packs
of three wolves (one deer per 12 days per wolf). However, it compares favorably with
the actual kill rate of one deer per wolf per 17.6 days found for a pack of eight wolves in
Ontario.[19]
Once the average rate of kill is known, the average food consumption per wolf can be
calculated. The average deer (considering both fawns and adults) from the Superior
National Forest during winter weighs about 113 pounds (calculated from Erickson et al.
1961), and an arbitrary 13 pounds can be deducted from this for inedible portions. This
leaves 100 pounds of deer per wolf per 18 days, or 5.6 pounds per wolf per day. This
figure is much less than the 10 to 14 pounds estimated consumption rate for wolves

feeding on moose on Isle Royale (Mech 1966a). However, much variation can be
expected in an animal whose physiology must be adapted to a feast-or-famine existence.
Wolves can be maintained in captivity on 2.5 pounds of meat per day, and large active
dogs (Canis familiaris) require 3.7 pounds per day, so it is likely that the minimum daily
requirement for wolves in the wild is about 4.0 pounds per day (Mech 1970). This figure
agrees well with the estimated consumption rate for our study area.

FOOTNOTES:
[19] Kolenosky, G. B. Wolf movements, activities and predation impact on a wintering deer
population in East-Central Ontario. (Manuscript in preparation for publication.)

Relative Population Density

Censusing wolves in a 1.5-million-acre study area is a difficult task, and we have no
direct information on which to base a population estimate. However, some deductions
can be made about the relative population densities in our study area between the period
1948 to 1953 and the period of the present study, 1967 to 1969.
R. A. Rausch (1967a) hypothesized that the frequency of large packs is higher when
population density is high, and presented evidence supporting this idea. On this
assumption, a comparison of pack-size distributions between various periods can
indicate relative population densities between periods. The advantage of this method is
that it eliminates the usual type of year-to-year biases in wolf censuses such as might
result from differences in precise census route, type of aircraft, skill of observers, and
other conditions. Only a difference that would cause a bias in the size of the packs seen
would be of importance.
Therefore, we tested the difference in size distributions of population units between the
1948-53 study period and the present period (table 1), using a Kolmogorov-Smirnov
two-sample test (Siegel 1956). The average "pack" size in the earlier years was 2.8,
compared with 4.2 at present; thus pack sizes are significantly larger at present (95
percent level). This indicates that the population density from 1967 to 1969 may have
been higher than from 1948 to 1953. This apparent change may be attributable to a
reduction in snaring, trapping, and aerial hunting that took place between the two
periods as a result of changes in State game regulations.
A similar comparison between our observations from 1967 and those from 1968-69
(table 1) shows no significant difference between these years, so it appears that the
density of wolves in our area has remained about the same over the period of three
winters. This agrees with the results of several other studies summarized by Mech

(1970) in which wolf populations unaffected by man have been found to remain
relatively stable from year to year.

DISCUSSION AND CONCLUSIONS

The movements, behavior, and ecology of the wolves in our study area during winter are
variable, and are influenced considerably by snow conditions. This may explain the fact
that in late February 1969 wolves 1051, 1053, and 1055 suddenly extended their travels
and range (fig. F-34 and table 4).
However, increased travel may have resulted from other factors. For one thing, the
wolves apparently did not need to spend so much time hunting as before. Because of the
deep snow, the ability of wolves to capture deer increased, and the animals had a surplus
of food. Perhaps under such conditions wolves may use more of their energy for
traveling than for hunting.

Figure 34.—Net weekly (straight-line) distances traveled by three
radiotagged wolves.

In this respect it is interesting that 1051 moved right out of his area and traveled into
country that presumably was unknown to him. Wolves 1053 and 1055 each ventured
into an area that was almost devoid of deer and that even had few moose in it. Without

sufficient fat reserves in all these animals, it would seem disadvantageous for them to
have made these travels.
Evidently wolves can obtain enough food in much smaller areas than these three
animals used after February. Both 1059's pack of five and 1057 lived in relatively small
areas throughout the winter and seemed to survive well. Before late February, 1051,
1053, and 1055 did also. Thus some factor other than food must have influenced the
movements of these three animals from late February through April.
The fact that the increased movement began during the breeding season makes one
suspect a relationship between the two. One possibility is that the factors increasing the
hormonal flow associated with breeding in adults stimulate a hormone output in
immature or subordinate individuals that causes an increase in their movements. An
alternative is that the breeding behavior of resident packs involves the beginning of, or
an increase in, aggression toward neighboring nonmembers. This might force the lone
animals to shift about over large areas in avoidance of such aggression.
Whatever the cause of the changes in movements of these animals, the fact that the pack
used a much smaller area than any of the lone wolves may be of central importance in
trying to understand the organization of the wolf population. The following pieces of
information are also pertinent to such an understanding: (1) the pack, which can be
presumed to include a breeding pair (Mech 1970), chased other wolves in its area; (2)
the lone wolves, which apparently did not breed, were tolerant of, or indifferent to, other
lone wolves in their areas; (3) the ranges of the lone wolves overlapped considerably
(fig. 35); (4) the lone wolves seemed to avoid certain large areas that one might
logically think would have been visited by them (fig. 35); and (5) packs of wolves were
sometimes observed in these large areas (fig. 35).

Figure 35.—Locations of all radiotagged wolves and unmarked packs
observed during winter 1968-69, except dispersal of 1051 out of the
study area. Only selected lakes shown.
(Click for larger image.)

From the above information it can be hypothesized that the wolf population consists
basically of groups of breeding packs defending territories of limited size, with lone
wolves and other nonbreeding population units that are tolerant of each other shifting
about in much larger nonexclusive areas among these territories. The information from
Isle Royale (Mech 1966a, Jordan et al. 1967) is consistent with this idea, but the area of
that island (210 square miles) is too small to allow untested extrapolations to be made
about spacing in much larger wolf populations. Data from Algonquin Park, Ontario
(Pimlott et al. 1969) also strongly suggest this hypothesis. However, the packs studied
there could not be identified with certainty, and little information was obtained about
nonbreeding population units.
To test the proposed hypothesis with certainty, a larger number of identifiable breeding
and nonbreeding population units from the same general area must be followed during
at least one winter. This will be the main objective of our next study.

SUMMARY

During the winters of 1966-67, 1967-68, and 1968-69, aerial observations of timber
wolves (Canis lupus) were made in the Superior National Forest in northeastern
Minnesota, where the primary prey is white-tailed deer (Odocoileus virginianus). In 480
hours of flying during the study, 77 sightings involving 323 wolves were made. In
addition, during 1968-69, five radiotagged wolves and their associates were tracked via
receivers in aircraft for a total of 570 "wolf-days." Visual observations were made
during 65 percent of the times the wolves were located from December through April.
The average size of each population unit (including single wolves, pairs, and packs)
observed was 4.2, although packs of as many as 13 wolves were sighted. Radiotagged
wolves spent most of their daylight hours resting during winter, and when traveling,
hunting or feeding during the day, tended to do so before 11:00 a.m. and after 3:00 p.m.
Considerable variation was discovered in the movement patterns of individual wolves,
with straight line distances between consecutive daily locations ranging from 0.0 to 12.8
miles, and between weekly locations, 0.0 to 49.0 miles. A pack of five wolves used a
range about 43 square miles in extent, whereas lone wolves covered areas many times
this size. One animal in an apparent dispersal was tracked a straight line distance of 129
miles between extreme points.
A reddish male wolf was the leader of the pack of five and led two observed chases after
alien wolves in the pack's territory. This animal was also most active during scent
marking by the pack. Lone wolves were apparently indifferent to other wolves, and thus
exclusive areas, or territories, were not observed among lone wolves.
Hunts involving a total of seven deer were observed and described, and two successful
attacks on deer were interpreted from tracks in the snow. Wolves generally consumed all
the flesh and much of the hair and bones from kills, except during February and March

1969 when extreme snow conditions increased the vulnerability of deer to an unusual
degree. At that time kills were found that were partly or totally uneaten. The kill rate by
radiotagged wolves and associates during the winter of 1968-69, based on 468 wolf-
days of data, varied from one deer per 6.3 days to one per 37.5 days per wolf, with the
average being one deer per 10 to 13 days. The rate was much lower per wolf for
members of the pack of five than for lone wolves, and much lower before February 1,
1969, than after. The average rate of kill during more usual winters was estimated to be
about one deer per 18 days. This is a consumption rate of about 5.6 pounds of deer per
wolf per day.
Indirect evidence based on comparisons of pack-size distributions for different periods
indicates that the wolf density in the study area may have increased since 1953, but that
it has remained the same from 1967 to 1969.
On the basis of data presented in this paper, the following hypothesis about the
organization of the wolf population studied is proposed: The wolf population consists
basically of groups of breeding packs defending territories of limited size, with lone
wolves and other nonbreeding population units, tolerant of each other, shifting about in
much larger nonexclusive areas among these territories.

ACKNOWLEDGMENTS

This study was supported by Macalester College, the U.S. Bureau of Sport Fisheries and
Wildlife, the USDA Forest Service, the Minnesota Department of Conservation, and the
New York Zoological Society. Special thanks are due the following for their help and
cooperation with this project: Mr. J. O. Wernham, former Supervisor, Mr. L. T. Magnus,
Wildlife Biologist, numerous District Rangers, and other supporting personnel of the
Superior National Forest, Mr. J. T. Morgan, North Central Forest Experiment Station;
and Mr. S. E. Jorgensen and Mr. C. E. Faulkner, U.S. Bureau of Sport Fisheries and
Wildlife.
Drs. C. T. Cushwa, L. F. Ohmann, Catherine Ream, and D. G. Schneider aided in the
field work. Mr. W. W. Cochran provided advice and suggestions on the radiotracking
technique, Dr. U. S. Seal furnished the drugs and the advice on their use with wolves,
and Mr. R. Himes contributed significantly in the wolf trapping. Mr. L. Ringham,
Ontario Department of Lands and Forests, granted permission for research personnel to
radiotrack wolves crossing into Quetico Park, Canada. Numerous students from
Macalester College also contributed to the field effort.
Thanks are also due pilots Robert Hodge, Pat Magie, Ken Bellos, Don Murray, Jack
Burgess, and several others, who along with pilot-biologist John Winship, expertly flew
the aircraft used in the study.

This report was reviewed by the following biologists: Mr. G. B. Kolenosky, Dr. P. A.
Jordan, Mr. M. H. Stenlund, and Dr. D. L. Allen.
Mr. Wallace C. Dayton, Miss Elizabeth Dayton, and the Quetico-Superior Foundation,
all of Minneapolis, generously contributed funds to support Mech during the preparation
of the paper.

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to the wolf. Sci. Mon. 46: 323-336.
Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The ecology of the
timber wolf in Algonquin Provincial Park. Ont. Dep. Lands and Forests
Res. Rep. (Wildlife) 87, 94 p.
Pulliainen, E. 1965. Studies of the wolf (Canis lupus L.) in Finland. Ann. Zool.
Fenn. 2: 215-259.
Rausch, R. A. 1967. Some aspects of the population ecology of wolves, Alaska.
Amer. Zool. 7: 253-265.
Rutter, R. J., and Pimlott, D. H. 1968. The world of the wolf. 202 p. Philadelphia
and N. Y.: J. B. Lippincott Co.
Schenkel, R. 1947. Expression studies of wolves. Behaviour 1: 81-129.
(Translation from German by Agnes Klasson.)
Seal, U. S., and Erickson, A. W. 1969. Phencyclidine hydrochloride
immobilization of the Carnivora and other mammals. Fed. (Symp. Lab.
Anim. Anesthes.) Proc. 28: 1410-1419.
Siegel, S. 1956. Non-parametric statistics for the behavioral sciences. 312 p. New
York: McGraw-Hill.
Stenlund, M. H. 1955. A field study of the timber wolf (Canis lupus) on the
Superior National Forest, Minnesota. Minn. Dep. Conserv. Tech. Bull. 4, 55
p.
Young, S. P. 1944. The wolves of North America, Part I. 385 p. Washington, D.
C.: The Amer. Wildl. Inst.

AN ANALYSIS OF THE AGE, SEX, AND CONDITION
OF DEER KILLED BY WOLVES IN NORTHEASTERN
MINNESOTA
L. David Mech and L. D. Frenzel, Jr.
The selective effect of predation on prey populations is of significance in studies of
evolution and population dynamics. Selective predation can be an important agent in the
process of natural selection, and it influences the extent to which predators limit the
numbers of their prey.
One of the predators most commonly chosen for investigating the selective effect upon
prey is the wolf (Canis lupus). Because animals preyed upon by wolves generally are
large, their remains can be more easily located and examined. It already has been
established that in most areas wolves kill primarily young, old, and other inferior
members of such prey populations as Dall sheep (Ovis dalli), moose (Alces alces),
caribou (Rangifer tarandus), bison (Bison bison), and musk-oxen (Ovibos moschatus);
evidence for this generalization has been summarized by Mech (1970).
However, only recently has it been shown that this generalization may extend to
predation on the smallest hoofed prey of the wolf in North America, the white-tailed
deer (Odocoileus virginianus). Pimlott et al. (1969) demonstrated a difference between
the age structure of 331 deer killed by wolves during winter in Algonquin Park, Ontario,
and 275 deer assumed to represent the actual population in the same area. Whereas only
13 percent of the deer from the population at large were estimated to be more than 5
years old, 58 percent of the wolf-kills were in this age category.
We employed a similar analysis for deer killed by wolves in northeastern Minnesota, but
used a more refined aging technique and included comparisons of the age and sex
structures of various subsamples of wolf-kills. Whereas the Ontario research involved a
prey population unlimited by man, our work was carried out on both a hunted
population and on one relatively unhunted. Further comparisons were made between
deer killed during periods of normal snow conditions and those taken during unusually
high snow accumulations. The incidence of various abnormalities in wolf-killed deer
was also compared with that in hunter-killed animals.
The study was carried out in the Superior National Forest in northern St. Louis, Lake,
and Cook Counties of northeastern Minnesota (fig. 1), in conjunction with other aspects
of wolf research (see Mech et al. p. 1).

Figure 1.—The study area showing locations where wolf-killed and
hunter-killed deer were taken. Line arbitrarily separates the hunted area
from the wilderness area.
(Click for larger image.)

METHODS

The investigation began in February 1966 and continued through March 1969; the basic
objective was to examine as many wolf-killed deer as possible and compare their ages,
sex, and condition with a large sample of deer from the population at large in the same
area. Wolf-kills were examined only during December through March when they could
be found from the air. Aircraft ranging in size from an Aeronca Champ to a Cessna 206
were used to fly over frozen lakes at altitudes up to 2,000 feet to locate wolves (fig. 2),
wolf tracks, or kills (fig. 3). We often discovered kills by tracking a wolf pack.
During the winter of 1968-69 this method of finding kills was supplemented by
radiotracking five wolves and their associates via aircraft (see Mech et al., p. 1). The
latter technique resulted in increased discovery of inland kills.
A deer carcass was judged killed by wolves if the death had been recent, if tracks or
other sign indicated that wolves had fed upon it, and if no other possible cause of death
was discovered. Carcasses fed on by wolves but not clearly identifiable as kills were
labeled "probable" wolf-kills. Although the cause of death of the specimens in this latter

category could not be determined
with certainty, there was no reason
to believe other agents were
involved.
In addition to the wolf-kills
examined by project personnel, data
and lower jaws from deer judged
killed by wolves were contributed
by other biologists, game wardens,
forest rangers, and others whose
competence was known.
Nevertheless, if certain
identification of carcasses as wolf-
kills was not possible, the data were
relegated to the "probable" wolf-kill
category.
Whenever possible, kills discovered
from the air were examined on the
ground (fig. 4). Often only skeletal
parts remained, but soft parts were
also examined when available.
Femur marrow, heart, lungs, liver,
kidneys, reproductive tracts, and
omenta were usually inspected in
the field for fat, parasites, and
abnormalities, and the degree of
Figure 2.—Wolves were located from the air, subcutaneous back fat was also
usually on frozen lakes. (Photo courtesy of L. D. noted. Hoofs and lower legs were
Mech.) checked, and those showing
pathological conditions or
abnormalities were collected and
examined by the Veterinary Diagnostic Laboratory of the University of Minnesota. All
lower jaws found were collected, aged, and examined for dental abnormalities and
pathological conditions.
In November 1967 and 1968 hunter-check stations were operated on the study area (fig.
5), and deer bagged by hunters were field-checked for age (Severinghaus 1949) and
hoof abnormalities. As many lower jaws as possible were collected from field-checked
deer and other deer killed in the area for age determination and examination for
abnormal dentition.

An
assumptio
n was
made that
the age
structure
and
incidence
of
abnormalit
ies in the
sample of
hunter-
killed deer
would be
reasonably
representat
ive of
those in
the
population
at large, an
assumptio
n also Figure 3.—Wolf-kills were easily spotted from aircraft. (Photo courtesy
implicit in of L. D. Mech.)
a similar
comparison made by Pimlott et al. (1969). In this respect, the following statements by
Maguire and Severinghaus (1954, p. 109) about deer in New York State are pertinent: "It
may be concluded that, considering the open season as a whole, wariness does not
significantly distort the age composition of the [deer] kill in relation to that of the
corresponding wild population, except possibly for buck seasons of only 1 or 2 days
duration.... A reliable appraisal of the age composition of the kill by hunting may be
obtained through the operation of roadside checking stations." However, in critically
reviewing the present paper Severinghaus stated that in States such as Minnesota, with
fewer hunters and higher hunter success rates, age compositions of deer from checking
stations may not be the same as those of wild populations. Reviewers Peek and
Downing also made similar comments.
Nevertheless, for our comparison with wolf-killed deer it is not necessary that the
hunter-kill age structure be exactly representative of the age structure of the actual deer
population. All that is required is that there be reasonable agreement between the two.
The hunting regulations in our study area allow a 9-day period of taking deer of any age
or sex, and a single hunter may legally shoot as many deer as he and his party or

associat
es have
permits
for.
Thus
there is
no
reason
for
selectiv
e
Figure 5.—Information about
hunting,
hunter-killed deer in the study area
and we
was obtained through hunter-check
feel
stations. (Photo courtesy of L. D.
confide
Frenzel.)
nt that
the age
structure of the hunter-kill in our study area
Figure 4.—As many wolf-killed does basically represent that of the deer herd at
deer as possible were examined large.
from the ground. (Photo courtesy of
Two laboratory techniques were used for
L. D. Mech.)
determining the ages of deer from the lower
jaws or mandibles—a tooth replacement and
wear technique (Severinghaus 1949) and an
incisor-sectioning method (Gilbert 1966). The tooth-wear technique requires only the
molariform teeth but it is more subjective and inaccurate, particularly in older deer
(Ryel et al. 1961). Incisor sectioning requires only incisors and appears to be much
more accurate.
However, because the incisors had been lost from many of the wolf-kills, and because
the tooth-wear technique was used at checking stations, both methods were applied in
the laboratory. Mr. David W. Kuehn (1970) sectioned and aged the incisors. Fortunately
there was a sufficiently large sample of mandibles with molariform teeth and incisors
from both wolf-killed and hunter-killed deer to enable us to devise a table showing the
actual ages (based on incisor-sectioning) of each of the jaws assigned to various tooth-
wear classes. This table was then used to distribute the ages of specimens that contained
only molariform teeth. For example, because it was found that 37 percent of the jaws
aged 4½ years old by tooth wear were actually 5½ years old, we assigned 37 percent of
the incisorless jaws aged 4½ by tooth wear to the 5½-year category. Similarly, another
conversion chart comparing field age determinations of hunter-killed deer with ages
based on incisor sectioning of the same jaws was employed to distribute the ages of
field-aged, hunter-killed deer for which jaws or incisors could not be collected.

RESULTS

We flew a total of 480 hours during
this and related research, mainly
during January through March 1967
and December 1968 through March
1969; about one-third of this time
was devoted primarily to searching
for kills. Jaws were examined from
93 wolf-kills and 49 probable wolf-
kills.
Hunter-check stations yielded
information from 335 deer (fig. 6),
and data on 98 additional hunter-
killed deer were contributed by
other hunters. Incisors were
collected from 82 of 214 hunter-
Figure 6.—All hunter-killed deer examined
killed deer checked that were older
were checked for age. (Photo courtesy of L. D.
than yearlings; comparisons were
Frenzel.)
then made between ages of the deer
based on incisor sectioning and
those based on field checks using the wear method. Similarly, incisors were sectioned
from 195 wolf-killed and hunter-killed deer older than yearlings that had been aged by
the tooth-wear method in the laboratory, so that these two methods could be compared
(Kuehn 1970). (Note: incisor-sectioning is unnecessary for fawns and yearlings because
animals of these ages can be aged objectively by the progress of tooth replacement.)
Because age or sex distributions might differ in the various subsamples of deer
examined during this study, these parameters were compared in subsamples of both
wolf-kills and hunter-kills (table 1). No significant differences were found in the age or
sex structures between the known wolf-kills and "probable" wolf-kills, so these
subsamples were pooled and considered wolf-kills for all subsequent comparisons.
Three significant differences in sex ratio were found among the subsamples of wolf-
kills: (1) wolves killed more female fawns than male fawns, but more male adults than
female adults (table 2); (2) more of the adults killed in the hunted area were females,
while in the wilderness more males were taken (table 3); and (3) after January 1969,
when snow was unusually deep, 57 percent of the deer killed were females, compared
with only 38 percent before this date.

Table 1.—Results of statistical comparisons between various samples of deer kills
from northeastern Minnesota

Results of comparisons
Sample Sample Sample Sample Direction of
VS Age Sex
size description size description difference
structures[20] ratios[21]
Wolf-kills:[22] Wolf-kills:[22]
93Known 49 Probable Nonsig.[23] Nonsig. —
Dec. 1968-
42Jan.-Mar. 1967 83 Nonsig. Nonsig. —
Mar. 1969
66Male 61 Female Nonsig. — —
50Wilderness area 92 Hunted area Nonsig. Nonsig. —
More
Adult, hunted Sig., 99
41Adult, wilderness 64 — females in
area percent
hunted area
96 Lakes[24] 32 Inland Nonsig.[25] Nonsig. —
More
After Jan. Sig., 95
66Before Feb. 1969 77 Nonsig.[26] females after
1969 percent
Jan.
Sig., 95 More female
105Adults 22 Fawns —
percent fawns
Hunter-kills: Hunter-kills:
Field aged,
110 Field aged, 1967 225 Nonsig. Nonsig. —
1968
335Field aged 98 Lab. aged Nonsig. Nonsig. —
Lab. aged,
132Lab. aged, males 79 Nonsig. — —
females
Field aged, Sig., 95 More male
89Field aged, fawns 246 —
adults percent adults

Sig., 99 Older deer in
433Hunter-kills 142 Wolf-kills —
percent wolf-kill
Wolf-kills
Hunter-kills Sig., 99 Older deer in
321 118 excluding —
excluding fawns percent wolf-kill
fawns

FOOTNOTES:

[20] Kolmogorov-Smirnov two-sample test (Siegel 1956).
[21] Z test (Downie and Heath 1959).
[22] Because test showed no significant differences in age or sex structure between sample of
known wolf-kills and probable wolf-kills, these were combined for all subsequent tests and
the pooled sample considered "wolf-kills."
[23] At 95 percent level or greater. (NOTE: Lack of a significant difference does not prove
that no difference exists. Rather, it means only that the available evidence does not allow the
positive conclusion that a difference does exist.)
[24] Wolf-kills found on lakes were compared with those located inland because of the
possibility that kills on lakes may not be representative of kills in general.
[25] Sample too small for test, but no apparent difference.
[26] No significant difference in entire age structures. However, when the percentage of
yearlings is compared between the two groups, the difference is almost significant at the 95
percent level.

Table 2.—Sex ratios of hunter-killed deer and wolf-killed deer from northeastern
Minnesota

Age Hunter-killed deer Wolf-killed deer
Percent Percent Percent Percent
Number Number
male female male female
Fawns 108 50 50 22 41 59
Adults 315 68 32 105 54 46

In the comparisons of the subsamples of hunter-kills, the only statistically significant
difference found was that the adult subsample had a higher proportion of males than the
fawn subsample. No significant difference was found in the age structures of the
subsamples, so these were all pooled into a sample of 433 hunter-kills for comparison
with the wolf-kills. For the same reason, the entire sample of 142 wolf-killed deer was
used for a comparison with the hunter-killed sample.

Table 3.—Sex ratios of wolf-killed deer from wilderness areas and from hunted areas

Age Wilderness area Hunted area Total
Percent Percent Percent Percent Percent Percent
Number Number Number
male female male female male female
Fawns 4 0 100 18 50 50 22 41 59
Adults 41 71 29 64 44 56 105 56 44

Wolf-killed deer in our sample, with an average age of 4.7 years, were significantly
older (99 percent level) than hunter-killed deer, with an average age of 2.6 years. For
example, deer 5 years of age and older made up 48 percent of the wolf-kills but only 10
percent of the hunter-kills (table 4). The oldest hunter-killed deer in our sample was 9½
years old, but the oldest wolf-killed deer was 14½ (fig. 7).
Because of a possible bias against fawns in the method of collecting data from wolf-
kills (to be discussed later), the age structure of the sample of wolf-kills excluding
fawns was tested against that of the sample of hunter-kills excluding fawns. The result
once again was a highly significant difference between these two age structures (table
1).
As an additional test of the degree to which the age structure of the wolf-killed deer
might differ from that of the actual population, we compared our wolf-kill age structure
with the age structure of a hypothetical deer population. This was considered advisable
just in case the hunter-kill data were poorly representative of the age structure of the
actual deer herd. Several hypothetical age structures were constructed and compared
according to advice from Downing.[27] In all cases, the comparisons produced the same
basic results as the tests with the hunter-killed sample. An example of one comparison is
given in figure 7.
A further result obtained by aging the wolf-killed deer pertained to the young
individuals killed. The deciduous first incisors of fawns and the deciduous premolars of
yearlings are usually replaced with permanent teeth by December (Severinghaus 1949).
Of 24 wolf-killed fawns examined, however, three (13 percent) taken during January,
February, and March had not yet replaced their deciduous first incisors. Of the 13
yearlings found during this same period, nine (70 percent) had failed to replace their
deciduous premolars, and two (15 percent) had just replaced them (one deer killed in
February and one killed in March).

FOOTNOTES:
[27] R. L. Downing. Personal correspondence to L. D. Mech, October 2, 1969.

Table 4.—Age and sex distribution of deer killed by wolves and hunters in
northeastern Minnesota

Age Wolf-killed deer Hunter-killed deer
(years) Number of: Percent Number of: Percent

MalesFemales Unknown Total MalesFemales Unknown Total
Fawns 9 13 2 24 17 54 54 4 112 26
1+ 5 7 1 13 9 63 26 1 90 21
2+ 3 8 5 16 11 42 19 2 63 15
3+ 2 4 2 8 6 47 16 1 64 15
4+ 6 3 4 13 9 32 22 1 55 13
5+ 12 9 — 21 15 15 12 1 28 6
6+ 9 2 1 12 8 3 — — 3 —
7+ 12 4 — 16 11 7 4 — 11 3
8+ 4 2 — 6 4 5 1 — 6 1
9+ 4 2 — 6 4 1 — — 1 —
10+ — 3 — 3 2 — — — — —
11+ — 1 — 1 — — — — —
12+ — — — — — — — — —
13+ — 1 — 1 }4 — — — — —
14+ — 2 — 2 — — — — —

Total 66 61 15 142 100 269 154 10 433 100

Mandibles from the 142 wolf-killed deer and 259 hunter-killed deer were examined
closely for abnormal dentition (table 5, figs. 8-10) (Mech et al. 1970) and pathological
conditions (table 6), and the lower limbs of 75 wolf-kills and 126 hunter-kills were also
checked for abnormalities and pathology (table 7, fig. 11). Statistical comparison
showed that the incidence of each condition was significantly higher in the sample from
wolf-killed deer (table 8).
Jaw necrosis found in our specimens was similar to that described by Murie (1944) for
Dall sheep and Mech (1966a) for moose. Generally animals with this condition are old,
and ours were no exception.

Figure 7.—Comparison between the age
structures of deer killed by wolves, deer killed by
hunters, and a theoretical population from the
same general area of northeastern Minnesota.

Figure 8.—Deciduous first premolar (arrow), usually not
present in deer, was found in specimen M-31.

Figure 9.—A permanent first premolar (arrow) was
discovered in M-8.

Figure 10.—An extra set of fourth premolars (arrows)
occurred in specimens M-96.

Table 5.—Abnormalities in the mandibular dentition of deer from the Superior
National Forest, Minnesota

Cause Side
Specimen
Sex Age[28] of of Abnormality
number
death jaw[29]
Years
M-8 F 3+ Wolves Right P1 present (fig. 9)
Normal; no P1 present
Left
outside or inside jaw
Deciduous P1 present
(fig. 8) and permanent
17
M-31 F Wolves Both P1 present inside left
mon.
ramus; right side not
examined internally
M-45 M 4+ Wolves Right P2 rotated 90°
Left P2 absent
M-52 M 4+ Wolves Right P2 absent
Left Normal

2 permanent P4s
M-96 F 2+ Hunters Right present; both crooked
in orientation (fig. 10)
P2 diagonal; P3
normal; P4 below
gumline, pointed
posteriorly and
Left
wedged against M1;
appears to have
pushed out original P4
(fig. 10)
Third column of M3
M-117 M 5+ Hunters Right
reduced
Third column of M3
absent although
M-191 M 4+ Wolves Right
rudimentary root
present
Third column of M3
Left much reduced, peg-
like, and almost
separate
M-225 — 4+ Wolves Right P2 absent
Left P2 situated diagonally
Third column of M3
M-234 F 5+ Wolves Right
reduced
P slightly crooked in
M-254 M 2+ Hunters Right 2
orientation
P2 slanting posteriorly
Left
and crowding P3
Third column of M3
M-272 M 5+ Hunters Right reduced, peg-like, and
almost separate
Third column of M3
peg-like and separated
Left
from second column
by 4 mm.
M-296 F 5+ Wolves Right Normal
Left Extra permanent P4
crowding original P4;

much like M-96
Permanent P2 still not
emerged but appears
M-369 M 3+ Hunters Right
to be wedged against
root of P3

FOOTNOTES:
[28] Based on incisor sectioning method of Gilbert (1966) except that underlined figures are
based on tooth replacement or wear (Severinghaus 1949).
[29] Where only one side is listed, the other was not available.

Table 6.—Pathological conditions in the lower jaws of deer killed by wolves or
hunters[30]

Cause Approximate
Specimen
Sex Age of date of Condition
number
death death
Years
Lump in left
side of
M-70 M 6½ Wolves Feb. 1968 mandible
near M1 and
M2
Large lump
in left
diastema
M-192 M 7½ Wolves Jan. 1969
apparently
from healed
fracture
Light
necrosis
M-206 M 8½ Wolves Jan. 1969
around base
of teeth
M-218 M 3½ Wolves Feb. 1969 Large lump
in left
diastema

apparently
from healed
fracture
Heavy
necrosis
around
molars and
extending
M-228 F 11½ Wolves Mar. 1969 into bone;
half of each
M3
destroyed,
both roots
and crown
Light
necrosis
M-236 F 14½ Wolves Feb. 1969
around base
of teeth
Heavy
necrosis and
M-402 F 10½ Hunters Nov. 1968 lumps on
both sides of
mandible

FOOTNOTES:
[30] Not including dental abnormalities, which are described in table 5.

The following organs were excised from wolf-killed deer and examined grossly in the
field for parasites and abnormalities (fig. 12): lungs (six animals, normal); heart (seven
animals, normal); liver (four animals, one small unidentified tapeworm cyst). Twin
fetuses were found in each of two adult does examined.
Twelve deer were checked for body fat in one or all of the following areas: back
(subcutaneous), kidneys, heart, omenta. Of these animals, seven had large amounts of
fat, but five were almost depleted of fat from these stores. These five were all killed in
February or March 1969; three were fawns, and two were yearlings that had not yet
shed their deciduous premolars.

Of 69 animals examined for femur marrow condition, two had fat-depleted marrow. One
was a fawn killed in March 1969 that had not shed its deciduous first incisors, and the
other was a 5½-year-old buck killed in February 1966.
A fawn and a yearling that had died in February 1969 from unknown causes also had
fat-depleted, marrow. These animals might have been killed by wolves, for wolves had
fed on them. However, they could have died from malnutrition and been eaten as
carrion.

Table 7.—Pathological conditions in the lower limbs of deer killed by wolves or
hunters

Cause
Specimen
Sex Age of Condition
number
death
Years
Right hind foot: "Old healed ankylosis
of the pastern joint ... a spontaneously
healed bacterial arthritis with the
M-28 M 5½ Wolves
destroyed joint cavity filled in by solid
bone. This deer probably had defective
gait"[31] (fig. 14).
Front foot: "A 3×4×5 cm. fibrous mass
in the subcutis about the digital flexor
tendon on the volar surface of the
metacarpus. The surface was denuded,
M-29 F 5½ Wolves
ulcerated, and superficially infected by
surface bacteria.... Probably did detract
from the animal's speed of flight"[31]
(fig. 15).
Hind foot: "Probable that the lesion was
at one time an active bacterial bone
marrow infection that had eventually
fistulated to the skin.... Regional tendons
M-37 F 7½ Wolves and their sheaths were also present
among this inflammation and scarring,
and it would be fair to assume that the
animal's agility was impaired to some
extent."[31]
M-115 M 4½ Hunter Right front hoof: Broken at tip.

Left front foot: "Two severe transverse
lacerations on the volar surface. Each
was approximately 4 cm. in length. One
was located at the margin of the heel,
and the other was located several cm.
M-196 F 4½ Wolves proximad. The more proximal wound
had severed the flexon tendons, and the
consequent uselessness of the limb was
suggested by the splayed toes, the
unmarred hoof wall and unworn soles"
[32]
(fig. 16).
Left hind leg: "A diffuse swelling of the
distal metatarsal bone, the surface of
which was studded with small
osteophytic spicules. The major flexor
and extensor tendons were forced to
assume a convex course over the
summits of the dorsal and plantar
M-227 M 9½ Wolves
surfaces of the defect, but the tendon
sheaths were clean and the normal wear
on soles of the involved toes suggested
that functional deficit and pain were
probably minimal ... quite certainly a
callus from previous fracture"[32] (fig.
17).

FOOTNOTES:
[31] D. M. Barnes. Personal correspondence to L. D. Mech, April 11, 1967.
[32] D. M. Barnes. Undated laboratory report transmitted to L. D. Mech in 1969.

Figure 11.—The jaws and legs of kills were inspected closely
for abnormalities. (Photo courtesy of L. D. Frenzel.)

Figure 12.—When internal organs were present in kills,

they were examined in the field. (Photo courtesy of L. D.
Mech.)

DISCUSSION AND CONCLUSIONS

It has been established that wolves hunting Dall sheep (Murie 1944), caribou (Crisler
1956), moose (Mech 1966a), and other species usually have a low percentage of
success. In the case of a pack of 15 wolves hunting moose on Isle Royale during winter,
only 4.6 percent of all the moose detected by the pack were killed; considering only the
moose that the wolves caught up to or held at bay, the kill rate was 7.6 percent (Mech
1966a).
What little evidence there is about wolves hunting deer indicates that the success rate is
also low with this prey species, at least in winter. The senior author has now observed a
total of 14 deer being chased by wolves in northeastern Minnesota, mostly by packs of
five, seven or eight wolves (Mech 1966b, and see Mech et al., p. 1). In only one case
(6.7 percent) did the wolves (a pair) succeed in catching their prey.
Low hunting success rates imply that the circumstances influencing hunts are seldom
favorable enough, or the prey animals encountered are seldom vulnerable enough for the
wolves to succeed. When the evidence cited earlier that most wolf-killed animals are
inferior members of their populations is considered, the most cogent explanation for the
low hunting success of wolves is that relatively few prey animals are vulnerable.

Table 8.—Incidence of various abnormalities and pathological conditions in wolf-
killed deer compared with that in hunter-killed deer

Wolf-kills Hunter-kills
Level of
Condition Deer in Deer with Deer in Deer with significance
sample condition sample condition
Number Number Percent Number Number Percent Percent
Dental [34]
142 8 5.6 259 5 1.9 90
abnormalities
Jaw necrosis,
lumps, [34]
142 6 4.2 259 1 0.4 95
or
fractures[33]
Pathology of
75 5 6.7 126 1 0.8 95
lower limbs

FOOTNOTES:
[33] Two mandibles from wolf-killed deer had large lumps from healed fractures in the region
of the diastemas.
[34] If all dental and jaw abnormalities are pooled, the difference between the incidence in the
wolf-kill sample (9.8 percent) and that in the hunter-kill (2.3 percent) is significant at the 99
percent level.

Age Structure

Our data strongly indicate that in northeastern Minnesota wolves prey much more
heavily on the older members of the deer population, at least during winter (fig. 7).
Substantial vulnerability to wolves seems to begin at about the age of 5 years (fig. 13),
because the percentage of wolf-killed deer in each year class increases from 9 percent
for 4½-year-old animals to 15 percent for 5½-year-olds (table 4). Indeed, 48 percent of
the wolf-kills were aged 5½ and over, which compares favorably with the Ontario figure
of 58 percent for these age classes (Pimlott et al. 1969).
These figures assume added significance when compared with a sample of deer killed
by hunters in the same general area (fig. 1). Only 10 percent of the hunter-killed deer
were 5½ years old or older, and the percent killed in each year class dropped off
suddenly from 13 percent aged 4½ to 6 percent aged 5½. If the age structure of the
hunter-kill sample is reasonably representative of the age structure of the population at
large, the wolf-kill data show that wolf predation in our study area during winter has a
definite selective effect on the deer population.
There is no direct way of knowing that the age structure of the hunter-killed deer
represents the age structure of the deer population at large. However, sampling hunter-
kills is the most practical means available for gaining an index to the age structure of the
existing herd. Further, there are three indirect pieces of evidence indicating that the
hunter-kill sample represents the actual age structure of the population, just as Maguire
and Severinghaus (1954) found in New York. First, our sample has the basic theoretical
form expected of a stable deer herd; i.e., the youngest year class contained the most
members, and each older cohort included fewer (fig. 7). Second, the age structure of our
sample has the same form as most other deer age structures from widely diverse areas,
(Ontario, Pimlott et al. 1969; southern Minnesota, Erickson et al. 1961; Massachusetts,
Shaw 1951). Third, there is no reason to believe that in our area rifle hunting is
especially selective for any particular age classes. In talking with large numbers of
hunters, we have learned that most shoot at any and all deer they happen to see.
Even if the age structure of the hunter-kill sample did not approximate that of the actual
herd, the comparison of the wolf-kill with the theoretical population dictates the same

conclusion: the rate of kill of
older deer by wolves was several
times greater than that of
younger deer, excluding fawns
(fig. 13). In any case, if the
actual deer population in our
study area had an age structure
similar to that of our sample of
wolf-kills (which would be the
only age structure that would
contradict our conclusion), its
numbers would be declining by
orders of magnitude each year,
and there would now be only a
remnant population. Such
obviously is not the case.
The only other question that
might arise from a comparison of
the age structure of our wolf-
killed deer with that of the
hunter-killed deer concerns the
area from which each sample
was taken. Fifty of our wolf-kills
came from a region almost
inaccessible to hunters (fig. 1).
However, the other 92 came
from the same general area as the
Figure 13.—Relative rates of predation on deer of hunter-kills. Nevertheless, there
various ages, based on comparisons of the ages of was no statistically significant
wolf-killed deer with those of a theoretical difference in age structure
population (dashed line) and those of the hunter- between the wolf-kills from the
killed population. See figure 7. wilderness versus those from the
hunted area (table 1). This fact
also suggests that the human
hunting in the area is relatively
light and has little effect on the age structure of the deer population in the area.
Wolves may also be taking a disproportionately high number of fawns, although our
data do not show this. Nevertheless, there may be a bias against fawns in our method. It
is not unusual to discover the remains of a wolf-killed deer so completely eaten that
there is no indication left of the animal's age. Because fawns often are only about half
the size of adult deer, and their skeletons have not yet completely ossified, the chances
are better that fawns will be more completely eaten. Pimlott et al. (1969) also

recognized this possible bias, although their data did indicate that wolves were killing a
higher percentage of fawns than occurred in the population.
Our study does support the other conclusion of Pimlott et al. (1969), based on a study of
331 kills, that wolf predation on deer during winter shows a definite selection for older
animals. It does not agree with the tentative conclusion of Stenlund (1955) that wolves
in the Superior National Forest do not prey disproportionately on old deer. However,
Stenlund's conclusion was based on 36 kills and on the assumption that only deer at
least 7 years old were "old." Deer 5 years old and older composed 33 percent of
Stenlund's sample, a figure considerably higher than the 10 percent in these age classes
in our hunter-kill sample (table 4). Thus Stenlund's data do not contradict our
conclusion.
The age of 5 years seems to be the beginning of the period of vulnerability for adult
deer. Although 5 years might not seem especially old, there are two aspects of
significance concerning deer of this age and older. First, they are in the second half of
the life span for most members of the species, and their alertness and ability to bolt
quickly away might be expected to decline. It is of interest in this regard that Klein and
Olson (1960, p. 87) believed 5 years of age to be "the upper limit of physiological
efficiency" of black-tailed deer (Odocoileus hemionus) in Alaska. Second, up to the age
of at least 4½ years, and perhaps beyond, the apparent weight-load-on-track of deer
increases with age (Kelsall 1969). Thus older deer would sink farther into the snow than
younger ones, and their escape might be slowed and hindered more. For further
discussion of the effect of snow on the vulnerability of deer, see Mech et al. (p. 51).

Sex Ratio

Statistical tests comparing a number of subsamples of both wolf-killed deer and hunter-
killed deer showed a series of significantly different sex ratios (tables 1)-3. The ratio of
males to females in the fawn cohort of the hunter-kill, which is probably the most
representative of the actual fawn sex ratio, was even (table 2). With wolf-kills, however,
a significantly higher percentage of females was taken in the fawn subsample (59
percent) than in the adult subsample (46 percent). These results compare favorably with
those of Stenlund (1955), who found that from 1948 to 1953 in the same area as the
present study 68 percent of 19 sexable fawn wolf-kills were females and 44 percent of
63 sexable adult wolf-kills were females.
If the sex ratio of fawns began even, and more females than males were killed by
wolves, then a higher proportion of males would be left in the adult population, unless
some other mortality factor kills more male fawns. Thus it is not surprising that in the
wilderness area, where little or no hunting is done, the sex ratio of wolf-kills in the adult
cohort is significantly heavy toward males (71 percent: 29 percent). This was also true
of the wolf-kills in Algonquin Provincial Park, where males made up 57 percent of the
total sexable wolf-kill (Pimlott et al. 1969). The latter figure may even have been higher

if calculated for adults alone, for a preponderance of female fawns in the Algonquin
Park data (such as occurred in our and Stenlund's samples) would tend to obscure the
preponderance of males in the adult sample.
The adult subsample of hunter-kills also contained a higher percentage of males (66
percent : 34 percent). Although this might also reflect the influence of wolf predation on
female fawns, it probably is more a result of the greater movement of bucks during the
hunting season, which overlaps with the rutting season. Even the sex ratio of adult deer
killed in wolf-free areas shows a preponderance of males (Erickson et al. 1961).
However, it appears that the higher harvest of bucks by human hunters does markedly
affect the sex ratio of the deer population in the hunted area, for the wolf-kill of adults in
that area contained a significantly higher percentage of does (56 percent) than did the
wolf-kill of adults in the wilderness area (29 percent).
Evidently the hunter harvest is not heavy enough to affect the age structure of the deer
population to any marked degree, for no significant difference in age structure was
found between the wolf-kill in the hunted area and that in the wilderness area (table 1).
This does not conflict with the conclusion that hunting affects the sex ratio of the deer
herd, because it would take much less to influence a population characteristic having
two classes (sex) than one having 14 (age).
One additional difference in the sex ratio was found between two other subsamples of
the wolf-kill—that is, the wolf-kill before and after an unusually high snow
accumulation, which reached its peak about February 1, 1969 (table 1). Of a total of 77
animals killed before this snow condition occurred (including those from previous
years), 38 percent were females. Of 44 animals killed after the heavy accumulation, 57
percent were females. One possible explanation for this is that females may normally be
less vulnerable to wolf predation, for Kelsall (1969) has shown that they probably have
a lighter weight-load-on-track than males. Thus when snow conditions changed greatly,
making deer generally much more vulnerable to wolves (see Mech et al., p. 35), a
preponderance of does suddenly might have become available. There is some evidence
that does may be generally less vulnerable under most conditions, for all seven of our
wolf-killed deer over 10 years old were females, and the oldest was over 14.

Condition of Wolf-Killed Deer

Because the data show that wolves in our study area tend to kill a disproportionate
number of older deer, it is not surprising to discover that wolves also tend to capture a
disproportionate number of individuals with abnormalities and pathological conditions
(table 8). The explanation for such selection is obvious in regard to the abnormalities of
the lower limbs (figs. 14-17): deer with injured or abnormal limbs simply cannot run as
fast or as agilely as normal animals (table 7). Our observations show that deer usually
depend on their alertness and speed to escape approaching wolves (Mech 1966b, Mech

et al., p.
1). Any
trait or
conditio
n that
tended
to
interfere
with
either
alertnes
s or
speed
would
decrease
an Figure 15.—Infection and fibrous
individu mass in a front foot of specimen M-
Figure 14.—Arthritis in right hind al's
foot of specimen M-28. (Photo 29. (Photo courtesy of University of
chance Minnesota Veterinary Diagnostic
courtesy of University of Minnesota of
Veterinary Diagnostic Laboratory.) Laboratory.)
escape.
It is
more
difficult
to
explain
how
dental
abnorm
alities or
patholo
gical
conditio
Figure 17.—Healed fracture of left
ns of the
hind leg of specimen M-227. (Photo
mandibl
courtesy of University of Minnesota
e (figs.
Veterinary Diagnostic Laboratory.)
8-10)
would
predispo
se an individual to wolf predation. However, in
the case of dental abnormalities the genetic or
environmental conditions that caused the Figure 16.—Injury to left front foot

abnormality might also have caused some other of specimen M-196. (Photo courtesy
trait that increased the animal's vulnerability. Or of L. D. Mech).
the abnormal condition itself may have caused a
further, more critical, disruption of the animal's physiology or behavior, which in turn
predisposed it to wolf predation.
The finding of several wolf-kills with poor fat stores could indicate that primary or
secondary malnutrition was a factor in the animals' deaths. However, it would take a
statistical comparison between the fat stores of the deer at large and those of the wolf-
kills to establish this.
The discovery that 13 percent of the fawns and 84 percent of the yearlings killed during
January, February, and March had not yet shed their deciduous incisors and premolars,
respectively, also fits well with the rest of our information. Evidently some unusual
factor had caused the delay in tooth development and replacement. One possibility is
that the animals were born in August or September, much later than normal. Although
most deer in Minnesota are born in May and June, there are records of births in July and
August. In addition, a fetus 181 to 200 days old was found in a doe killed on September
26 (Erickson et al. 1961).
An alternate explanation for the delay in tooth replacement is that the animals were
suffering from malnutrition or nutrient deficiency. Severinghaus[35] has evidence that
yearling bucks that have not replaced their deciduous premolars during November, and
thus are aged at 17 months (Severinghaus 1949), generally have shorter, narrower
antlers and fewer points than 18-and 19-month-old individuals. Degree of antler
development in turn is considered related to nutritional state (Latham 1950). Thus it is
reasonable to conclude that animals behind in tooth development and replacement,
whether this is caused by age or diet, are physiologically inferior.
Most of the abnormal conditions discussed above pertain to the skeletal parts of wolf-
kills. If the soft parts of a large number of kills could be examined thoroughly, one
might discover a much higher incidence of diseases and other pathological conditions.
In conclusion, our data on both age and condition of wolf-killed deer show that at least
during winter, wolves in our study area usually do not kill just any deer they discover,
although they do try to. Evidently, most deer can usually escape wolf predation. The
most frequent exceptions are those 5½ years old and older, those born late, those
suffering from poor nutrition, those with abnormalities or pathological conditions, and
possibly fawns.
The above conclusions parallel those of Murie (1944), Crisler (1956), Mech (1966a),
and Pimlott et al. (1969) for wolves preying on Dall sheep, caribou, moose, and deer
respectively, and further substantiate the claim by Mech (1970) that they can be
extended to wolves preying on most, if not all, species of large mammals under most
conditions. It is also apparent from the data presented above that deer over 5 years of

age and those with abnormalities of the jaw or lower limbs represent such a small
percentage of the total population that they are seldom taken by human hunters. In this
respect, competition between timber wolves and human hunters appears to be minimal
in the study area.

FOOTNOTES:
[35] C. W. Severinghaus. Unpublished data.

SUMMARY

White-tailed deer (Odocoileus virginianus) killed by wolves (Canis lupus) during winter
in a relatively unhunted wilderness area and in an immediately adjacent hunted area of
Minnesota were compared with deer killed by hunters in the same general area, and with
a hypothetical population. Deer killed by wolves were significantly older. Statistical
comparisons also showed the following: (1) hunters generally killed an even sex ratio of
fawns, and a disproportionate number of adult bucks, (2) wolves took a higher
percentage of female fawns than female adults, a disproportionate number of bucks in
the wilderness area, and a higher percentage of does in the hunted area. The latter fact
evidently reflects the higher hunter success on males in the hunted area. Significantly
higher incidences of abnormalities and pathological conditions of both mandibles and
lower limbs were found in wolf-killed deer than in hunter-killed deer, and these
conditions are described. It is concluded that wolf predation on white-tailed deer in the
study area during winter generally is selective in that it tends to remove members of the
prey population that are old, debilitated, or abnormal. Apparently these classes of deer
represent such a small percentage of the population that they are seldom taken by
human hunters.

ACKNOWLEDGMENTS

This study was supported by Macalester College, the New York Zoological Society, the
Minnesota Department of Conservation, the USDA Forest Service, and the U.S. Bureau
of Sport Fisheries and Wildlife. Pilots Robert Hodge, Pat Magie, John Winship, Jack
Burgess, Don Murray, and Walt Neumann aided substantially in obtaining jaws from
wolf-killed deer. Students from the Macalester College Biology Department and
personnel of the USDA Forest Service and the Minnesota Department of Conservation
helped secure mandibles from both wolf-killed and hunter-killed deer. The interest of
Mr. John E. Peninger and of many deer hunters in contributing the jaws is also greatly
acknowledged.

Mr. David W. Kuehn sectioned the incisors of the deer jaws and determined their ages.
Dr. Donald M. Barnes of the University of Minnesota Veterinary Diagnostic Laboratory
examined the abnormal lower limbs, described their pathology, and provided photos of
specimens used herein.
Mr. Wallace C. Dayton and Miss Elizabeth Dayton and the Quetico-Superior
Foundation, all of Minneapolis, financed Mech during the preparation of this paper.
The following individuals read the manuscript and offered many helpful suggestions:
Mr. R. L. Downing, Mr. C. W. Severinghaus, Mr. J. M. Peek, Dr. C. T. Cushwa, Mr. M.
H. Stenlund, and Dr. R. R. Ream.

LITERATURE CITED

Crisler, Lois. 1956. Observations of wolves hunting caribou. J. Mammal. 37: 337-
346.
Downie, N. M., and Heath, R. W. 1959. Basic statistical methods. 289 p. New
York: Harper and Bros.
Erickson, A. B., Gunvalson, V. E., Stenlund, M. H., Burcalow, D. W., and
Blankenship, L. H. 1961. The white-tailed deer of Minnesota. Minn. Dep.
Conserv. Tech. Bull. 5, 64 p.
Gilbert, F. F. 1966. Aging white-tailed deer by annuli in the cementum of the first
incisor. J. Wildl. Manage. 30: 200-202.
Kelsall, J. P. 1969. Structural adaptations of moose and deer for snow. J. Mammal.
50: 302-310.
Klein, D. R., and Olson, S. T. 1960. Natural mortality patterns of deer in southeast
Alaska. J. Wildl. Manage. 24: 80-88.
Kuehn, D. W. 1970. An evaluation of the wear method as a criterion for aging
white-tailed deer. M.S. Thesis., Univ. Minn.
Latham, R. M. 1950. Pennsylvania's deer problem. Penn. Game News, Spec. Issue
1. (Cited from: Allen, D. L. 1962. Our Wildlife Legacy.)
Maguire, H. F., and Severinghaus, C. W. 1954. Wariness as an influence on age
composition of white-tailed deer killed by hunters. N. Y. Fish and Game J.
1: 98-109.
Mech, L. D. 1966a. The wolves of Isle Royale. U.S. Nat. Park Serv. Fauna Ser. 7,
210 p.

Mech, L. D. 1966b. Hunting behavior of timber wolves in Minnesota. J. Mammal.
47: 347-348.
Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species.
389 p. New York: Natural History Press, Doubleday.
Mech, L. D., Frenzel, L. D., Jr., Karns, P. D., and Kuehn, D. W. 1970. Mandibular
dental anomalies in white-tailed deer from Minnesota. J. Mammal. 51: 804-
806.
Murie, A. 1944. The wolves of Mount McKinley. U.S. Nat. Park Serv. Fauna Ser.
5, 238 p.
Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The ecology of the
timber wolf in Algonquin Provincial Park. Ont. Dep. Lands and Forests
Res. Rep. (Wildl.) 87, 92 p.
Ryel, L. A., Fay, L. D., and Van Etten, R. C. 1961. Validity of age determination
in Michigan deer. Mich. Acad. Sci., Art, and Letters 46: 289-316.
Severinghaus, C. W. 1949. Tooth development and wear as criteria of age in
white-tailed deer. J. Wildl. Manage. 13: 195-216.
Severinghaus, C. W. 1955. P. R. Rep. W-28-R-9: Job 1A, April 13, 1955.
Shaw, S. P. 1951. The effect of insufficient harvests on an island deer herd. N.E.
Wildl. Conf. (Mimeo).
Siegel, S. 1956. Non-parametric statistics for the behavioral sciences. 312 p. New
York: McGraw-Hill.
Stenlund, M. H. 1955. A field study of the timber wolf (Canis lupus) on the
Superior National Forest, Minnesota. Minn. Dep. Conserv. Tech. Bull. 4, 55
p.

THE EFFECT OF SNOW CONDITIONS ON THE
VULNERABILITY
OF WHITE-TAILED DEER TO WOLF PREDATION
L. David Mech, L. D. Frenzel, Jr., and P. D. Karns
Wolves (Canis lupus) and deer (Odocoileus virginianus) having evolved together, no
doubt have become adapted to contending with each other's physical abilities. Thus it is
not surprising to learn that deer which succumb to wolf predation are generally weaker,
older, or abnormal compared with the total deer population (Pimlott et al. 1969, also see
Mech and Frenzel, p. 35).
However, the structural and behavioral adaptations of both species must have evolved
under environmental conditions that are average or usual; otherwise, an adjustment of
wolf to deer populations, and vice versa, could not have been maintained over long
periods. This implies that extreme or unusual conditions might sometimes occur, to
which either the wolf or the deer is poorly adapted.
One of the most important environmental factors that can influence the interactions of
wolves and deer is snow. The total fall, depth on the ground, and the density are all
aspects of snow that may vary considerably and affect the ability of wolves to capture
deer. Recent studies of wolves and deer in northeastern Minnesota (see Mech et al., p. 1,
also Mech and Frenzel, p. 35) afforded us opportunities to investigate the relationships
between snow and the interactions of wolves and deer.

METHODS

Two principal methods of study were used in this investigation. The first involved
recording the snow depth and support quality ("penetrability") in feet and tenths of feet
(Verme 1968). Snow measurements were taken during the winters of 1966-67, 1967-68,
and 1968-69, in which large differences in snow conditions existed. Ten such
measurements were made weekly near Isabella, Minnesota, in an open aspen (Populus
tremuloides) stand away from influences that might have caused drifting or other
unusual snow conditions; the measurements were averaged. Penetrability was
determined with Verme's snow-compaction gauge—a 3-foot piece of 1-1/8-inch (outside
diameter) copper tube filled with lead to total 3 pounds, which gives a weight per area
of 211 gm./cm.2. To obtain a measurement, the pipe is held vertically with its lower end
just flush with the snow, and then is released. The depth to which it sinks is considered
the penetrability of the snowpack by a walking deer.

Although the snow conditions measured at Isabella are not representative of the entire
study area, year-to-year comparison in the Isabella area should also apply generally
throughout the region.
The second technique used in this study was observing the movements of wolves and
deer. This was usually done from low-flying aircraft, and was facilitated by the use of
radiotracking, as described by Mech et al. (p. 1). Close inspection of wolf-killed deer
was made from the ground (Mech and Frenzel, p. 35).

RESULTS AND OBSERVATIONS

Snow measurements for each winter are shown in figures 1 through 3. The winter of
1968-69 was the most extreme of the three in terms of accumulated snow, and was
generally regarded as having one of the heaviest snowfalls and accumulations on record
for the study area. Snow depth on the level near Isabella reached 3.9 feet at one time,
and from January 3 to April 4 it exceeded 2.4 feet. The highest snow level reached
during 1966-67 was 2.4 feet, and the highest level reached during 1967-68 was 1.4 feet.
In the vicinity of Ely, some 30 miles from Isabella, the 1968-69 peak accumulation was
39 inches, the highest accumulation since 1948-49 when records were first kept.[36]
Thus we consider the winters of 1966-67 and 1967-68 to be within the normal range for
the study area, and the 1968-69 winter as being most unusual (fig. 4).

Figure 1.—Snow depth and penetrability by deer and wolves near
Isabella, Minnesota, 1966-67.

Figure 2.—Snow depth and penetrability by deer and wolves near
Isabella, Minnesota, 1967-68.

Figure 3.—Snow depth and penetrability by deer and wolves near
Isabella, Minnesota, 1968-69.

The snow penetrability in 1966-67 remained high throughout January, February, and
March. During the following winter, penetrability fluctuated more, but even at its
greatest, it was relatively unimportant to deer because the total snow depth was so low.
During 1968-69, however, penetrability was a very important aspect of snow condition.
It was so high during late January and early February, when snow accumulation was
also at its peak, that a walking deer would be expected to sink in 2.5 to 3.5 feet. Snow
penetrability then decreased through February and March to a point where a walking
deer would sink in approximately 0.6 foot on March 21. However, because snow
accumulation remained so high through February and March, the lower penetrability
during late February and March still afforded no relief to running deer, because they
must exert forces several times as great as when walking. On the contrary, the low
penetrability (which is an indirect measure of density) could be expected to hinder a
running deer in deep snow, for it would cause much more resistance.
Deer movements, like snow conditions, varied greatly during the three winters of the
study. During the first two winters, deer were generally found singly and in groups of

two to six, often around the shores of
lakes but also scattered about inland. In
late January and February 1967, running
deer were observed sinking deeply into
snow, but their movements still did not
seem to be hindered, no doubt because of
the high penetrability (low density) of the
snow that year (fig. 1).
However, during late January, February,
and March of 1969 the deer were much
more concentrated, mostly in conifer
swamps, along southwest-facing slopes,
or on lakes. Although groups of two or
three animals could be found in scattered
Figure 4.—During the winter of 1968-69, inland "pockets" throughout the winter,
the snow was unusually deep in the study groups of five or six were not uncommon
area. (Photo courtesy of L. D. Frenzel.) on lakes during January. The tendency to
concentrate continued to increase, and on
February 6, as many as 11 deer were
observed on one lake; by March 13,
group size had increased to as high as 22 deer in the same area. Throughout February
and March, heavy concentrations of deer tracks covered most wilderness lakes, further
evidencing much greater use of shorelines than had occurred in the two previous winters
(fig. 5).
No doubt deer tended to concentrate on lakes because travel inland became so difficult.
On January 28, two deer were seen plowing through snow up to their necks. Although
the snow began settling in February, and the penetrability decreased, by late February
running deer still plunged chest-deep and had to hesitate at every bound. These
conditions persisted until about March 26, by which time a surface crust strong enough
to hold a running deer had formed.

In
consi
derin
g
wolf
mobi
lity
in
snow
, two
types
of
mov
emen
t
must
be
reco
gnize
d:
the
trot
used
durin
g
gene
ral
trave Figure 5.—Under unusually deep snow conditions, deer used lake
l, shores heavily. (Photo courtesy of L. D. Mech.)
and
the
bounding used while chasing prey. The trot is an easy gait of about 5 m.p.h.
on firm footing (Mech 1970), and can be continued for hours at a time.
During periods of deep snow and high penetrability, most wolf travel is on
frozen waterways, roads, snowmobile trails, and animal trails, including the
wolves' own pathways, which become well packed with frequent use (fig. 6,

7A, B). Such travel was observed during each of the three winters of this
study.

Figure 6.—Wolves travel single file in deep snow. (Photo courtesy of L.
D. Mech.)

The second type of wolf movement affected by snow is the leaping and
bounding associated with chasing prey. The shallower angle of the wolf's
bound (fig. 8) (compared with that of the deer) often causes the wolf to
flounder in snow that presents little hinderance to deer (Mech 1970). Such
was the case in January and February 1967 in our study area. During 1967-
68 no observations of wolves chasing deer were made by the authors, but
reports by other field workers indicated that running conditions were similar
to those of 1967.
During the winter of 1968-69, wolves also bogged down a great deal in snow
when chasing deer. However, after January 1969 the snow was so deep that
deer were floundering even more than wolves in many cases. The fact that
wolves could run in the trail broken by deer probably also gave the wolves
an advantage under the conditions that severely restricted deer movements.
The above observations of snow conditions, deer movements, and wolf
movements during the three winters of the study are in accord with
observations made on the differences in the ability of the wolves to capture

Figure 7.—(A) A single wolf must break his own trail through the snow.
(Photo courtesy of L. D. Frenzel.) (B) Regular use by a pack keeps trails
open. (Photo courtesy of L. D. Mech.)

Figure 8.—Wolves run at a shallow angle, thus hindering them in deep
snow. (Photo courtesy of D. H. Pimlott.)

deer during the same period. Two indices support the conclusion that wolves

had a much easier time catching deer during February and March 1969 than
earlier in the winter and in the two previous winters: (1) the degree of
utilization of wolf-killed deer, and (2) the kill rate of radiotagged wolves.
During the winters of 1966-67 and 1967-68, and in December and early
January 1968-69, most wolf-killed deer found had been thoroughly eaten,
and the bones—if present at all—were well chewed and scattered at each kill
(fig. 9). All skin and flesh from the skull were eaten, and the mandible was
usually separated from the skull. During late February and early March
1967, few fresh kills were even found, and wolves were returning several
times to old kills that had been cleaned up many days before.
However, in late January 1969 a
substantial change began taking
place. The skeletons of most kills
found were almost intact, the flesh
having been eaten from around the
bones (fig. 10). Appreciably more
skin was usually left on the carcass,
especially on the side lying on the
snow, and the neck and head were
generally intact. This was true even
of fawns, which in the past often
were almost completely consumed.
In several cases, only about half of
the flesh had been eaten from the
carcasses. On February 2, 1969, four
deer recently killed by wolves were
found along a 1½-mile stretch of
Birch Lake and nearby Polaris Lake
Figure 9.—Usually the remains of a (Minnesota-Ontario border). One
wolf kill are well chewed and large doe was completely uneaten
scattered before the wolves abandon and remained so for at least 24 hours
them. (Photo courtesy of L. D. after discovery from the air. Further,
Mech.) one fawn had only a few pounds of
flesh eaten, a yearling doe was half
eaten, and another fawn was about
75 percent eaten. Hazardous landing

conditions during this period
severely limited the number of
carcasses that could be examined
from the ground, but on February 6 a
yearling doe was discovered that had
only about 5 to 10 pounds of flesh
eaten, and on February 8 an adult
doe was found that was completely
intact except for wounds.
In past winters some kills had been
located that had been only partly
eaten, but in each case the carcasses
were soon revisited and cleaned up
(Mech 1970). This was often not the
case in 1969. For the rest of the
winter most of the deer killed by
wolves in our study area were not as
completely consumed as in previous Figure 10.—During a period of
winters. Pimlott et al. (1969) found a especially deep snow, wolves
similar relationship between the abandoned many kills before
severity of the winter and the degree pulling apart the skeletons. (Photo
to which wolf-killed deer were courtesy of L. D. Mech.)
utilized.
Correlated with the above information was the kill history of our radiotagged
wolves (Mech et al., p. 1). From December 1968 through January 1969 No.
1051 had killed three or possibly four deer, and generally had spent 6 or 7
days feeding on each. However, throughout most of February this animal
visited a new deer carcass (which presumably he killed) every 3 days, and he
spent only 1 or 2 days at each. In two cases two new carcasses were found in
the immediate vicinity of this animal during the same day, and in each case
the wolf spent only 1 day in the area. A second wolf (1053) which had spent
most of December and January scavenging on the remains of both deer and
moose (Alces alces) that had died long before, made her first known kill of a
deer on January 31, 1969. The kill rate of the other three radiotagged wolves
also increased, although the data for them are less complete. The average kill
rate for all radiotagged wolves and their associates was one deer per wolf per

16 to 20 days before February 1, and one per 8 to 12 days after February 1
(see Mech et al., p. 1).

FOOTNOTES:
[36] M. H. Stenlund. Personal correspondence to L. D. Mech, Oct. 10, 1969.

DISCUSSION AND CONCLUSIONS

Under usual snow conditions throughout most of the range of the white-
tailed deer, healthy vigorous individuals can probably escape most attacks by
wolves. Observations by Mech (1966), Rutter and Pimlott (1968), and Mech
et al. (p. 1) indicate that a high percentage of attempts by wolves to kill deer
during winter are unsuccessful. This is further implied by the figures of
Pimlott et al. (1969) and Mech and Frenzel (p. 35) showing that at least
during winter wolves tend to kill a disproportionate number of old deer as
well as those with various abnormalities and pathological conditions.
However, during a winter with extremely deep snow, the usual relationships
seem to change somewhat. Fewer deer are able to escape wolves, and a
surplus is killed. This means that some individuals not vulnerable under the
usual snow conditions become vulnerable during extreme conditions. There
are two main possible reasons for this, the effect of the extreme weather
conditions on the health and vigor of the deer, and the physical effect of the
snow on the escapability of the deer.
In regard to the first possibility, there was limited evidence that during
February and March 1969 some fawns and yearlings in our study area were
losing their fat stores. Two of three yearlings, and both fawns intact enough
for examination during this period lacked back fat, and the marrow in one of
six fawn femurs was partly fat depleted. Nevertheless, the third yearling
inspected still had back fat, and a 3½-year-old doe had heavy omental, renal,
heart, and back fat during the same period. Thus, although an abnormal

decline in the physical condition of some deer in the late winter might partly
account for the increased kill by wolves during February and March 1969,
the effect of snow on the escapability of the deer probably was also
involved.
The key difference in snow conditions between the two periods—(1) the
winters of 1966-67, 1967-68, and December-January 1968-69, and (2)
February and March 1969—was the heavy, persisting accumulation of snow
during the latter period, combined with the increasing density of the snow.
As our observations show, this greatly hindered the movements of deer
fleeing from wolves.
Under more usual conditions, a running deer might sink through the snow to
the ground and thus obtain a firm footing from which to spring again. In
discussing wolf-caribou relations in snow, Kelsall (1968, p. 249) stated the
following: "While caribou (Rangifer tarandus) will sink into snow even
deeper than wolves, their longer legs permit them to run efficiently where a
wolf will bog down. Nasimovich (1955) considered that roe deer and sika
deer could be taken by wolves when snow was not more than 30 cm. (11.8
inches) in depth. At depths above that their pursuit becomes difficult or
fruitless."
However, it appears that when snow becomes extremely deep, wolves then
gain the advantage. With 22 to 48 inches or more of snow to plow through, a
deer would have trouble even touching a firm foundation. According to
Kelsall (1969), deer measure only 20 to 24 inches from hoof tip to chest,
with legs extended.
It is true that wolves stand even shorter than deer and so might be expected
to flounder even more. However, this is where another factor becomes
important, the "weight-load-on-track" or total weight per area of track. As
Kelsall (1969) has pointed out, the mean weight-load-on-track for deer is
extremely difficult to measure directly, because the actual under-surface of
the deer's foot slants vertically, and a much greater area may be used to
support an animal in snow than on a hard surface. This probably explains the
discrepancy between Kelsall's measurements and work done by Verme
(1968) in Michigan. According to Kelsall, deer weight-load-on-track (hoof
only) varies between 431 and 1,124 gm./cm.2. However, Verme stated that
his compaction gauge (with a weight load of about 211 gm./cm.2, described

earlier in this paper) sank in virtually the same amount in snow as did deer.
Under the snow conditions in our study area, we found that the same type of
compaction gauge generally penetrated to a depth within a half inch of that
to which deer were sinking. On this basis, it seems reasonable to suggest that
a deer in snow is supported by more of its foot than just the hoof, and that
the actual weight-load-on-track of deer in snow is about 211 gm./cm.2.

For wolves, this measure varies from 89 to 103 gm./cm.2 (Foromozov 1946).
This means that for the same amount of force applied during running, a wolf
would have twice as much support as a deer. It also means that in deep snow
a walking wolf generally is much less restricted than a walking deer. Late in
February 1969, for example, when deer were seriously limited in their ability
to travel, wolves were able to travel widely (Mech et al., p. 1).
Even though wolves have much greater support than deer, when running
they still sink into the snow almost as much as deer under most conditions,
probably because both run with such force that snow usually offers little
support. Nevertheless, with extremely deep snow, the difference in support
factor between wolves and deer could become critical, and this is probably
what happened during February and March 1969. With deer seriously
restrained by the deep snow, even a slight advantage in favor of the wolf
could increase hunting success. A high snow density during that period
would accentuate this advantage. This is because until the snow becomes
dense enough to hold a running deer, each increase in density would further
the advantage of the wolf, which would require only half the density to
support it, while it would hinder the deer.
One result of the extreme snow conditions of early 1969 was that deer
tended to gravitate to lakes, where snow was shallow and footing was firm.
Initially upon disturbance by human beings, and probably by wolves, these
deer usually headed inland, but it is apparent from a number of kills
examined that when pressed hard by wolves inland, deer headed out onto
lakes where possible. Apparently they could run there with better footing.
However, frozen lakes also provide wolves with good running conditions,
and even seem to give them an advantage (Rutter and Pimlott 1968, Mech
1970), so many of these deer were killed (fig. 11).

Figure 11.—On frozen lakes, wolves often seem to have the advantage
over deer, such as in this case where the wolf (center) has just killed a
deer and is trying to discourage a raven from joining him in the feed.
(Photo courtesy of L. D. Frenzel.)

Stenlund (1955, p. 44) reported as follows on years of low snowfall, the
opposite condition, which demonstrated the same relationship between snow
depth and kills on lakes: "The winters of 1951-52 and 1952-53 were
abnormally mild with little early snow. As a result, few wolf-killed deer
appeared on the lakes and most deer attempted to outrun wolves in the
woods."
Thus it appears that extreme snow conditions in our study area increase the
vulnerability of deer to wolf predation in three ways: (1) by causing a
decline in the health and nutritional state of some members of the deer
population; (2) by hindering the escapability of the deer; and (3) by causing
deer to congregate on frozen lakes where wolves have the advantage in
running.

SUMMARY

During the winters of 1966-67, 1967-68, and 1968-69, the interactions of
wolves (Canis lupus) and white-tailed deer (Odocoileus virginianus) were
observed in northeastern Minnesota from aircraft. Snow depth and
supporting ability were also measured during these winters, and the ability of
wolves to capture deer was compared for a period of usual snow conditions
versus a period of extreme snow conditions.
It was found that during February and March 1969, when snow remained
from 2.5 to 3.9 feet deep and failed to support running deer, wolves were
able to capture deer more easily. This was evidenced by kills that were left
partly or completely uneaten, and by a higher rate of predation by
radiotagged wolves and their associates.
Although both wolves and deer floundered in the extremely deep snow, the
relatively lighter weight-load-on-track of wolves evidently gave them a
greater advantage than under the usual snow conditions, when wolves were
observed floundering more than deer. This factor, plus a decline in the health
and vigor of some segments of the deer population and a tendency for deer
to congregate on frozen lakes, where wolves have an advantage, help explain
the increased vulnerability of deer to wolf predation during the winters of
deep snow.

ACKNOWLEDGMENTS

This study was supported by Macalester College, the Minnesota Department
of Conservation, the USDA Forest Service, the U.S. Bureau of Sport
Fisheries and Wildlife, and the New York Zoological Society. Pilots John
Winship, Pat Magie, Jack Burgess, and Don Murray flew the observation
planes during radiotracking. Miss Elizabeth Dayton, Mr. Wallace C. Dayton,
and the Quetico-Superior Foundation, all of Minneapolis, financed Mech
during the writing of this report.
Thanks are also due L. J. Verme, J. P. Kelsall, and J. M. Peek for their
helpful reviews.

LITERATURE CITED

Foromozov, A. N. 1946. The snow cover as an environment factor and
its importance in the life of mammals and birds. (Moskovskoe
obshchestvo ispytatelei priroda) Materialy k poznaniyu fauny i
flory SSSR, Otdel. Zool. n. 5 (XX). (Translation from Russian
published by Boreal Institute, Univ. Alberta, Edmonton,
Alberta.)
Kelsall, J. P. 1968. The caribou. Can. Wildl. Serv. Monog. 3, 340 p.
Kelsall, J. P. 1969. Structural adaptations of moose and deer for snow.
J. Mammal. 50: 302-310.
Mech, L. D. 1966. Hunting behavior of wolves in Minnesota. J.
Mammal. 47: 347-348.
Mech, L. D. 1970. The wolf: the ecology and behavior of an
endangered species. 389 p. New York: Natural History Press,
Doubleday.
Nasimovich, A. A. 1955. The role of the regime of snow cover in the
life of ungulates in the U.S.S.R. Moskva, Akademiya Nauk
SSSR. 403 p.
Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The
ecology of the timber wolf in Algonquin Provincial Park. Out.
Dep. Lands and Forests Res. Rep. (Wildl.) 87, 92 p.
Rutter, R. J., and Pimlott, D. H. 1968. The world of the wolf. 202 p.
Philadelphia and New York: J. B. Lippincott Co.
Stenlund, M. H. 1955. A field study of the timber wolf (Canis lupus)
on the Superior National Forest, Minnesota. Minn. Conserv.
Dep. Tech. Bull. 4, 55 p.
Verme, L. J. 1968. An index of winter severity for northern deer. J.
Wildl. Manage. 32: 566-574.

THE POSSIBLE OCCURRENCE OF THE
GREAT PLAINS WOLF
IN NORTHEASTERN MINNESOTA
L. David Mech and L. D. Frenzel, Jr.
The timber wolf (Canis lupus) of northeastern Minnesota occupies an area
within the range given by Goldman (1944) for the eastern timber wolf (C. l.
lycaon Schreber). However, this area is within 150 miles of the eastern edge
of the former range of the Great Plains wolf (C. l. nubilus Say), and there is
some question as to whether the Minnesota wolf is really an intergrade
between these two subspecies. Writing of nubilus, Goldman (1944, p. 444)
stated: "Specimens from eastern Minnesota and Michigan seem more
properly referable to lycaon, but relationship to nubilus is shown in
somewhat intermediate characters."
In describing lycaon as basically a gray wolf, Goldman made no mention of
the occurrence of black or white color phases in that subspecies. However, in
discussing nubilus, Goldman (1944, p. 442) wrote the following: "Many
color variations are presented. Individuals may be nearly white at any
season, except for a sprinkling of black hairs over the back, a small, narrow,
but conspicuous, black patch over the tail gland, and a more or less distinctly
black tip. Black individuals may occur in the same litter with those normally
colored." Goldman also referred to nubilus as "now probably extinct."
In the eastern part of the range of
lycaon, color phases other than gray
appear to be rare as Rutter and
Pimlott (1969, p. 188) attest: "The
uniformity of the color of timber
wolves in many areas is evidenced
by the work in Algonquin Park, in
Ontario. There, over the past eight
years, dozens of packs have been
Figure 1.—A few wolves observed
observed from the air. However, we

in the study area were jet black. have never been able to discriminate
(Photo courtesy of L. D. Mech.) between any of them on the basis of
the color variation of individual
animals."
Thus it seems significant to report on incidences of black and white color
phases in wolves that we have observed in northeastern Minnesota during
some 480 hours of flying associated with wolf research (Mech et al., p. 1).
The observations took place in the Superior National Forest, in northern
Cook, Lake, and St. Louis Counties during the winters of 1966-67, 1967-68,
and 1968-69. A total of 309 sightings were made of wolves that could be
classified by color; of these, 11 (3.6 percent) were jet black (fig. 1) and two
(0.6 percent) were creamish white, with the cream color the most intense on
the back. No doubt some of the grays, and perhaps the blacks and whites,
were repeated observations, but the figures should provide a reasonable
approximation of the incidence of these color phases in this area. All black
or white animals except one were observed with gray wolves (table 1 and
fig. 2).
A number of black wolves, and a few white wolves, have been seen by other
observers, all in the three counties listed earlier. To gain some idea of the
past incidence of these color phases in the same general area, we asked
Conservation Officers Robert Hodge, Robert Jacobsen, and Frank Baltich of
the Ely, Minnesota, area about the numbers of each phase that they took
before 1960. They reported killing an approximate total of 580 wolves, of
which four were black and three were white or creamish white.

Table 1.—Observations of wolves of black and white color phases

Color combinations
Date Location
within each pack
Feb. 24, 1967 T64N-R8W-S1 Vera Lake 3 grays; 1 black; 1 white
Mar. 4, 1967 T63N-R9W-S27 Lake Two 3 grays; 2 blacks
Dec. 18, 1968 T63N-R8W-S35 Lake Insula 2 grays; 2 blacks[37]
Jan. 17, 1969 T65N-R8W-S27 Carp Lake 1 gray; 1 white

Feb. 1, 1969 T63N-R8W-S13 Lake Insula 4 blacks; 2 grays[38]
Feb. 5, 1969 T63N-R8W-S8 Benezie Lake1 black
Feb. 6, 1969 T63N-R10W-S33 Clear Lake 3 grays; 1 black

FOOTNOTES:
[37] These animals were near the shore of the lake, so others may have been
inland where they could not be seen.
[38] This group might well have been the same as that seen on Dec. 18, 1968.

Figure 2.—A pack of four blacks with two grays (first and third). (Photo
courtesy of John Winship.)

Because black and white color phases have rarely if ever been reported for
lycaon, yet were well known for nubilus, it is not unreasonable to conclude
that the race of wolves now occupying northeastern Minnesota does show
strong nubilus influence. Goldman examined the skulls only of 10
Minnesota specimens assignable to lycaon and only one referable to nubilus.
Because wolves in the known range of nubilus are thought to be extinct, and
because the animals in northeastern Minnesota are legally unprotected and
subject to a control program, it seems highly desirable that the question of
their taxonomy be studied intensively while specimens are still available.

ACKNOWLEDGMENTS

This study was supported by Macalester College, the New York Zoological
Society, the Minnesota Department of Conservation, the U.S. Bureau of
Sport Fisheries and Wildlife, and the USDA Forest Service. Mr. Wallace C.
Dayton and Miss Elizabeth Dayton, and the Quetico-Superior Foundation,
all of Minneapolis, financed Mech during the preparation of this paper. We
would also like to thank Dr. J. L. Paradiso, Dr. H. L. Gunderson, and Mr. M.
H. Stenlund for reviewing this manuscript.

LITERATURE CITED

Goldman, E. A. 1944. The wolves of North America, Part II.
Classification of Wolves. p. 389-636. Washington, D.C.: The
Amer. Wildl. Inst.
Pimlott, D. H., Shannon, J. A., and Kolenosky, G. B. 1969. The
ecology of the timber wolf in Algonquin Provincial Park. Ont.
Dep. Lands and Forests Res. Pap. (Wildl.) 87, 94 p.

SOME RECENT RESEARCH PAPERS
OF THE
NORTH CENTRAL FOREST EXPERIMENT
STATION
Tree Improvement Opportunities in the North-Central States Related
to Economic Trends, A Problem Analysis, by David H. Dawson
and John A. Pitcher. USDA Forest Serv. Res. Pap. NC-40, 30 p.,
illus. 1970.
Relation Between the National Fire Danger Spread Component and
Fire Activity in the Lake States, by Donald A. Haines, William
A. Main, and Von J. Johnson. USDA Forest Serv. Res. Pap. NC-
41, 8 p., illus. 1970.
Thinning and Fertilizing Red Pine to Increase Growth and Cone
Production, by John H. Cooley. USDA Forest Serv. Res. Pap.
NC-42, 8 p., illus. 1970.
The Impact of Estimation Errors on Evaluations of Timber Production
Opportunities, by Dennis L. Schweitzer. USDA Forest Serv.
Res. Pap. NC-43, 18 p., illus. 1970.
User Evaluation of Campgrounds on Two Michigan National Forests,
by Robert C. Lucas. USDA Forest Serv. Res. Pap. NC-44, 15 p.,
illus. 1970.
System Identification Principles in Studies of Forest Dynamics, by
Rolfe A. Leary. USDA Forest Serv. Res. Pap. NC-45, 38 p.,
illus. 1970.
Skiing in the Great Lakes State: the Industry and the Skier, by William
A. Leuschner. USDA Forest Serv. Res. Pap. NC-46, 42 p., illus.
1970.

Proceedings of the Ninth Lake States Forest Tree Improvement
Conference, August 22-23, 1969. USDA Forest Serv. Res. Pap.
NC-47, 34 p. 1970.
A Water Curtain for Controlling Experimental Forest Fires, by Von J.
Johnson. USDA Forest Serv. Res. Pap. NC-48, 7 p., illus. 1970.
Wildness Ecology: A Method of Sampling and Summarizing Data for
Plant Community Classification, by Lewis F. Ohmann and
Robert R. Ream. USDA Forest Serv. Res. Pap. NC-49, 14 p.,
illus. 1970.

ABOUT THE FOREST SERVICE....
As our Nation grows, people expect and need more
from their forests—more wood; more water, fish, and
wildlife; more recreation and natural beauty; more
special forest products and forage. The Forest Service
of the U.S. Department of Agriculture helps to fulfill
these expectations and needs through three major
activities:

Conducting forest
and range research
at over 75 locations
ranging from Puerto
Rico to Alaska to
Hawaii.

Participating with
all State forestry
agencies in
cooperative
programs to protect,
improve, and

wisely use our Country's 395 million acres of
State, local, and private forest lands.

Managing and protecting the 187-million acre
National Forest System.

The Forest Service does this by encouraging use of
the new knowledge that research scientists develop;
by setting an example in managing, under sustained
yield, the National Forests and Grasslands for
multiple use purposes; and by cooperating with all
States and with private citizens in their efforts to
achieve better management, protection, and use of
forest resources.
Traditionally, Forest Service people have been active
members of the communities and towns in which
they live and work. They strive to secure for all,
continuous benefits from the Country's forest
resources.
For more than 60 years, the Forest Service has been
serving the Nation as a leading natural resource
conservation agency.

Transcriber's Notes

This is a compilation of four separate reports, each having their own table
and figure numbers. I have retained the original table and figure numbers
due to all the references made to them within the text. However I did reindex
the footnotes for the complete compilation. I made minor punctuation
corrections, modified the table formats, moved some illustrations, and made
the following typo corrections:

Table of Contents: Changed "Occurence" to "Occurrence".
Originally: The Possible Occurence of the Great Plains Wolf in
Northeastern Minnesota

Page 5: Added missing parenthesis after "individuals".
Originally: the same color (with the exception of a few black or white
individuals (see Mech and Frenzel, page 60)

Page 27, Deleted repeated word "the".
Originally: When still on the the ice about 15 feet from shore,

Page 34, Literature Cited: Changed "Vegetatation" to "Vegetation".
Originally: Ohmann, L. F., and Ream, R. R. 1969 Vegetatation studies in
the BWCA

Page 37: Changed "repreductive" to "reproductive".
Originally: lungs, liver, kidneys, repreductive tracts

Page 40: Changed "wildnerness" to "wilderness".
Originally: while in the wildnerness more males were taken

Page 41: Changed "decidous" to "deciduous".
Originally: The deciduous first incisors of fawns and the decidous

Page 42: Changed "end" to "and".
Originally: from wolf-killed deer end examined grossly in the field

Page 42, Figure 9: Changed "discoverd" to "discovered".
Originally: A permanent first premolar (arrow) was discoverd in M-8.

Page 47: Changed "wildnerness" to "wilderness".
Originally: not surprising that in the wildnerness area

Page 57, Footnote 36: Deleted duplicate "to".
Originally: Personal correspondence to to L. D. Mech, Oct. 10, 1969.

Page 58: Changed "diffference" to "difference".

Originally: Nevertheless, with extremely deep snow, the diffference

Page 59, Literature Cited: Changed "roll" to "role".
Originally: Nasimovich, A. A. 1955. The roll of the regime of snow

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