HomeMy WebLinkAboutAPA2691N E L C H I N A
BY
Warren B : Ballard
Robert o. Stephenson
and
Ted H. Spraker
STATE OF ALASKA
Jay s. Hammond, Govern9r
DEPARTMENT OF .FISH AND .GAME
Ronald 0. Skoog, Commissioner
DIVISION QF .~~
Ronald J.' Somerville, Director
Donald E. 1-Ic:::Kni'ght, aesearch Chief
FINAL REPORT
Federal Aid in Wildlife Restoration
Projects W-17-8, W-17-9, W-17-10, and W-17-11
Jobs 14.8R, 14.9R, and 14.10R
~th additional support from the .Alaska Power Authority
(printed June 1981)
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FINAL REPORT (RESEARC
State: Alaska
Cooperators: Warren Ballard, Robert Stephenson,
Ted Spraker, Sterling Eide, James Foster,
Albert Franzmann, Art Flynn, Dan Holleman,
SuzAnne Miller and John Schlotthauer
QL
"131
. c'LL
6 35
Project Nos.: W-17-7, W-17-8,
W-17-9, W-17-10 and
W-17-11
Project Title: Big Game 1981
Investigations
Job No.:
Job No.:
Job No.:
Period Covered:
14.8R
14.9R
14.10R
Job Title:
Job Title:
Job Title:
April 1, 1975 to June 30, 1980
SUMMARY
Wolf Populations
and Movements
1n Relat1on
to Those of
Prey Spec1es
Wolf Food Habits
Impact of Wolf
Predation Upon
Ungulate Populations
Between 1 April 1975 and 30 June 1980, 103 individual
timber wolves representing up to 22 different wolf packs were
captured and radio -collared in Game Management Unit 13 (Nelchina
Basin) of southcentral Alaska. Cost and problems associated
with helicopter darting are discussed.
The 103 radio-collared wolves were individually located on
3, 525 separate occasions resulting in 6, 927 wolf sightings.
Pack and individual wolf histories are presented and discussed.
Wolf territories were, for the most partr nonoverlapping.
Overlaps which did occur were seasonal in nature or due to the
method by which territqries were plotted. There appeared to be
changes in terri tory boundaries from year to year. Terri tory
sizes 2for 14 intensively ~tudied packs ranged 2from 268 2 to
864 mi (691 to 2738 km ), averaging 537 mi (1390 km ).
Terri tory size appeared to be larger for larger packs and for
those packs in areas of low moose density.
Wolf den and rendezvous site usage is described. The
earliest that radio-collared wolves were observed at a natal den
. was 13 April. Most wolves began visiting den sites in late
April and early May. Parturition appeared to occur throughout
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the month of May. Natal dens were abandoned between 4 June and
1 August. Pups were observed first traveling with adults from
late August to mid-September of each year.
· Radio-collared wolf packs were observed on 360 individual
prey kills, 38 ( 10. 6%) of which were also occupied by one or
more brown bears. Moose of varying ages comprised 72 percent of
the observed kills. Calf and short yearling moose comprised 20
percent of the total kill. Wolves were preying upon short and
-long yearling age classes from January through July
disproportionately to their presence in the moose population.
Moose calves 0-6 months of age comprised only 6 percent of the
kills observed.
Four thousand two hundred and ninety food items were
identified in 3,624 wolf scats collected at den and rendezvous
sites during this 5-year period. Overall, calf moose was the
most frequently identified food item (44%). Percent occurrence
of various prey i terns in wolf scats was generally related to
prey abundance. Occurrence of calf moose in scats was
correlated with subsequent fall calf:cow ratios, suggesting that
wolves were preying upon calf moose in proportion to their
abundance. Scat data were converted to numbers of individual
prey eaten which was then extrapolated to GMU 13 spring wolf
population estimates. This analysis suggested that wolves in
GMU 13 were preying upon from 434 to 1,013 moose calves annually
from mid-May through mid-July.
One hundred and twenty-five moose and 25 caribou kills were
examined in situ to determine both cause of death and age and
physical condition of prey taken b y wolve s . On e i n sta nce of
surplus killing o f caribou b y wolve s was repo rted . Perce nt
marrow fat of calf and short yearling moose ki lled by wolves was
significantly higher than that of calves dying from both
accidental causes and winter kill. We concluded that wolves
were preying upon relatively healthy calf and short yearling
moose.
Age and condition of wolf-killed adult moose examined from
1970-1972 were combined with data collected during this study.
Overall, ages of adult moose killed by wolves were significantly
(P<O.OS) different from those tagged moose, winter-killed moose,
and moose dying from accidental causes.
Age and condition of wolf-killed adult moose were compared
with those of moose tagged during the same year pre dation
occurred. We .concluded that during severe winters wolves preyed
upon adult moose in proportion to their occurrence in the moose
population, while during average or mild winters relatively
older adult moose were being preyed upon .
Marrow fat percent of wolf-killed adults was significantly
(P<O.OS) higher than that of winter-kill e d moose, but not
significantly (P>O.OS) different from those dying of accidental
causes. We concluded that wolves were preying upon relatively
healthy adult moose during winter.
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Overall, 17 wolf packs averaged a kill every 4 .9 days with
a range of 3.1 to 12.7 days per kill. Differences between packs
and problems associated with methods of calculating predation
rat~s are discussed.
During winters 1978-79 and 1979-80, five wolf packs were
intensively monitored to determine rates of kill according to
pack size. Ungulate kill rates varied from one kill/8. 3 days
for a pack of two to one kill/3. 6 days for a pack of nine
wolves. Large wolf packs generally appeared to have a higher
kill rate than smaller wolf packs .
During summers 1977 and 1978, activity patterns of two
denning wolf packs were studied and are discussed. It was
~determined that adult males (presumed to be alpha males) were
nearly always present when ungulate kills were made.
Wolf densities in GMU 1~, excluding the wolf removal area,
varied from ~wolf/37.6~ {97.3 km) in fall 1975 to
1 wolf/121.7 mi (315.2 km ) in spring 1978. Wolf numbers in
GMU 13 have declined since 1975. Hunting, trapping -and
dispersal were ·identified as the main reasons for the decline.
GMU 13 wolf densities were compared with those reported
elsewhere in North America .
Annual GMU 13 wolf harvests are presented and discussed.
Rates of harvest from individual radio-collared wolf packs were
exami ned in relation to productivity and ab ility of packs to
replace losses. Losses in excess of 41 percent of the fall
population resulted in pack population declines the following
fall. It was suggested 2 that a post hunting-trapping wolf
density of 1 wolf/100 mi might be suitable to keep wolf
predation on moose to a minimum yet maintain a reasonable vlolf
population .
Of 103 wolves radio-collared during this study, 14 ( 14%)
were known to be alive on 30 June 1980. Twenty-five percent
were also known to have dispersed during the 5 years of study.
The largest source of wolf mortality was human induced (77%).
Ground shooting and suspected illegal aerial hunting, accounted
for 76 and 11 percent, respectively, of the man-caused
mortality. Natural forms of mortality accounted for 23 percent
of the mortalities.
During this study at least 26 radio-collared wolves were
known to have dispersed from their original pack area.
Sixty-eight percent of the dispersals were males. Approximate
average age of dispersed males was 35 months 1 while females
averaged 37 months. Dispersal was most prevalent . from April
through June. Average distance dispersed was at least 6 7. 7
miles (106 km). The longest documented movement was 460 miles
(736 km), constituting a record movement for this species.
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The effect of wolf predation on moose calf survival was
studied in two areas of GMU 13. In one area, referred to as the .
Susitna River Study Area, wolf densities were lowered by
Department personnel. In the other area of GMU 13 (remainder of
the · Unit generally east of Talkeetna Mountains) , wolves were
intensively studied to enumerate population den sities and food
habits.
From January 1976 through July 1978, 60 wolves were killed
by Department personnel in an effort to test the hypothesis that
lowered wolf densities would improve moose calf survival. Wolf
densities il2 the Susitna River study area were estimated at
1 wolf/98 mi of habitat in spring 1975 to 1 wolf/232 mi of
habitat in spring 1978. By spring 1980 wolf densities had
increased to within at least 89 percent of the spring 1975
estimate due to reproduction and immigration .
Fall moose sex and age composition count data and annual
harvests were compared between the wolf removal area and other
comparative count areas in GMU 13 where Department wolf removals
had not been conducted. Statistical analyses revealed no
significant (P>O.OS) differences in either calf-cow ratios,
moose observed per hour of survey, nor in ratios of harvested
moose. Had wolf control increased moose calf survival we would
have anticipated some significant differences in these ratios.
Results of wolf food habits, moose calf mortality, and bear food
habits studies indicated that the rates of ·predation on moose
cal ves by wo l ves we re far l ess than by brown bears . This tended
to explain the lack of response by the moose popul ation to
reductions in wolf densities. Results of the bear transplant on
moose survival are compared and discussed with this study.
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CONTENTS
Summary. . . . . . . . . . . . . . . i.
Background . . .
Objectives . . . .
Procedures . . . . . . . . . .
Results and Discussion . . . . . . . . . .
Wolf Pack and Individual Histories ..
Wolf Territories. . . . . . . . . .
Den and Rendezvous site Usage . . . . .
Food Habits in Relation to Availability
Food Habits Summary . . . . .
Age and Condition of Prey .... .
Wolf Predation Rates ... .
Summer Activity Patterns ...... .
GMU Unit 13 Wolf Densities .. .
Trapping and Hunting Mortality.
Wolf Dispersal. . . . . . . . .
Experimental Wolf Removal . . . . . .
Wolf Repopulation of Removal Area .
Evaluation of Wolf Removal on Moose .
. • . 1
• • • • 2
2
• • • 6
• • • 9
• • • • J6
• • • • • • 3 9
of Prey . . 41
• • • • 82
• • • • 98
• 99
• • ·120
• • ·126
• • • • ·128
• • • • ·14 9
• • • • ·152
• • . • ·157
. • • • ·157
Recommendations_. . . . . . • .
Acknowledgements . . . . . .
• • • . • ·171
. • • . • • • ·171
Literature Cited . . ..... . ·173
BACKGROUND
Rausch (1969) su.'l'['.marized the status of wolves (Canis lupus)
in the Nelchina Basin (Game Management Unit 13) for the period
1957 through 1968. From 1948 to 1953 extensive poisoning by the
Federal Government reduced populations of predators to low
levels. In 1953 only 12 wolves were believed to remain in the
Nelchina Basin. Rausch believed the wolf population gradually
increased after 1953 and reached a peak of 400 to 450 in 1965.
Although no formal wolf studies were conducted from 1969 through
1974, Mcilroy (1976) speculated that a second peak occurred in
1970 after a low of 300 animals in 1968.
Rausch (l969L Bishop and Rausch, (1974), and Mcilroy
(1974) described the history of the Game Management Unit 13
moose (Alces alces) population. All pointed to an apparent
inverse relationship between numbers of predators and numbers of
ungulates. Moose apparently began declining after the severe
winter of 1961-62. This decline continued and was hastened by
severe winters in 1965-66, 1970-1971 and 1971-72. Fall calf:cow
ratios declined sharply and reached a record low for the Basin
in 1975. Although wolf predation was not suggested as the main
reason for the population decline, it was thought to have at
least amplified the decline and, more importantly, prevented
recovery during mild winters (Rausch et al. 1975). This concern
coupled with the findings of Stephenson and Johnson (1972,
1973), which revealed a high percentage of calf moose in wolf
scats, suggested that wolf predation on calves was preventing
the moose population from recovering. Consequently a series of
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studies was initiated to obtain information on wolf-moose
relationships in the Nelchina Basin. Preliminary results of
this study were presented by Stephenson (1978) and by Ballard
and Spraker (1979).
OBJECTIVES
To delineate wolf pack territories and determine wolf ~
densities in an experimental area and a control area in Unit 13.
To compare seasonal wolf movements with seasonal movements
and abundance of major prey species and environmental
parameters.
To assess wolf food habits in the experimental and control
areas of Unit 13.
To quantitatively assess the impact of wolf predation upon
ungulate populations in Unit 13.
PROCEDURES
Wolves were studied in two different portions of Game
Management Unit 13. In the susi tna River Study Area (Fig. 1}
wolf numbers were experimentally reduced with the aid of
helicopter and fixed-wing aircraft to assess the effects of wolf
predation on moose calf survival. Moose calf survival there was
to be compared to survival in the Nelchina Study Area, where
wolf numbers had not been reduced.
Game 2 Management ~it 13, · an area of approximately
23,782 mi , has 7,258 mi over 4,000 foot elevation. Detailed
descriptions of its vegetation, climate and geography were
provided by Skoog ( 1968).
The Susi tna River Study Area is located in the northern
portion of the Unit (Fig. 1). Its boundaries are as follows:
the Alaska Range on the north; the Maclaren River on the east;
the Maclaren and Susitna Rivers on the south; the confluence of
Deadman Creek with the Susitna River northward to headwaters of
Brushkana Creek, downstream to Brushkana Creek's confluence with
the Nenana River and then northwest upstream to the Alaska range
on the west. The Nelchina study area generally comprised those
portions of Game Management Unit 13 lying east and south of the
Susitna River study area.
Wolves were captured for radio-telemetry studies with a
Cap-Chur gun and dart "(Palmer Chemical Co.) fired from a Jet
Ranger 206B helicopter using methods similar to those described
by Baer et al. (1978). Our capture technique differed from
theirs in that we darted in all types of vegetative cover and
also while the animal was moving. Initially, a drug mixture of
0.7 ml, 100 mgjcc of phencyclidine hydrochloride (Sernylan,
Parke-Davis Co.} with 1.0 ml, 100 mgjcc of promazine
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hydrochloride (Sparine, Wyeth Laboratories) was used to
immobilize wolves. Later, the dosage of Sernylan was increased
to 1. 0 ml ( 100 mgjcc) to decrease the the latent period and
obtain more complete immobilization. Twelve wolves were also
immobilized with 2 mg of etorphine (M-99, D-M Pharmaceuticals,
Inc., Rockville, MD). After being processed and radio-collared
these wolves received an equivalent cc dosage (2 mg/ml) of the
antagonist diprenorphine (M-50-50, D-M Pharmaceuticals, Inc.,
Rockville, MD) injected into the radial vein. Four live wolves
were purchased from trappers and radio-collared.
Captured wolves were initially equipped with an adjustable
machine belt radio collar manufactured by AVM Instrument Company
(Champaign, IL) and later with an adjustable collar made of
fiberglass and urethane manufactured by Telonics (Mesa, AZ).
Hair and blood samples were taken from each wolf using
methods similar to those described for calf moose
(Ballard et al. 1979). All hair samples were sent to
Dr. Arthur Fly11n; Case Western Reserve University, Cleveland,
Ohio, for mineral element analysis. · When practical, the
following body · measurements were recorded: weight, total
length, heart girth, chest height, neck circumference, shoulder
height, tail length, and length of canines. Preliminary
analyses of hair and blood data were presented in Ballard and
Spraker (1979). Results of these studies will be combined with
those obtained from the Kenai Peninsula wolf study (Peterson
1978) and prepared for publication. Data reflecting blood and
hair parameters not yet analyzed, are presented in Appendices I
and II, respectively.
Initially we used a portable radiotelemetry receiver
manufactured by AVM Instrument Company. The receiver contained
four bands with 12 channels per band and covered frequencies in
the 150.000 to 152.000 MHz range. Later, we began using a
programmable scanning receiver manufactured by Telonics (Mesa,
AZ).
Radio-collared wolves were tracked and, when possible,
visually observed from fixed-wing aircraft using the methods
described by Mech (1974). Monitoring intensity varied from pack
to pack but consisted of at least bi-monthly monitoring during
winter months.
Approximate ages of captured wolves were determined on the
basis of tooth eruption and wear. Wild wolf age estimates were
based upon their relative size and by criteria described by
Jordon et al. (cited by Mech 1970). In some cases, age and sex
structures of certain packs were not ascertained until the
animals had been killed by hunters or trappers. We encouraged
hunters and trappers to provide us with wolf carcasses taken in
Units 11 and 13 by offering $10.00 per carcass. Ages of
harvested wolves were determined by tooth eruption and wear, and
by examining epiphyseal cartilage of the long bone according to
methods described by Rausch (1967).
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The sex and age of moose and caribou (Rangifer tarandus),
killed by wolves were often determined from fixed-wing aircraft
based on size, pelage and antler growth. Moose kills were
categorized as calves, yearlings or adults. Both calves and
yearlings were aged to the nearest month using an assumed
birthdate of 1 June.
Size of wolf territories was determined by plotting all
radio locations for individual packs and then connecting the
outermost observations (Mohr 1947). Locations for individual
radio-collared wolves which had dispersed were not included.
Sizes of wolf territories and study areas were determined with a
compensating polar planimeter. All study areas and wolf
territories were pl~met~ed at least three times and then
averaged to compute mi (km ).
Active wolf dens located through observations of
radio-marked wolves, or during associated flying, were inspected
on the ground after they were vacated by wolves. The vicinity
of each den was searched and all scats collected and food
remains identified. scats were placed in individual paper bags,
then autoclave_d and analyzed using previously described
techniques (Stephenson and Johnson 1972), except that hair scale
impressions ( Adorj an and Kolenosky 1969) were used to confirm
identification of prey remains when identification was
uncertain. Comparisons of hair scale impressions were made with
known samples by imprinting them on a slide containing clear
fingernail polish.
When practical, we examined wolf kills on the ground. We
determined the cause of death according to methods described by
Stephenson and Johnson (1973} and Ballard et. al. (1979). A
femur or metatarsal and the mandible were collected from each
kill to aid in establishing the animals' physical condition on
the basis of percent marrow fat using methods described by
Neiland ( 1970). Ages of moose killed were determined on the
basis of tooth eruption and cementum annuli, using methods
described by Sergeant and Pimlott (1959). Caribou were aged on
the basis of tooth eruption and wear (Skoog 1968}.
During winters 1978-79 and 1979-80 an attempt was made to
locate and examine all kills made by selected radio-collared
wolf packs during a 2-3 month period. These selected packs were
radio-tracked at least every other day and were backtracked to
their previous location. In an effort to maintain study packs
at stable numbers portions of GMU-13 were periodically closed to
hunting and trapping of wolves. Boundaries and season lengths
of closed areas were described by Eide (1979) and Tobey (1980).
The effects of reductions in wolf density on calf and adult
moose survival in the Susitna River study area were determined
by several methods . Fall moose sex and age composition counts
were used to compare calf: cow ratios in areas of high and low
wolf densities. If wolf reductions had increased moose
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survival, it was anticipated that the increase would also be
reflected by increases in the sport harvest of bull moose.
Moose harvests were determined by tabulating the numbers of
successful hunters as determined from 11 mandatory 11 harvest
reports. Statistical differences between annual calf: cow and
harvest ratios involved arcsine transformation of observed
ratios in calculations of the statistics (Sokal and Rohlf
1969:607). Comparisons of trends in calf:cow ratios within and
between areas involved analysis of residuals (Everith 1977:46)
to determine which cells of a chi-square table were significant
in rejection of the null hypothesis. The latter analysis was
conducted using BMDP canned programs (University of California,
Los Angeles, Dept. of Biomathematics, School of Medicine).
Statistical differences in means and deviations in ratio data
were tested with t-tests and chi-square analyses (Snedecor an9
Cochran 1973), respectively.
Carcasses of wolves removed from the exoerimental area were
necropsied and aged, and nutritional condition and reproductive
status were assessed. Preliminarv results of laboratory
examinations of these and other wolves, including those taken
during 1976 and .1977 in control programs in GMU-20A and in
northwestern Alaska, have been presented by Nielson (1977).
Additional results from these studies are being analyzed and
will be presented in future publications. Stomach and
intestinal tracts were submitted to John C. Schlotthaver,
University of Minnesota, for parasite investigations. Results
of this study are currently being prepared for publication
(Averbeck, et al. In Prep.). Tissue samples from the
above-mentioned wolves have also been radio-assayed for
radiocesium ( cesium-137) by Dan Holleman, Institute of Arctic
Biology, University of Alaska, Fairbanks, in an effort to
elucidate the relative importance of moose and caribou in the
diet of wolves. Results of portions of these analyses are also
being prepared for publication (Holleman and Stephenson In
Press).
RESULTS AND DISCUSSION
One hundred and three different wolves were captured and
radio-collared in GMU-13 from April 1975 through June 1980.
Sixteen wolves were recaptured on one or more occasions. All
wolves, with the exception of two mortalities (wolf #122062 and
one unlisted), appeared to recover from the drugs within several
hours and most returned to the pack within 12 hours of initial
immobilization. One collaring mortality resulted from drowning
which apparently occurred as the animal was recovering from the
drug in deep snow, while the other resulted from hitting the
animal between the vertebrae with the dart. Wolves immobilized
with etorphine recovered within 60 seconds after the antagonist
was administered.
Costs associated with radio-collaring wolves in GMU 13 were
discussed in Ballard and Spraker ( 1979). Costs consisted of
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charter charges for helicopter and fixed-wing aircraft and
darting equipment but did not include manpower nor the price of
the radio collars. Based on 1978 prices, an average of $578 was
expended to radio-collar each wolf. Wolves from packs that did
not alreaqy have at least one radio-collared member were more
costly ( x = $624. 00 per animal) than those which did
(x = $511.00 per wolf). This difference was reflected in
increased search time with fixed-wing aircraft. Four trapped
wolves were purchased from trappers and radio-collared for
approximately $350.00 each. Two of these animals had been
injured by the traps and appeared to have difficulty keeping up
with other pack members. Nevertheless, trapped animals were
useful in establishing contact with a pack so additional members
could be subsequently radio-collared from a helicopter.
Our helicopter darting technique was most efficient in
moderate to sparse spruce habitat and when the depth of powdered
snow exceeded 1 foot. These conditions usually slowed the
animal and allowed the helicopter to get within easy shooting
range. We experienced difficulty capturing wolves in dense
spruce habitat and on open tundra with hard snow pack. Dense
spruce often allowed an animal to evade the helicopter and
required many shots for each successful hit. Open tundra
allowed a wolf considerable maneuverability at high speeds and
although we were always successful, it was usually necessary to
tire the animal by running it.
From· April 1975 through June 1978 the 103 radio-collared
wolves representing up to 22 individual packs (a pack is defined
as two or more wolves) were located on 3,525 separate occasions
resulting in 6,927 sightings (Table 1). These observations
represent 1,805 wolf pack days (pack day is defined as any day
on which a pack was located one or more times) . Monitoring
intensity was variable from pack to pack, being dependent upon
proximity to the field station, short-term objectives of the
project (i.e., predation rates study), reliability of
radio-transmitters, and loss of radio-collared wolves to hunting
and trapping.
Observability of individual packs was also variable,
ranging from 14 to 95 percent and averaging 84 percent. Pack
observabili ty appeared to be primarily a function of different
habitat types and individual wolf behavior. Wolves occupying
thick spruce habitats were less frequently observed than those
occupying open areas. Some wolves seemed to be evasive when we
attempted to observe them from aircraft. Also, small packs (2-3
individuals) were harder to observe than relatively large packs
(5 or more individuals). Observability was also influenced by
the experience of the pilot and observer.
We had a disproportionate number of visual sightings of
certain wolves. Most evident was our inability to make visual
contact with both black and white wolves, especially black
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Table 1. Summary of location observations of radio-collared wolf packs l in the Susitna and Nelchina River Basins of southcentral
Alaska from April 1975 through June 1980.
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Combined Number
Combined number different 0 number individual Number pack Number
radio-collared radio wolf radio pack
Pack wolves location sightings location days --
Brushkana 1 31 109 31 34 .
Butte Lake 1 23 41 23 31 -~ Coal Creek 1 19 18 16 17
Deadman 2 63 67 47 38
Deep Lake 6 210 343 183 145
Delta 4 27 43 24 20
,-
Ewan 5 181 354 164 151 L
Gakona 3 11 13 8 8
Hogan Hill 5 306 746 225 176 r Jay Creek 1 47 49 41 43 ' Keg Creek 8 568 958 287 240 [_
Maclaren 1 51 119 51 52 r--
Mendeltna 4 391 913 151 128
Middle Fork 5 49 60 35 27 ~--
Saint Anne 11 181 353 97 76
Sinona ., 304 844 """ 170 I !00
Stephan Lake 1 24 16 24 24
Susitna 11. 452 700 129 135 L
Tolsona 5 153 542 98 89 ,......
Tsusena 4 16 7 1 1
Tyone 14 359 568 185 170 ~--=-
Watana 5 57 64 29 30
Totals 10s.!/ 3,525 6,927 2,035 1,805 b
y Does not correspond with total number of wolves radio-collared
because some wolves became members of other packs.
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wolves during snow-free periods and white wolves during periods
of snow cover. This suggests that the numbers of wolves of
these color phases could be underestimated during aerial
surveys.
Histories of individual radio-collared wolves and their
respective packs are described in the following section.
Individual radio-collared wolves are identified in the text by
the last three digits of their assigned accession number.
Wolf Pack and Individual Histories
Butte Lake Pack -The Butte Lake pack included one collared
wolf (#984) captured while traveling alone near Butte Lake in
late April 1975. An adult female (#983), member of the
Brushkana pack, was collared the same day 3 miles (5 km) from
wolf 984, and tracks suggested that both wolves had been feeding
on an old moose kill in the area. Subsequently, wolf 984 was
accompanied by three black wolves and later ~y one blac~ and one
gray wolf and inhabited an area of 1,188 mi (3,077 km ) almost
totally separate from the range of the Brushkana female. The
associates of the Butte Lake male did not include female wolves,
and a den was not located. The summer movements of this group
were much more extensive than those of denning packs. No scats
representing the summer diet were collected, but the small
number of carcasses located suggested that both moose and
caribou were used by the pack. Relatively few radio locations
were obtained during early summer 1975 due to poor weather and
the mountainous terrain inhabited by the pack during part of the
study period. Three males were removed from this pack in
January 1976 as part of the wolf removal experiment.
Brushkana Creek Pack -The Brushkana pack included only one
radio-collared member, an adult female (#983). At least three
and possibly four adults were associated with this pack, and six
pups were reared in 1975. An adult male was shot within the
territory of the pack by a local trapper in October 1975,
constituting the only known mortality prior to the pack's
removal in connection with the wolf reduction experiment in
January 1976 . Based on the movements of the c?t-lared fema~e,
the pack inhabited an area of at least 468 mi ( 1, 212 km ) .
Coal Creek Pack ·-This pack consisted of an uncollared
yearling gray male and wolf 002 which was originally
radio-collared as a member of the Keg Creek Pack in April 1975.
The origin of the yearling male is unknown. Shortly after being
radio-collared, wolf 002 exhibited a tendency to travel
independently of the other Keg Creek pack members during late
winter and summer 1976.
In mid-January 1976 wolf 002 was often found alone but
stayed for the most part within the Keg Creek boundaries. On
8 March 1976 wolf 002 was located near the center of the Hogan
9
Hill wolf territory (Figs .. 2 and 3) and was accompanied by the
radio-marked Hogan Hill male ( #988) . On 9 March these two
wolves were observed traveling within a few miles of each other
in the northern portion of the area where the Hogan Hill and Keg
Creek territories overlapped. On this date, wolf 988 was
accompanied by another wolf, presumably also a Hogan Hill pack
member. on the morning of 12 March 1976, wolf 002 and wolf 988
were again found traveling together in the area. Observations
during the remainder of March showed that these wolves
subsequently returned to their respective packs and territories.
However, wolf 002 tended to remain some distance from the four
remaining Keg Creek pack members during late March and April.
On 22 April 1976 she was located in the southern portion of the
Ewan pack terri tory, 49 miles ( 80 km) south of the Keg Creek
den. She returned to the terri tory by 28 April but showed a
tendency to remain apart from other pack members and in late May
again left the Keg Creek territory. On 29 and 30 May, the wolf
was located in the Susitna River study area north of the
Maclaren River, 24 miles (39 km) north of the 1975 den.
Extensive aerial searches during the next 2 weeks failed to
locate the wolf, but on 13 June 1976 she had moved west across
the susitna River to the vicinity of Coal Creek. It is probable
that during this time, wolf 002 was either in the Watana or
Clearwater Mountains (which could not be adequately searched due
to inclement weather).
On 26 June wolf 002 and an uncollared gray male were
observed 8 miles (13 km) east of the 13 June 1976 location, east
of the Susitna River on Clearwater Creek. By 28 June the pa1r
had moved 29 miles (47 km) west to the Jay Creek area where they
were observed to detect and pursue the radio-marked Tsusena
female (#199) which in mid-June had also entered the
experimental area. Details of this encounter are provided in
the section on the Jay Creek Pack. On 30 June the pair was
located in the Watana Creek drainage, 8 miles (13 km) east of
the den used in 1975 by the previously removed Deadman pack, and
on 2 July they were found on the south side of the Susitna River
near Kosina Creek.
Our observations of these wolves during June ~d July 1~76
indicated that they maintained a territory of 315 mi (816 km ).
They were not associated with a den during this period. During
July 1976 they were only observed on two kills; one of unknown
species and one beaver (Castor canadensis). Both wolves were
killed in late July as part of the experimental wolf removal
program.
Deadman Lake Pack -This pack inhabited a relatively small
area in the vicinity of Deadman and Watana Creeks north of the
Susitna River. An adult male (#981) and female (#982) were
collared in April 1975 and three pups were reared that year.
Although radio locations suggested the presence of a den on a
brushy hillside, none was located despite an extensive ground
search. Two rendezvous sites were located, and scats were
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• . • • Boundary of experimental area
Boundary of control area
Observed boundaries of radioed 1-----pack territories
1. Sinona Ck.
2. Ewan Lake
3. Deep Lake
4. Hogan Hill
5. Middle Fork
6. Delta R.
7. Keg Ck.
8. Mendeltna Ck.
9. Maclaren R.
10. Butte Lake
11. Brushkana 16. Lower Gakona
12. Deadman Ck. 17. Jay Ck.-Coal Ck.
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13. Oshetna R. Circles indicate general
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14. Tsusena Ck. of nonradio-marked packs ineluded
15. Stephan Lake in population estimates.
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F~gure 3. Wolf pack territories derived from monitoring radio-collared animals in Game Management Unit 13
of Southcentral Alaska from 1 July 1976 through June 1978.
St. Anne's ----.J<..JV<..J<..AAIVV\
Ewan ----------~~~X.c>~
Deep Lake -----: ::: :: : ::::: : :
Mendeltna -----·.,. • ~ ._,-~: :~:
Tyone River ----~~~~~ Keg Creek -----1:111111111111
Hogan Hill ----.. -.. ,. .. " "' ...
S inona Ill -----';. v ';..' /-; '-/.
Sinona 112 -----::::::::::::::::::::::::::
Middle Fork ---Jjjjj-tDj
De 1 ta ---------:!:t::~:;g;:::::::::\'.'-\'~~
Tyone male ----=:~·:.•;.~~~··:·~·::~
Coal Creek ----::: · . : .
Jay Creek -----~---------~
Wa tana --------H-:i:'t~l:-t~l:t :1~
Stephan Lake --:··•:··~·:··•:~·
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collected at one of these. The three pups were removed in
January and the two adults in March 1976 in connection with the
wolf removal experiment. The ?2'me rang~ suggested by radio
locations, included only 108 mi (280 km ) • The small home
range may be the result of the small number of radio locations
obtained. However, the pack was small in number and inhabited
an area with relatively abundant and varied food resources.
Deep Lake Pack -Information on the numbers and movements
of the Deep Lake pack was not obtained until March 1976 when two
adult wolves (#'s 067 and 993) were radio-collared. An active
den, at which an unknown number of pups were reared, was located
in the area in June 1975 however, and signs indicated the
presence of the pack in the area during early and mid-winter
1974-75. For unknown reasons wolf 993, collared at the same
place as wolf 067 in March, subsequently inhabited an area
southeast of the observed home range of the other seven members
of the Deep Lake pack, remaining in the southern one-third of
the adjacent Ewan pack terri tory. Wolf 993 was never observed
with other wolves. We lost radio contact with her in late
September 1976 at her most southerly radio location along the
Trans-Alaska Pipeline, 2 mi south of the Tazlina River. We
suspect she moved south along the Trans-Alaska Pipeline into the
Chugach Mountain range where her transmitter eventually failed.
In July 1976 the Deep Lake pack consisted of at least four
and probably five wolves. We suspected there was a den site
approximately 2 mi southeast of Swan Lake but were unable to
find it. Subsequent sightings and examination of carcasses
turned in by trappers indicated that the pack probably did
reproduce in 1976.
In late July 1976 wolf 067 began traveling alone near the
·northeast edge of the pack's territory. He was last observed in
mid-August 1976, 2 mi south of Lone Butte, approximately 18 mi
west of the Deep Lake pack terri tory. In March 1977 he was
killed with members of the Keg Creek pack, having become a
member of that pack.
Contact with the Deep Lake pack was reestablished in
October 1976 when one wolf was captured and radio-collared
(009). Five wolves, including two judged to be pups on the
basis of size, were present. During winter 1976-77 ground
shooting, and perhaps one dispersal, reduced the pack to two by
spring 1977.
After February 1977 we had no contact with the pack except
for track sightings and public observations until November 1977.
At that time the pack numbered eight (three adults and five
pups) indicating they had reproduced. Wolf 009 was recaptured
and one additional male (#095) was radio-collared. The location
of the den site was unknown, however, the 1975 site described by
·Stephenson (1978) was not used. During winter 1977-78 ground
shooting reduced the pack to one adult and two pups . The pack
1 3
did not den in 1978. In June, wolf 204 began periodically
leaving the Deep Lake Territory and was last observed with the
pack in November 1978.
The two remaining members (wolves #009 and 1 uncollared
yearling) of the Deep Lake Pack continued to occupy the
traditional pack terri tory through fall 1978 and early winter
1979. In late October, one black wolf, which was subsequently
radio-collared (#212), began periodicaly traveling with the
pack, but he also demonstrated an affinity for traveling with
three other wolves which we suspected were members of the
original Ewan pack. These latter wolves were regularly seen on
the Copper River near Gulkana during winter 1978-79.
In late February 1979, wolf 009 was killed by the Susitna
wolves in the eastern portion of the Deep Lake territory. The
Susi tna wolves returned to their terri tory shortly after this
incident and, thus, were probably trespassing. At the time,
wolf 212 was with the Ewan wolves. The fate of the uncollared
yearling which accompanied wolf 009 is unknown.
During March 1979, radio contact with wolf 212 was lost due
to radio failure and we subsequently lost contact with the Deep
Lake pack until winter 1979-80. Sign and reports from trappers
indicated that very few wolves remained in the Deep Lake
terri tory. In March 1980, wolf 212 and a young gray female,
suspected to have been. the wolf accompanying wolf 009 the
previous year, were both ground shot at Minnesota Lake, within
the Deep Lake terri tory. We suspect that by early spring 1980
the Deep Lake terri-t::ory was vacant.
Annually from 1975 through early 2 1979, the ~eep Lake pac~
occupied fi.. terri tory of from 392 mi ( 1, 015 k.,."ll. ) to 701 mi
( 2, 214 km ) • The largest terri tory occurred from 1975 through
early 1978. Following the death of wolf 009 in February 1979,
examination of location sightings of the neighboring wolf pazk
suggests that this territory was considerably smaller {392 mi }
in 1979-80. If the pack was indeed only comprised of two
wolves, then it seems probable that a smaller hunting area would
be necessary.
The den site occupied by this pack in 1975 was not used
from 1976 through the denning season in 1980.
Delta River Pack The Delta pack included only one
collared wolf, an adult male (#997),2 during 1~75-76. The pack
inhabited an area of about 288 mi (746 km ) in the Delta
River-Tangle Lakes area and its range appeared to partially
overlap the range of the Middle Fork wolves to the south. As
many as six adults may have been present in early summer 1975.
In June 1975, an adult lactating female was killed by an
automobile near Summit Lake and a yearling male wolf was shot in
defense of life and property at the same location in early July
1975. These two wolves and another had often been seen near a
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lodge on Summit Lake during a period of 3 weeks, and the two
that eventually died were described as being in very poor
nutritional condition, lacking caution toward humans and
appearing sick. The description of the adult female matched
that of a wolf seen at the Delta River den in early June 1975,
but which subsequently disappeared. Four pups were successfully
reared at the den, however, and four adults were seen with the
pups in early winter 1975. In mid-November, wolf 997 was found
dead, apparently from natural causes, on Phelan Creek. Necropsy
revealed no signs of physical injury and, although in poor
nutritional condition, the wolf was not totally emaciated. At
least two pups, both in poor nutritional condition, were trapped
in early winter 1975. Little is known about subsequent
movements of the remainder of the pack, and no signs of activity
were found at the den or in the surrounding area in late winter
or early summer 1976.
The Delta pack inhabited an area which had a very low
year-round density of big game prey species which may account
for the poor physical condition of pack members. For exaT.ple,
the four 1975 pups appeared small throughout summer, and
examination of two pups trapped in early winter showed that they
were small for their age and in very poor nutritional condition,
weighing 50 and 55 pounds (23 and 25 kg), respectively. It
appears that most of the 1975 pack members died or dispersed
during winter 1975-76, leaving the territory without a denning
pack.
From spring 1975 through June 1980, very little additional
information was collected on this area except for public
sightings and track counts. Radio contact with wolves in this
area was maintained only between 21 March 1977 and 5 May 1977
through eight locations of two adult radio-collared wolves (#063
and 064}. Both radio-collared wolves were observed together on
three occasions, but neither was observed with other wolves.
Contact with these animals was lost in May 1977 and at the time
we suspected radio failure. However, we never saw additional
sign of wolf activity in the Delta pack's area during several
hours of surveying from fixed-wing aircraft under ideal snow
conditions, and after several checks at the old den site. It
was unknown whether these wolves were members of the original
Delta pack or if they represented new wolves attempting to
colonize an area.
In March 1979, wolf 064 was located on the Wood River in
GMU 20A and shot by Department personnel as part of a wolf
control program. At the time, wolf 064 was accompanied by an
uncollared gray male. Comparison of color and weight indicated
that it was not wolf 063. Further discussion of this wolf's
movements is provided in the dispersal section of this report.
Two wolves were reportedly harvested from this area in
1976-77, but none were reported during the 1977-78 season. Two
wolves were observed on the Denali Highway at Mile 10 by a
1 5
member of the public in March 1978. We suspect that the
terrain, sparseness of vegetation, low density of available prey
and snow conditions make wolves occupying the Delta area highly
susceptible to trapping and ground shooting. Therefore,
post-winter wolf densities have probably been consistently low
in this area during recent years.
Ewan Lake Pack Radio contact with this pack was
established in April 1975 when one adult female (#990) was
captured and radio-collared. Two additional wolves, an adult
gray male (#991} and a male pup (#992}, were radio-collared in
November 1975. The pack denned in 1975 and included at least
seven adults and four pups during most of the summer 1 however,
one pup was apparently lost to unknown causes in August.
There were indications of relatively poor nutrition in this
pack; thoughout summer 1975 the Ewan pups appeared smaller than
those observed in other litters with the exception of the Delta
litter. Also the male pup (#992), collared in November, was
small and thin while the old adult male ( #991) was in fair
nutritional condition.
During winter I wolf 990 tended to remain apart from the
remainder of the pack but was usually accompanied by one and
sometimes two associates. This wolf was regularly found with
the pack only during November. During December and January, it
was accompanied by one and during February 1976 by two wolves.
They were last observed in the Ewan terri tory on 25 February
1976 after which contact was lost until 30 April when wolf 990
was discovered in a new pack (Mendeltna pack} in an area 40
miles (64 km} west of the 1975 Ewan den. This pack, which
contained at least two wolves never before seen with the Ewan
wolves (and was later determined to include at least 5. adults),
raised six pups at a den in the area during 1976. Wolf 990 and
possibly two other Ewan pack members had successfully integrated
into another breeding pack during March or April 1976 and
subsequently remained in that association.
Following the dispersal of the radio-collared female the
remainder of the Ewan pack could not be located. Extensive
reconnaissance in late winter revealed no indications of more
than a single wolf in the area previously used by the pack, and
frequent aerial checks of the 1975 den showed no sign of
activity. Although the loss of three, and possibly four, pack
members from the original pack (2 mortalities and 1 or 2
dispersals) was suspected, the remaining members of the pack
were unaccounted for and may have disappeared through dispersal
and/or mortality, or were not detected by our survey methods.
After wolf 990 had dispersed ·in April 1976 we did not
reestablish radio contact with the pack until March 1978. This
pack's territory contained dense spruce stands which made
tracking from a Super Cub difficult.
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We were, however, able to maintain records on the numbers
of wolves in the pack from· track counts and public sightings.
On the basis of track counts and public sightings this pack
probably included five wolves in fall 1976. If public
observations were correct/ the five wolves present in fall 1976
either represented immigration or indicated the pack had
reproduced in 1976. We suspect the latter. No wolves were
reported taken from this area during the 1976-77
hunting-trapping season/ although public sightings and track
counts indicated only three wolves· remained by spring 1977.
Trappers reported ·taking two pups in this area during
winter 1977-78/ indicating denning and successful reproduction
in 1977. Radio contact was established with this pack in
February 1978 when a gray female pup (#151) was radio-collared.
The pack numbered four at the beginning of spring 1978 season.
We again lost radio contact with this pack in March 1978 when
wolf 151 died 1 apparently from wounds inflicted by an adult
moose. Radio contact was not reestablished until fall 1978.
Nu..rnbers present in this pack according to public observations
are presented.in the Wolf Density section.
On 17 November 1978 we captured and radio-collared a gray
male pup ( #219) which 1 on the basis of tracks I may have been
accompanied by one or two associates. Following its capture the
wolf was observed alone in the vicinity of Crosswind Lake.
Contact with the animal was lost by late November, and at that
time we suspected the animal had been harvested. However, in
March 1979 wolf 219 was trapped north of Butte Lake, 80 miles
(128 km) from its November 1978 location. From tracks/ the
trapper suspected wolf 219 was associated with two or three
other wolves and 1 therefore, we suspected this wolf had become
associated with a new pack.
The 1'!"an pack terri tory encompassed approximately 864 m2
(2/238 km ). The den site occupied by the Ewan pack in 1975 was
not used from 1976 through the denning season in 1980.
Hogan Hill Pack Radio contact with this· pack was
established in June 197 5 when a yearling female ( #987) was
captured and radio-collared. The pack included seven adults (2
additional adults were observed on 1 occasion). In 1975 the
pack denned 2 miles (3.2 km) west of the Trans-Alaska Pipeline.
We could only confirm that two pups were raised from the 1975
litter. Reasons for the small litter size were unknown, but
canine distemper may have had an influence (Stevenson, et al.
In Prep). In November 1975, a young male (#988) was captured
and radio-collared.
Information on this pack's activity between June 1976 and
April 1978 was limited because wolf 987 was relocated only once
due to a faulty radio transmitter. We do not know if the pack
denned in 1976 but the 1975 den site/ located about 2 mi
(3 .2 km) west of the Trans-Alaska pipeline/ was not used. No
1 7
increase in pack size was evident between spring and fall 1976
according to public reports and Department track counts.
Wolf 988 remained in the eastern half of the pack's
terri tory during most of fall 1976. However, he traveled at
least once to the east fork of the Chistochina River, about
30 mi (48 km) east of the normal territory boundary. In
December 1976 he was again observed in the Hogan Hill territory
accompanied by two gray wolves. Shortly after the first of the
year he apparently dispersed from the Hogan Hill pack, since
radio contact was lost until March 1977 when he was relocated
with one other gray wolf north of Mankomen Lake, about 36 mi
(58 km) from the Hogan Hill terri tory. Radio contact with this
wolf was lost shortly after it was observed in this area.
Track counts and observations made by Department personnel
and the public indicated that at least four or five wolves
remained in the Hogan Hill terri tory after wolf 988 left the
pack. When radio contact with the pack was reestablished in
April 1978 it contained eight wolves. ~wo (#205 and #206) of
three wolves that were subsequently radio-collared were
11-month-old pups; indicating the pack had denned in 1977. They
did not use the 1975 den site again during this reporting
period.
On 9 May 1978, members of the Hogan Hill pack were observed
at a new den site located about 10 mi (17 ~m) NNE of Fish Lake,
Pups were first observed on 30 May with the largest number (5)
observed on 20 June. The pups were moved to the first
rendezvous site, located about 1 mi (1.6 km) north of the den,
on 18 June. The pups had moved to a second rendezvous site,
located 5 mi (8 km) north of the den, by 6 July.
By late fall 1978 the pack numbered 12: 7 adults and 5
pups. Radio contact with wolf 206 was lost in late September
1978. Eleven or 12 members of the pack, including both
radio-collared males, were ground shot in January 1979 and thus
radio contact with this pack was terminated.
The year-round home range of the Hogan Hill pack overlapped
to various degrees, the ranges of the Ewan, Deep Lake, Keg Cree~
and the2 Middle Fork 2wolves. T~rritory size varied from 345 mi
(894 km ) to 567 mi (1,469 km ) from 1975 through early 1978.
Overall, from 1~75 through 1978, the territory encompassed
597 mi 2 (1,546 km ).
Jay Creek Pack -The Tsusena gray female (#199) moved from
the Tsusena pack's territory shortly after being radio-collared
in 1976 (a young adult male was also radio-collared at the same
time but was never relocated). On 28 June 1976, wolf 199 was
located in the Susitna River Study Area where she was observed
bedded down just east of Jay Creek. Wolf 002 and her associate,
a yearling gray male, both from a different pack, were observed
within 1/2 mi (0.8 km) of ~nd heading towards the Tsusena female.
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The Tsusena female appeared to become excited, running in short
tight circles, then attempted to hide in nearby brush. She hid
for a few minutes, then burst out at a full run with the Coal
Creek wolves chasing some distance behind. This chase lasted
for 20 minutes and covered a distance of approximately 4 mi
(6 km). During most of the chase the wolves did not appear to
be in visual contact and the pursuing wolves stopped often to
pick up the scent. At one point the pursuing wolves were within
l/4 mi (0.4 km) of the Tsusena wolf but for the most part they
were easily outdistanced. Following the chase the Coal Creek
wolves rested at a creek while the Tsusena female continued at a
slower pace toward the Susitna River.
After this chase_by Coal Creek wolves, wolf 199's movements
became more widespread in a southerly direction. She was
observed 14 mi (22 km) up Kosina Creek late in June 1976 and
then moved back to the mouth of Watana Creek. She remained
along the Susitna River until the end of July, then moved 40 mi
(64 km) southeast to the head of Joe Creek. She stayed in that
area and the vicinity of upper Daisy Creek, which comprised the
summer territory of the Mendeltna pack, until 21 August 1976
when she was observed 5 mi {8 km) southeast of Clarence Lake.
During the remainder of summer, fall and winter of 1976-77
this wolf's movements were restricted primarily to the Susitna
River lowlands. She was observed alone until 29 September 1976
when she was seen with a large, dark gray wolf which, on the
basis of size, was judged to be an adult male. She was observed
with this male on seven different occasions and it became
apparent that a social bond had been formed.
We lost contact with wolf 199 after 19 March 1977 due to
radio failure and, ~~erefore, have no location sightings until
she was recollared in March 1978.
During winter 1977-78, tracks of three or four wolves were
common along the Susitna River in the same locale where the
Tsusena female (199) had been observed in 1976. That winter 13
moose kills were observed on the river. We attributed these
kills to this pack on the basis of tracks. In March 1978, when
wolf 199 was recollared just east of Clarence Lake, she was
pregnant and accompanied by three wolves, an adult dark gray
male and two female pups. All but wolf 199 were killed by
Department personnel as part of the experimental wolf removal
study.
Wolf 199 was observed only three times during May and June
1978 because of dense vegetation which prohibited visual
observation. She was tracked, however, to an aspen vegetated
knoll at Jay Creek on several occasions indicating that she was
denning in that area. On 28 June, she was observed there
accompanied by at least three pups.
1 9
Radio contact with wolf 199 ended in July 1978, probably
due to radio failure. During winter 1978-79, tracks of four to
five wolves, in the same area occupied by the pack earlier,
indicated that wolf 199 successfully raised pups in 1978.
During winter 1979-80 aerial· trappers reported 10 to 11
wolves within the Jay Creek pack area. Six wolves were ground
shot at that time. At least four wolves were observed on the
Susi tna River near Jay Creek during a moose survey in March
1980, indicating that the pack area was still occupied, probably
by descendants of wolf 199 which we suspect survived through
1980.
Keg Creek Pack -Three adult wolves, two adult males (#986
and #083) and one adult female (#002) were captured and
radio-collared in April 1975. Two additional wolves, an adult
female (#068) and a female pup (#201) were radio-collared in
March 1976.
During winter 1975-76, mortality within this pack wa13 high;
four wolves were taken with the aid of aircraft in December 1975
and we suspect that three to five wolves were illegally shot
from aircraft in mid-March 1976.
Wolf 068 was suspected of being the dam of the 1975 litter
and was the dam of the 1976 litter. Dense vegetation prevented
observation of pups until 21 Auyust, but radio locations of
female 068 indicated that by 9 July the pups had been moved to a
rendezvous site about 3 mi (4.8 km) north of the den site. By
29 August the PU!)S began regularly traveling with the adults.
Between late August 1976, when pups were first observed,
and March 1977 t.he size and composition of the pack remained
stable at three adults and five pups. No mortality was observed
during this period.
Studies of moose movements indicated that some moose from
the Susitna River Study Area wintered and calved in the
territory of the Keg Creek pack (Ballard and Taylor 1978). The
decision was made to expand the study area to include the
year-round range of these moose. Therefore, in late March 1977,
Department personnel removed seven of eight Keg Creek pack
members. The wolf which escaped was the gray adult female
(#201) collared in 1976, with which we lost contact because of a
faulty transmitter. One of the grays which we had observed with
the pack for the previous several months was the radio-collared
adult gray male ( 06 7) which had dispersed from the Deep Lake
pack in August 1976.
Wolf 201 apparently did not breed during 1977, because we
never found any indication of more than one set of wolf tracks
until late March 1978 when she was recollared with an adult gray
male (#203)~ When collared, wolf 203 was blind in one eye, had
one foot missing and was judged to be 8-9 years old. From what
area this wolf originated is not known.
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Wolf 201 was pregnant when recaptured in 1978. She was
first observed at the den site previously used by the pack in
1975 and 1976 on 9 May 1978. Pups were first observed at the
site on 29 May 1978. The largest number of pups observed was
five (on 12 June). By 25 August the pack moved to a rendevous
site approximately 34 miles (54 km) to the west. On
30 September 1978 a total of 8 wolves were observed: 7 grays
and 1 black, 6 of which were pups (5 grays and 1 black).
This pack remained stable in number until February 1979
when at least three to four members of this pack were ground
shot. Examination of the carcasses indicated that at least two
wolves had been illegally shot from aircraft. Contact with the
remaining two pack members was lost in late February 1979 due to
unknown causes. No public or Department wolf observations were
recorded for this pack area during 1979-80. Therefore, this
pack area was considered to be vacant or occupied by only one
pair of wolves. The den site used in 1975, 1976 and 1978 was
not used in 1979 or 1980.
During the reporting
territory ranging from 106
annually.2 OVerall, the
(1,310 km ).
pe2iod the ~g Creek pa~ occupied 2 a
mi (275 km ) to 432 mi (1,119 km 2
year-round territory was 506 mi
Maclaren Pack -The Maclaren pack included one collared
adult female (#985) which was captured in April 1975. Other
pack members included an adult male and four pup~ reared ~n
1975. This pack inhabited an area of 279 mi (723 km )
encompassing the upper portion of the Maclaren River. On one
occasion the Maclaren wolves were found within the Keg Creek
pack territory where they killed a yearling bull moose.
Although radio locations suggested the presence of a den in
1975, none was observed from the air nor during ground searches
after the area was vacated. However, a rendezvous site 1 mile
( 1. 6 km) from the probable den area was located. Despite the
presence of only two adult wolves, the pack was able to
regularly kill adult moose during winter. One pup was lost
(possibly trapped) in mid-November prior to the removal of the
pack in January as part of the experimental removal program.
Mendeltna Pack -Contact with this pack was established in
April 1976 when the adult gray female (#990) from the Ewan Pack
became integrated into a new pack. The new pack was comprised
of wolf 990, a light gray thought to be a female, 1 dark gray, 1
slate black, and 1 black wolf.
The pack denned in 1976 at a site located 1.5 mi (2.4 km)
SE of Marie Lake. At least six pups, three grays and three
blacks, were raised. Pups were moved to a rendezvous site,
located approximately 1 mi ( 1. 6 km) south of the den, between
28 June and 2 July. The pups remained here until 2 August when
they moved approximately 14 mi (22 km) northwest to the head of
Daisy Creek. Between ~ and 8 September the pups began
2 1
accompanying the adults on a regular basis. On 8 September we
observed wolf 990 with three gray and two black pups. The third
black pup was observed alone at the Daisy Creek rendezvous site.
Following this observation, our counts of the entire pack were
short one black pup. We suspect the pup observed at Daisy Creek
either dispersed, which seems unlikely, or died.
In October 1976, radio contact with wolf 990, originally
from the Ewan pack, was lost. We never again observed this wolf
with the pack nor was she reported in the harvest records. We
speculate that she dispersed or died of natural causes.
Wolf 083, an adult gray male originally tagged in April
1975 as a member of the Keg Creek pack, was discovered with the
Mendeltna wolves in April 1977. We thought that wolf 083 had
been killed by illegal aerial hunting in the Keg Creek
territory. Wolf 083 may have had contact with wolf 990 from the
Ewan pack in October 1975 when both radio signals were located
close to a moose kill. Radio contact with wolf 990 was lost in
February 1976 and consequently reestablished in the Mendeltna
pack in April 1976, while radio contact with Keg Creek wolf 083
was lost in mid-March 1976. This sequence of events suggests
that both wolves may have emigrated to the Mendeltna area about
the same time. Our contact with the Mendeltna pack in late June
1976 indicated wolf 083 was probably present at that time.
The Mendeltna pack contained seven wolves at the beginning
of the 1977 deP-ning season. By 21 April +~ey began visiting +~e
old den site utilized in 1976. After this date, observations at
the den became more frequent. During the 1977 season this pack
maintained two den sites and raised two litters of pups. Adult
wolves were first observed at a second den site, located at
Nickolson Lake approximately 5 mi (8 km) northwest of the main
den, on 20 May. Wolf 007 was the dam of this second litter.
Pups were first observed at the main den on 1 June 1977 but
not at the second den until 12 June 1977. At least seven pups
were raised at the main den while at least two were reared at
the second den. It was possible, however, that 10 or more pups
were born because we made one sighting of three pups at the
Nickolson den and perhaps eight at the main den. However, these
observations occurred during a period when pups may have been
transferred between den and rendezvous sites.
Nickolson Lake pups were first moved-to a rendezvous site
approximately 1 mi (1.6 km) east of the den on 22 June. At the
main den we saw two adult wolves begin to move five pups to the
Kelly Lake rendezvous site in the morning of .23 June, arriving
at the site over 4 mi (6 km) straight line distance away, at
least 12 hours later. We noted that as the pack moved from the
den, the light gray wolf which we believed to be the dam kept
returning to the den site. Because we were certain that at
least seven pups were being raised at this den we speculate that
two pups died. Our observations in the fall, when snow
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conditions were optimum for observing wolves, could only confirm
the presence of eight pups in the entire pack. Therefore, at
least one or maybe two or more pups were lost during the summer.
The circumstances surrounding these losses are unknown.
By 25 July the Nickolson Lake rendezvous site was abandoned
and the pups were moved to the Kelly Lake site. Between 8 and
17 August all the pups were moved to a. third rendezvous site at
White Sand Creek located approximately 9 mi (14 km) northwest of
the Kelly Lake site. By 13 September, the pups began traveling
with adults on a regular basis.
By early fall 1977, the pack numbered seven adults and
eight pups. In early November wolf 008 was discovered dead at
the northern edge of the pack's terri tory west of Moose Lake on
Tyone Creek. Examination of the kill site indicated that he had
been killed by several other wolves. Tracks of four or five
wolves were followed down Tyone Creek, indicating they were
members of the Tyone Creek pack. The dead wolf had been fed
upon and a portion of the rib cage and most of the viscera were
gone. Most of this appeared to be the result of scavenging
activity by ravens (Corvus corax) and gray jays (Perisoreus
canadensis). Necropsy revealed several punctures on the neck
and considerable subcutaneous hemorrhaging in both the neck area
and on the upper left portion of the cranium. We surmise that
this wolf was killed by another pack, apparently while
trespassing in the Tyone Creek territory,
During winter 1977-78, the Mendeltna pack slowly suffered
attrition from ground shooting and illegal aerial hunting until
early February when the pack numbered eight. Heavy snowfall
during February followed by clear sunny weather provided
excellent wolf tracking conditions. On 9 February 1978 the
remaining eight known members of the Mendeltna pack were
harvested by ground shooting. Members of this pack were
accounted for by wolf hide sealing documents with the possible
exception of three, a radio-collared gray male pup and two black
yearlings. The carcass of the radio-collared pup was found
buried in snow, where it had been shot with buckshot and not
retrieved. The two blacks, one a crippled yearling male, still
remain unaccounted for; either their presence has been
undetected after several hours of census, or they actually were
harvested and their harvest location falsely recorded.
From June 197~ to February 1978, this pack occupied a
territol2Y of 559 mi 2 (1,448 km) (Fig. 2). Summer territory wa~
515 m~ (1,334 km ) while winter territory was 266 mi
( 689 km ) • Members of this pack traveled to the extreme edges
of the eastern, western and southern portions of their territory
during summer while the northern extreme was frequented
primarily during winter.
No wolves were known to occupy this area until late in 1978
when at least one member. of the Tolsona pack began traveling
2 3
through the area. Repopulation of this area is described in the
Tolsona pack section.
Middle Fork Pack Radio contact with this pack was
established in March 1976 when wolves 062, 078 and 996 were
captured and radio-collared. Unfortunately the collars were
chewed off within 2 weeks and only seven radio locations were
obtained. At that time the pack contained nine animals.
our only contact with the Middle Fork pack during fall 1976
was a track sighting by a member of the public which indicated
the pack still numbered nine. During winter 1976-77 two wolves
were reportedly shot, one was wolf 078 which had been collared
in March 1976.
We reestablished radio contact with the pack in March 1977
when it numbered four. Two wolves were captured. Wolf 062 was
again recaptured along with a 10-month-old pup (#061) both of
which were radio-collared. Wolf 062 drowned while still
partially immobilized. &1 additional pup (#085) was
radio-collared in April 1977. We believed the remaining pack
·member was also a-pup, were unable to account for three other
wolves which should have been present in the pack.
During summer 1977, the Middle Fork wolves did not den and
were rarely observed together. They were never observed on any
kills and we believe that they were preying on small game.
In early September, these wolves began associating with
each o~~er once again and did so through winter 1977-78. During
January 1978 both radio-collared wolves, and probably the third
unmarked member of the pack were shot, leaving the Middle Fork
pack territory vacant. An aerial census in late April supported
this conclusion.
2 The Middle Fork pack occupied an area of approximately 514
mi (1,331 km 2 }. Their territory appeared to have considerable
overlap with that of the old Delta pack's area. Like the Delta
area, the somewhat open nature of the terrain and vegetation
would probably make members of this pack highly susceptible to
winter aircraft activities. We suspect that late winter wolf
densities in this area have always been and will remain low.
St. Anne Pack -Data pertaining to this pack during 1976
consisted of track counts, public locations and harvest records.
Although the pack denned in 1976 it was not known how many pups
were produced; however, by fall the pack numbered eight.
In April 1977, three wolves (#086, #087 and #088) were
radio-collared from a pack of eight. The pack denned in 1977 at
the same site used in 1976. Pups were first observed on 11 May.
By fall 1977 the pack numbered 14.
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The rate of loss of radio-collared wolves from this pack
was high. Some of the losses were probably due to radio failure
and hunting-trapping mortality. However, harvest locations on
sealing documents did not account for them. Eight of 10 of the
losses were definitely due to dispersal which will be discussed
more thoroughly in the Dispersal section of this report.
By early spring 1978, the pack contained 11 wolves. The
pack denned at a new den site in 1978, located on Nickel Creek
about 6.5 mi NW of the site used in 1976 and 1977. The number
of pups raised was not determined, but by fall the pack numbered
seven. By spring 1979, their number was reduced to four. They
denned at the site used in 1976 and 1977 and by fall again
numbered eight. Dispersal reduced the pack to four members by
spring 1980. The pack denned in 1980 at the Nickel Creek den
used in 1978.
Wolf 094 appeared to be the most stable member of the pack
since she was always present with the pack after October 1977.
At capture we estimated this wolf to be approximately 9 years
old based on tooth wear. Guides in the area report that for
several years ·prior to our contact with wolf 094 a white wolf
was observed in the St. Anne area. Whether it was the same wolf
is not known but since we have observed only one totally white
wolf during this 5-year study this is probably the case.
Whether she was the dam of the litters raised from 1976 through
1980 is unknown but her affinity to the den site in 1979 and
1980 suggest that she was.
From 1976 through 19802 the stz Anne wolvzs occupie<2 a
territory ranging from 366 ~i (948 ~) to 397 mi (1,028 km ).
overall, an area of 513 mi (1,329 km ) was occupied. Although
our monitoring intensity for the 1978-1980 period was not as
thorough as it had been, it appeared the pack had shifted its
intensive use· area to the south and east toward the Klutina
River. Reasons for this possible shift are unknown but may have
been related to differences in prey abundance.
sinona Creek Pack - A young female (#989) collared in April
1975 provided all radio-tracking data until November 1975 when
two males (#'s 038 and 048) were collared. This pack included
seven adults and four pups reared during 1975. The pack of 11
apparently suffered no mortality during winter 1.975-76 and two
litters totaling nine pups were reared in 1976 at two dens 11
miles (18 km) apart. By early June, pups from the northern den
site were moved to the main den site.
During the 1976 denning season, we never observed the
Sinona pack using a rendezvous site that was any significant
distance from the main den site. On 19 July, however, the pups
and adult pack members began using a slightly elevated ridge
approximately 100 yds south of the main site. This site
resembled a den site and was used for several weeks. We began
to observe the pups travelj,.ng. with the adults on a regular basis
in late August.
2S
By fall 1976 the pack numbered 11 adults and nine pups.
Males 038 and 048 provided most of the radio locations obtained
for this pack. The adult gray female (#989) was believed to be
present in fall 1976 but her radio was not operational.
During the 1976-77 hunting-trapping season, most of the
Sinona pack members were ground shot. We were able to account
for the harvest of at least 16 and perhaps as many as 18 wolves
from this pack. We suspected, however, that more than just two
members escaped being harvested. At least two wolves, an adult
with a distinctive white head and the adult gray female (#989),
were not presented for sealing. If the latter wolf lost its
radio collar, it would be the first adult wolf to do so during
this study. Further, when we reestablished radio contact with
the known remaining members of the pack in March 1977 tracks
indicated that five wolves may have been present. However, only
the two radio-collared wolves were present in subsequent
observations. Aerial trappers also believed that more than two
wolves remained on the Gakona River in the Sinona pack
territory.-
Following the heavy harvests from this pack in 1976-77, we
never observed the remaining two known pack members ( #048 and
#084) in the terri tory used the previous year. They occupied
the area Stephenson ( 1978) thought was occupied by the 11 lower
Gakona pack. 11 We refer to this area and the two wolves as
Sinona #2 pack.
During summer 1977, the Sinona #2 pack denned close to the
Gakona River approximately 14 mi (22 km) SW of the 1976 main den
site. Wolf 084 began frequenting the den site by 12 May 1977.
The den site was difficult to observe from the air because of
dense aspen stands, but in 1977 we were able to confirm that the
site was no longer used by the pups after 20 July. In 1977 we
first observed the pups regularly traveling with adults by
15 September. At least six pups were raised and the pack
numbered eight in fall 1977.
We suspect that at least two wolves, perhaps from the 1976
Sinona pack, continued to occupy the old Sinona terri tory, now
referred to as Sinona #1 pack terri tory. A pack's presence. was
partially confirmed during the 1977-78 hunting-trapping season
when at least eight wolves, three adults and five pups, were
shot in the middle and upper Gakona River areas. Both of the
1976 den sites were checked, but no activity was observed.
Thus, if any wolves from the old Sinona pack reproduced they
denned at a new site.
During the 1977-78 hunting-trapping season, four pups
harvested from the Sinona #2 pack. The pack numbered
during spring 1978. Denning occurred approximately 1
(1.6 km) north of the site used in 1977.
were
four
mile
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Six pups were raised in 1978 and the pack numbered 10 by
fall 1978. During winter 1978-79 at least eight of 10 members
of the pack were ground shot by aerial trappers. Wolf 048 may
have escaped being harvested because, after most of the pack had
been shot, he began traveling alone. However, we lost radio
contact with this wolf in January 1979. Radio contact with wolf
213, however, was lost at the same time the remainder of the
pack was harvested and we believe she was killed.
Following loss of radio contact with wolves 048 and 213 in
January 1979 we had neither contact nor public reports of wolves
occupying this area until January 1980. At that time wolf 229
and possibly two associates from the Susi tna pack immigrated
into this area and were joined by one additional gray wolf.
This new pack began occupying the area previously occupied by
the Sinona #2 pack and in spring 1980 denned close to the site
used by that pack in 1977. The den sites utilized by the Sinona
pack in 1976 and the one site used in 1978 were not used again
during this period of study.
Based upon our observations of these packs, we calculate2
two ter~itory areas. Sinona #1 pack occupied an area of 301 mi
(780 krn ) during summer and early winter 1976-77. Pack #2, on
the ot~r hand, ~cupied a year-round territory of approximately
235 mi ( 600 km ) • Had both packs functioned as one, thz
combined 2 territories would total approximately 623 mi
(1,614 km ). .
Stephan Lake Pack The stephan Lake female (#016) was
. radio-collared in February 1976. A total of 14 radio locations
were obtained from 18 June through 18 October 1976, but this
wolf was observed on only two occasions because of dense spruce
cover. Only once was she accompanied by another wolf, a small
dark gray. No kills were ever observed in the vicinity of the
radio locations, although both cow and bull moose were pres~nt
within 1/4 to l/2 mi ( 0. 4 to 0. 8 km) of the wolves on many
occasions. On 14 July, the radio signal was located along
Prairie Creek. Salmon were numerous in this creek and possibly
were being fed upon by this wolf. On 18 October I she was
located at her most southerly location. She was found at the
same location on 22 November I and on 3 December when she was
found dead there under a spruce tree. The cause of death was
not immediately diagnosed because the carcass had been
scavenged. Necropsy revealed internal hemorrhage and what
appeared to be tooth and/or fang marks on the neck in addition
to a 3-inch lacerati9n across the top of the skull. It appeared
that this wolf may ·have been killed by another wolf or by a
brown bear ( Ursus arctos) . This wolf's left rear femur was
greatly deformed and was only 185 nun long, indicating that at
one time she had suffered a fracture which had not healed
properly.
27
Stephenson (1978) speculated that this pack of two denned
in 1976 but our subsequent observations in the suspected denning
area revealed no denning activity.
Following the mortality of wolf 016 we did not attempt to
reestablish contact with this pack because of the logistic
obstacles involved in routinely monitoring the radios. We had
too few radio locations to assess territory size.
Susitna Pack -Radio contact with this pack was established
in February 1979 within the Deep Lake territory. At that time
we observed the Susitna wolves fleeing from the carcass of the
Deep Lake female (#009) which they had just killed. Reasons for
this conflict and the fate of the uncollared gray which had been
accompanying wolf 009 are not known-, particularly since this
pack never returned to the Deep Lake area.
When collared, the pack was comprised of at least two
adults and seven pups. We suspected, on the basis of size and
later capture records, that the tenth wolf was an adult male.
Following capture, the pack moved to the area south of the big
bend in the Susi tna River. Whether these wolves had always
occupied this area is unknown, but seems likely based on the
gaps between territories illustrated in Figs. 2 and 3 for the
periods 1975 through 1978. By late spring 1979, the pack
numbered six or seven. Pack losses between fall and spring were
probably the result of one to two wolves being ground shot and
at least one dispersal.
The pack was first observed at the 1979 den site by
13 April. At least six pups, which were not observed until
3 August, were raised. Between late summer and October 1979,
the pack declined to 10, possibly due to dispersal.
During December and January 1979-80, large concentrations
of Nelchina caribou were found within the Susitna pack territory
as they began migrating toward the Wrangell Mountains. In late
January 1980, wolf 229, a yearling male, and at least two gray
associates appeared to follow the caribou migration and
dispersed to the east. Further details are presented in the
Dispersal section of this report. By early February the pack
numbered seven.
In early March 1980, the pack was reduced by two members
because of a conflict with the Tyone pack. Details of this
conflict follow:
On 8 March, we tracked wolf 295 (the adult gray female of
the Susitna pack) to a location 2 miles (3 .2 km) south of
Vermillion Lake. She was observed alone. We backtracked her in
the snow for several miles to the west to the confluence of
Sanona and Tyone Creeks where we observed three wolves walking
through a medium dense spruce stand. We assumed these· wolves
were members of the Susitna pack but closer examination revealed
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a total of seven wolves, which was more than we had observed in
the Susitna pack during the previous 2 weeks. A check of other
wolf radio frequencies revealed that seven radio-collared
members of the Tyone pack, which was comprised of two adults and
six pups, were also present. We then searched for other
radio-collared members of the Susitna pack and found wolf 296,
the light adult gray male, which on the basis of tail posture
and leadership in the pack was assumed to be the alpha male,
0.5 miles (0.8 km) north of Tyone Creek. Wolf 296 was dead and
an examination revealed puncture marks on the neck and
shoulders. In addition, at least seven distinct wolf trails
radiated from the area, leaving little doubt that wolf 296 had
been killed by other wolves . From blood in the snow, we
backtracked wolf 296 to the location where the struggle had
begun. At this site, we discovered a fresh adult moose kill.
There were at least two wolf beds in the snow approximately
20 feet from the moose kill. We also found a moose fetus, a
dead ptarmigan (Lagopus sp.), and two wolf beds on the opposite
side of the creek from the moose kill. Tracks of a single wolf
(possibly wolf 296) indicated that it had fled from the moose
kill. The carcass of the moose and the carcass of wolf 296 were
about 100 yards apart.
One of the wolf trails radiating from the moose kill site
was spotted with blood. We followed this trail for
approximately 0.25 miles (0.4 km) upstream where the trails of
four wolves came together, suggesting that an apparent pursuit
continued. Approximately 0.25 miles (0.4 km) north of the
creek, members of the Tyone pack had apparently caught wolf 303
(a gray yearling male). Wolf 303 was still alive, but had lost
a considerable amount of blood.
We radio-located wolf 302 of the susitna pack at 1130 hr.
and found her 3.5 miles (5.6 km) east of the carcass of wolf
296. When originally located at 1030 hr. she had been within
0.25 miles ( 0.4 km} of the Tyone wolves. At this time, the
location of four of seven Susi tna wolves and eight of eight
Tyone wolves was known.
While leaving the site we observed an additional fresh calf
moose kill close to the adult moose kill. The calf. had been
killed by punctures in the neck and anal regions but had not
been fed upon.
On 9 March 1980, we located wolves 295 and 302 of the
Susitna pack. Wolf 295 had moved to the east side of the Tyone
River. woYf 302 was within 5 miles (8 km) of wolf 295 and
appeared to be heading directly toward her. The Tyone pack,
however, was in the same location observed on 8 March and had
revisited the kill site of wolf 296 and the site of injured wolf
303. Wolf 303 had moved approximately 50 feet where members of
the Tyone pack finally killed him; he had punctures in the neck
and around the ears.
2 9
Based upon our ground and aerial observations we concluded
that the susi tna pack had come upon a moose kill made by the
Tyone wolves. The moose kill was located close to the
territorial boundary area between the two packs {Fig. 4).
Therefore, in addition to competition for possession of the
kills, the conflict may have involved a territorial dispute.
The Susitna area appeared to have a relatively poor availability
of prey in comparison to the Tyone pack's area during this
winter. Comparison of prey abundance between the two areas will
be discussed in the Food Habits section.
Following the dispersal of wolf 229 and its associates, and
the deaths of at least wolves 296 and 303, the Susitna pack was
reduced to four wolves by late spring 1980. They denned at the
same site they had used in 1979 and were first observed there on
23 April.
During the 1980 denning season, three of four {#' s 295,
302, 305, and 306) remaining pack members were fitted with
activity radio transmitters. In ~ddition, ground obse.rtlations
at the den site were made from 1 May through the middle of June
1980. Methods and results from this study period are being
prepared for publication by James Foster, Woodland Park Zoo and
Warren Ballard, Alaska Department of Fish and Game. At least
six pups were raised during the 1980 denning season.
From earlv 1979 throuqh June 1960, the Susitna pack
occupied a territory of 462 mi~ (1,197 km~). This latter figure
does not include four locations recorded when the pack was
apparently trespassing into the Deep Lake territory in February
1979.
Tolsona Pack -Prior to mid-June 1978 our contact with this
pack consisted of public sightings, track counts, and harvest
records. These data indicated that in early fall 1977 the pack
had numbered at least 11. By the end of winter, the pack had
been reduced to three by trappers.
We established radio contact with this pack in early June
1978 by searching aspen-covered knolls from fixed-wing aircraft
as it searched for a potential den site. The den site was found
and a yearling gray male (#210) was radio-collared. At that
time the pack was comprised of wolf 210, an adult gray female,
and a small black wolf which may have been a yearling. At least
eight pups were raised at the site. Pups were moved to a
rendezvous site 3.5 mi (5.6 km) away from the den site between
24 and 26 June 1978. During late summer 1978, wolf 210 began
exhibiting a propensity to travel to the western extremes of the
old Mendel tna terri tory which was thought to have been vacant
since February 1978. During these forays, wolf 210 was always
observed alone. In mid-September 1978 radio contact was lost.
At that time the pack numbered 10.
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c:BlJJ [TI'--L._j LJ p:--· J c.....J ~ L..::J ['T~<O>] L..:.:J
\~olf pack t\!r'rito'i:-ies derived from plotting radio locations obtained
June 1980 in Game Management Unit 13 of Southcentral Alaska.
St. Anne' s--------::::::::::::
'/)..1'' .,_ v. ',,._;:L;; ~z¥-0'/..~:""'-~-;.:-. .~r,......c'~,~-~:.~.:., T olsona-----------J;.r.·!f.>.~---''"'''' ,,~;:~~;~·,t':tt'·f~ .......... ·. •, ' . .. . . .. Tyone-------------;_ ... • • : •••. ;
Susitna----------
[_ill L;;.J pr ··rl ~
from1 Juiy 1978 through
1 -
. We had no radio contact with this pack from September 1978
until late January 1979, at which time we purchased a black
yearling pup from a local trapper. The pup had been caught by
the toes and was in go.od enough condition to be radio-collared.
At the· time we were uncertain as to her pack affiliation.
Within 2 weeks of capture, however, she had joined the Tolsona
pack which then numbered seven (3 blacks and 4 grays).
By 10 May 1979, the pack began frequenting the den site
utilized in 1978. Pups were first observed outside the den on
25 June. We never obtained an accurate count of the number of
pups produced. In mid-October, however, the pack numbered 16
( 11 grays and 5 blacks). On the basis of size and the scruffy
appearance of pups at that time of the year, we believed at
least six and perhaps nine pups were rais-ed ( 3 blacks and 6
grays).
During summer 1979, members of this pack appeared to
continuously expand their range to the west into the old
Mendeltna territory. In mid-October, when we obtained ~~e
largest count (16) of the pack, they were located close to Moore
Lake which had been the northern terri tory boundary of the
Mendeltna pack (Fig. 3). Wolf 210, which had not been
radio-located since late August of 1978, was with the pack at
that time.
Durinq winter 1979-80, the oack suffered attrition from
trapping a-nd perhaps dispersal even though portions of their
territory were included in an area closed to hunting and
trapping. Radio contact was temporarily lost when ~olf .220 1 s
radio transmitter failed prematurely. We did not reestablish
contact with this pack until early June 1980 when they were
discovered at the Nickolson Lake den site which had been used by
the Mendeltna pack in 1977. Wolf 220 was recaptured in July
1980, and at that time the pack was comprised of at least two
black and seven gray adults. At least six pups were present at
this den site.
Based upon the presence of an adult black wolf in the
Tolsona pack from summer 1978 to 1980 and the known expansion of
this pack 1 s terri tory into the Mendeltna area, we suspect the
black wolf may have been a survivor of the Mendeltna pack. As
mentioned in the Mendeltna pack section, following winter
1977-78 all but two black wolves were accounted for according to
wolf sealing documents. Perhaps, following drastic reduction in
numbers, these two blacks dispersed and became integrated with
the Tolsona pack, which in· spring 1978 was thought to contain
only three wolves.
From June 1978 2through Jun~ 1980, the Tolsona pack occupied
an area of 821 mi ( 2, 126 km ) • Their range extended from
Tazlina Lake to Lake Louise, west to Tyone Creek and then east
several miles past Tolsona Creek (Fig. 4). During this time, no
other pack territories were believed to overlap the Tolsona
territory.
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Tsusena Creek Pack Radio contact with this pack was
established in mid-February 1976 in the area north of the
Susitna River between Tsusena and Portage Creeks. Two (#'s 199
and 994) of nine members were captured and radio-collared.
Shortly after being radio-collared, however, wolf 199 dispersed
to the southeast and after being chased by the two Coal Creek
wolves established a territory at Jay Creek along the Susitna
River. Wolf 994 remained with the Tsusena pack but relatively
few radio locations were obtained probably because of radio
failure. A den site was not located in 1976.
In March 1977 we located a pack of seven wolves at the head
of Clark Creek. From the location we assumed it was the
original Tsusena pack. We radio-collared an adult gray male
(#056) and a gray male pup (#057). The presence of this pup
indicated that the pack had denned in 1976. On the basis of
size and behavior it appeared that four or five of the pack
members may have been the previous year's pups. Unfortunately,
we never established contact with the radio-collared animals.
Unsubstantiated reports from the public suggested that the
radio-collared_ animals had been taken by illegal aerial hunting.
The nature of the terrain within this pack's territory and the
extensive snow cover which was present at that time would have
made this pack highly vulnerable to aerial shooting. No attempt
was made to reestablish contact with wolves in this area until
spring 1980, however, no wolf sign was observed during intensive
aerial searches in mid-April suggesting that this territory may
be vacant.
Tyone Male -Wolf 001 was radio-collared in February 1976
at the west end of the Alphabet Hills. During the subsequent 1
months, ~is wolf traveled alone over a wide area of 1,400 mi
( 3, 626 Jc_"D. ) , traversing parts of at least four of t.."IJ.e
radio-marked pack territories described in L~is study. During
this period wolf 001 was not observed at any kills but was seen
near small bands of caribou on three occasions. After ranging
widely over the lower portions of the Black, Oshetna and Tyone
River drainages and the west fork of the Gulkana River, the
Tyone male was found at an active den near Clearwater Creek in
the Susitna River study area on 20 June 1976. Only one other
adult wolf, a black female, and three pups were observed at this
den. Although it is possible that the female and male were not
previously associated, they may have been since the male was
radio-collared only 14 miles (23 km) from the eventual location
of the active den. The wolves associated with this den were
probably members of a pack which used a territory to the west of
and adjacent to the Maclaren and Keg Creek packs in 1975. The
den site was abandoned in early July and we suspected the pups
had been moved to a rendezvous site, possibly in the vicinity of
Valdez Creek.
During June a~d July the wolves occupied an area of
approximately 290 mi (751 km 2 ). They were observed at two kill
sites: a calf caribou at_Valdez Creek on 27 July 1976, and an
3 3
adult moose on 28 July 1976 which was also being eaten by a
brown bear with two yearling cubs. The latter kill may have
been made by the bears.
Department personnel attempted to remove this pack from the
experimental area in July 1976. However, only wolf 001 was
taken at that time. The remainder of the pack was believed to
have been removed during winter 1976-77 leaving this pack area
vacant.
Tyone Creek Pack -Prior to establishment of radio contact
with this pack in November 1977, our data consisted of track
counts and public sightings. Between spring 1976 and fall 1977,
the pack numbered from six to eight individuals. In fall 1977
the pack numbered 12. Apparently the pack denned in 1977
because one pup (#151) was radio-collared.
During the 1977-78 hunting-trapping season, 11 of 12 known
pack members were harvested in this area, one of which was not
retrieved. The remaining wolf (#116) dispersed from the area.
By 27 February he. was observed accompanied by a black adu1 t
female in the western edge of the Keg Creek terri tory. During
March both wolves emigrated to the Susitna River Study Area. In
late March the black female was removed by Department personnel.
Wolf 116 continued to reside in the Susitna study area and by
20 June was observed with a yearling female which was also
removed in mid-July. By late fall wolf 116 was observed alone
at Monahan Flats, having dispersed over 60 miles (96 km) from
his original capture location. Following this latter
observation we lost radio contact with wolf 116 ·and assume he
dispersed farther to the north or west.
During fall 1977 an?2 early· w~nter 1978, the Tyone pack
occupied an area of 253 mi (655 km ). Public observations and
track sightings indicated that the pack also ranged to upper
Goose Creek on the west and the Susi tna River to the north.
In November 1978 we reestablished contact with wolves in
this area when two adults (#215 and 216) were radio-collared.
Whether these wolves were descendants of the original Tyone
Creek pack, which we thought was eliminated by ground shooting
in 1978, or represented wolves colonizing a new area is not·
known. They did, however, occupy the area previously occupied
by the Tyone pack.
During winter 1978-79, no other wolves were observed with
this pair. They were first observed at the 1979 den site on
23 April. Seven pups, which were first observed on 6 July, were
reared.
In early March 1980, this pack killed two wolves from the
Susitna pack during a conflict near two recently killed moose.
Details of this conflict were provided in the description of the
Susitna pack.
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Wolf hunting and trapping within this pack's territory were
closed in both 1978-79 and 1979-80 so wolf numbers would remain
stable for predation rate studies. In March 1980, however, we
apprehended two individuals who were harvesting and attempting
to herd wolves from fixed-wing aircraft in the closed area.
Both admitted guilt and the Piper Super Cub was forfeited to the
State and one individual was given a suspended jail sentence and
placed on 2 years probation. Five wolves, three of which were
radio-collared (#215, 216 and 301)1 were removed from the pack.
One of these was legally ground shot when the pack ventured out
of the closed area in January 1980. These mortalities reduced
the pack to four pups by late March 1980.
Following the removal of the alpha male and female from the
pack, the remaining pups apparently were unable to kill either
moose or caribou. From mid-March through July these wolves were
never observed on a fresh kill and revisited many of the old
kills made prior to the reduction in pack numbers. In mid-April
t.~e pack bega11. exhibiting an erratic movement pattern: on
14 April they were observed at Kosina Creek approximately 20
miles (32 km) northwest of the original territory boundary, then
on 25 April they were observed on the middle fork of the Susitna
River approximately SO miles (80 km) north of the territory
boundary. In both cases, however~ the pack returned to the old
pack territory within a few days of the observation. Obviously
the pack did not den in 1980 but the pups did show a tendency to
linger around the old 1979 den site.
Before the adult members of the pack were ground shot in
March zl-980 the2 pack occupied a relatively small territory of
302 mi (782 km ).
Watana Pack -Contact was temporarily established with this
pack in March 1978 when three wolves were removed by Department
personnel as part of the experimental wolf removal program and
one adult male (#197) was radio-collared. Wolf 197 occupied the
area from upper Watana Creek to lower Fog Creek. We lost
contact with this wolf in April due to unknown causes.
From April 1978 to April 1980 our data for this pack
consisted only of track counts and Department observations. By
fall 1978, the pack numbered three and may have remained at that
size through spring 1979, but we were only certain of the
presence of two wolves. The pack apparently denned in 1979
because seven wolves were present by fall.
In late April and early May 1980, three adults {#308/ 310
and 311) and one pup (#309) were captured and radio-collared
near Watana Creek. They were first observed at a den site on
13 May where at least six pups were raised. Based upon a
limited number of radio locations obtained from April through
June 1980 the pack occupied on area of approximately 258 mi 2
( 668 km 2 ).
3 5
Wolf Territories
For the purposes of this report we used Etkin ( 1964)
definition of territorality; "any behavior on the part of an
animal which tends to confine . . . its movements to a
particular locality. 11 Most definitions of terri torali ty assume
that the terri tory is defended against intruders. Although
wolves in the Nelchina Basin apparently do at times defend their
area against other wolves, intrusions into a neighboring
territory often occur when the home pack is not using that
portion of the area.
Table 2 summarizes territory sizes for 19 wolf packs
studied from April 1975 through June 1980. Territory sizes were
calculated only for those packs which had been intensively
studied during that time period. We calculated the size of
territories for four separate time periods: 1) wolf packs
studied from April 1975 through June 1976 (includes some packs
killed during experimental wolf removal in 1976); 2) packs
studied from June 1976 through June 1978; 3) packs studied from
July 1978 through June 1980; and 4) a summary of all packs for
all study years. -The first two time periods correspond with
analyses reported in previous progress reports by Stephenson
(1978) covering the 1975-76 period and by Ballard and Spraker
(1979) for the 1976-78 period. We separated territory sizes in
this manner because of differences in sampling intensity and
differences or apparent shifts in wolf territories.
As this study progressed we began analyzing the number of
pack locations ·in relation to computed territory size. We had
anticipated that, as more locations were obtained, the increase
in terri tory size would level off, . and in turn provide us with
an indication of the nu.-rnber of locations necessary to adequately
measure territory size in the Nelchina study. This was not the
case, however. As the number of locations increased the rate of
increase in terri tory size decreased, but overall computed
territory size continued to increase. It became apparent that
although wolf packs continued to occupy the same general area
their use of outlying areas changed ·by season and year.
Therefore, the apparent. boundaries of individual territories
appeared to be in a constant state of flux even though core
areas remained fairly constant. Haber (1977) working in nearby
Mt. McKinley National Park, felt that territory boundaries
remained stable over long periods of time. This may well be the
case in a saturated, unhunted wolf population but does not
appear to be the case in GMU 13 where wolves are exploited and
for various other reasons exist at less than maximum densities.
Some of the apparent differences in terri tory size were
undoubtedly ·attributable to seasonal differences in moni taring
intensity. This was most evident for the following packs:
Hogan Hill from 1976-80, Keg Creek from 1978-80, and Sinona #2
from 1978-80. During these time periods these packs continued
to use the core areas.
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Table 2. Summary of territory sizes for wolf packs intensively studied in
Game Management Unit 13 of southcentral Alaska from April 1975 through
June 1980.
Maximum
annual
density
in terr.
1975-1976 1976-1978 1978-1980 Total ,eer wolf
Pack name .2 km2 .2 km2 .2 km2 .2 km2 .2 km2 m~ m~ m~ m~ m~
Brushkana 468 1212 468 1212 47 121
Butte Lake 1188 3077 1188fJ 3077 297 769
Deadman 108 280 108-280 22 56
Deep Lake 701 1816 548 1419. 392 1015 8281/ 2145 104 268
Delta 288 746 288-746 36 93
Ewan 864 2238 864 2238 79 203
Hogan Hill 567 1469 345 894 181 469 5971/ 1546 so 129
Jay Creek 288 746 288-746 29 75
Keg Creek 432 1119 414 1072 106 275 506 1311 39 101
Maclaren 279 723 279 723 47 120
Mendeltna 559 1448 559 1448 37 97
Middle Fork 514 1331 514 1331 47 121
Saint Anne 397 1028 366 948 513 1329 37 95
Sinona 1/1 432 1119 301 780 472 1222 24 61
Sinona 112 235 609 146 381 268 694 27 69
Susitna 462 1197 4621/ 1197 36 92
Susitna-Sinona 284 736 284-736 71 '184
Tolsona 821 2126 821 2126 51 133
Tyone 253 655 302 782 364 943 30 79
Mean 537 1390
Standard deviation 189 488
y Not included in mean due to inadequate data or nondenning pack.
3 7
Because of changes in use patterns by individual packs and
differences in sampling we believe the most representative
terri tory sizes are presented in the total column of Table 2.
These figures include the core use area and encompass changes in
peripheral . areas. Average t¥'ri tory si~e for 14 denning wolf
packs from 1975-80 ~s 537 m~ (1,390 km ). 2 Territoriez; ranged
in size from 268 mi ( 694 km ) up to 864 mi ( 2, 238 km ) . on2
nondennin~ pack of three males occupied an area of 1,188 mi
(3,077 km ) which overlapped the territories of four other
packs. zimilarly, o~e lone male (#001) ranged over an area of
1,400 mi (3,626 km ) which also overlapped four pack
territories. The movements of this latter wolf began resembling
those of a regular pack member when he became associated with a
female at a den site.
The Deep Lake, Ewan and Tolsona wolf packs had the largesz
territorizs of the denning packs, averaging over 800 mi
(2,072 km ). Of possible significance is that these three packs
accounted for 51 percent of the caribou kills observE'i during
this study. In view of the low density of moose on the Lake
Louise flats, and the relatively greater dependence of these
packs on caribou· during winter, we suspect the larger
territories were a result of the low year-round density of
ungulates in these areas.
Wolf pack territories derived from plotting radio-locations
obtained in 1975-76 are illustrated in Fig. 2. This diagram
differs slightly from the preliminary assessment provided by
Stephenson ( 1978). In an effort to provide comparable
illustrations, the outermost point of pack locations was used to
determine territory size consistent with the method described by
Mohr (1947). These territory boundaries represent what appeared
to be Lhe situation at the time wi~~ ~~e possible exception of
the Butte Lake pack, which was a non-denning group comprised of
males.
Wolf territories derived from radio-locations obtained from
July 1976 through June 1978 are illustrated in Fig. 3. During
this time period territories were similar to those presented in
Fig. 2 for the Deep Lake, Ewan, Hogan Hill, Keg Creek, Middle
Fork, and Sinona #1 packs. Differences in territory boundaries
were the result of excessive harvests by either experimental
wolf removal by Department personnel or ground shooting by the
public. Packs that established new territories through
dispersal or shifting use patterns include Coal Creek, Jay
Creek, Tyone male, and Sinona #2.
The greatest overlap in territory boundaries was exhibited
by the Delta and Middle Fork packs. The area occupied by the
Delta wolves from March through May 1977, and subsequent
movements described in the preceding section, indicate that
these wolves probably did not actively maintain a territory but
rather were in the . process of colonizing a new area. In
contrast, the area depicted for the Middle Fork wolves was
occupied during 1977 and 1978.
38
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Territories derived from radio-locations obtained from July
197B through June 19BO are illustrated in Fig. 4. Of particular
interest are the unoccupied areas between the Tyone and Deep
Lake packs and the Ewan and Mendeltna packs depicted in Fig. 3.
Contact with packs in the identified gap areas was established
in 197B and 1979, respectively, suggesting that these wolf packs
were present prior to July 197B.
From June 1978 through fall 1978 the Tolsona pack occupied
the void area depicted in Fig. 3 . In late fall, however, a
Tolsona pack member (a radio-collared yearling male) was
observed in the Mendeltna pack territory. All, or most, of the
Mendel tna pack had been killed by ground shooting in February
and March 1978. During the winter months, the Tolsona pack
shifted their movements to the west and by late 1979 they
occupied their own area and the old Mendeltna's pack territory.
The expansion of the Tolsona pack's terri tory appeared to be
directly related to the elimination of the Mendeltna pack.
Contact with the Susitna pack was established in February
1979 when two_ adults and eight pups appeared to be trespassing
into the Deep Lake terri tory. Because at least two adults were
present, and examination of scatter diagrams of wolf pack
sightings revealed little overlap with the Depp Lake and Tyone
Creek pack areas, this pack was probably present in 1977-78 but
was probably a relatively small pack.
The territory depicted in the sinona area probably
represents establishment of a new pack in a vacant area. During
winter 1979-80 at least one radio-collared wolf and probably
three wolves from the Susitna pack appeared to leave the pack
and follow migrating caribou to the sinona area. They did not
return to their fo£~er territory but began inhabiting the area
formerly occupied by the Sinona #2 pack. At least eight and
perhaps all 10 members of this latter pack were killed by ground
shooting in 1978. It was possible that one wolf remained in the
sinona area since we could only account for three members
missing from the Susitna pack while the new group numbered four.
c Terri tory boundaries depicted in Figs. 2, 3, and 4 were
essentially nonoverlapping during the course of any particular
year. What overlap did occur during a year was either
seasonally temporary in nature-or was the result of the way
territories were illustrated.
Den and Rendezvous Site Usage
The earliest date when the members of a radio-collared wolf
pack were observed at a natal den was 13 April (Table 3).
Wolves usually began visiting den sites in late April and May
when they appeared to be in the process of cleaning L~e site in
preparation for parturition.
3 9
Table 3. Chronology of den and rendezvous site usage of selected study wolf packs in the Nelchina Basin study area from 1975 through 1980.
Date
Wolves first found at natal den Date pups first seen outside den Date moved to lst rendezvous site Pups appeared to travel with adult
Pack 1975 1976 1977 1978 1979 1980 1975 1976 1977 1978 1979 1980 1975 1976 1977 1978 1979 1980 1975 1976 1977 1978 1979 1980
Butte Lake 5/21 6/19 8/1 9/26-
10/6
Deadman Lk 6/26 3/9 7/10 11/12
Delta Riv 6/3 6/6 7/29 9/12?
Ewan Lake 6/13 7/30 9/23?
Hogan Hill 5/14 5/23 6/13 5/30 7/23 6/19 ? 10/13
Keg ~reek 5/23 4/13 5/9 6/13 8/21 6/8 7/2 7/14 8/25 9/23-8/29 <9/12
9/30
Maclaren R 5/21 6/16 10/4
Mendeltna 5/25 4/21 7/2 6/1 6/28-6/22 9/1-8 <9/13
7/2
Sinona Ck 5/22 5/6 5/12 <5/29 4/16 7/22 7/19 9/15 9/12 7/11 7/20? ? ? 9/25-8/25 9/15 9/12
Susitna 4/13 4/23 8/3 6/6 7/19
1/ 10/4
6/6---9/11 8/12
St. Anne 5/11 5/23 5/24 5/12 8/17 9/25 9/12 8/15 <7/13 ? ? ? 8/17 9/25 9/12 8/15
Tolsona <6/4 5/10 6/16 6/25 <6/10 6/24 7/19 6i9!1 9/1
Tyone 4/26 7/6 ? 9/11
Watana 5/13 7/14 7/14
!/ May have been induced by human disturbance.
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Dates at which pups were first observed outside dens or
rendezvous sites ranged from 1 June to 15 September. Late
sightings were the result of infrequent radio contact and dense
vegetation which prevented observation. Mech (1970) indicated
that wolf pups were commonly observed outside of dens when 3
weeks old. Clark (1971) observed 10-day-old pups outside dens
on Baffin Island while Foster (pers. comm.) indicated that pups
of captive wolves begin going outside the den when 16 days old.
Aerial observations of wolf den site usage and pup
emergence suggest that in southcentral Alaska parturition occurs
from mid to late May. However ground observations in 1980 at
the Susitna wolf den revealed that parturition had occurred by
1 May (Foster and· Ballard, unpub. data). Based upon this
observation and our observations of obvious size differences in
pups of different litters, we suggest that parturition l.S
variable and probably occurs throughout the month of May.
Natal dens were, in most cases, vacated during July with
dates ranging from 4 June to 1 August. Distances between natal
dens and first rendezvous sites ranged from a few hundred yards
to 9 miles {14 km). In a few instances, we observed packs
moving to a second rendezvous site in August and early
September. Usually these latter sites were used only for
several days. Pups began traveling with adults on a regular
basis between late August and mid-September.
Most wolf dens were roughly centered within the observed
territorial boundaries, but the Hogan Hill, Keg Creek and
Maclaren dens were located near terri to rial boundaries. The
average· distance between eight natal dens used in 1975 was 22.8
air miles (37 km) and ranged from 16 to 28.5 air miles (25 to
46 km) • These figures represent only those cases in which we
are certain that no dens existed in intervening areas. This
average distance is somewhat less than that described in the
northcentral Brooks Range (Stephenson and Johnson 1973) where
the minimum average distance between dens was about 25 miles
(40 km).
Food Habits in Relation to Availability of Prey
The following section describes the summer and winter food
habits of wolf packs that were intensively studied from April
1975 through June 1980. Food habits of packs for which only
limited data were available were partially described in the
preceding section of this report and have been included in the
food habits summary section.
Brushkana Pack -From April through mid-November 1975 the
Brushkana pack was observed at five kills: four moose and one
of unidentified species. Scats collected from the 1975 den site
suggested that moose were the most important prey item in late
spring and summer (Table 4), and caribou the second most
important. overall, ungulates comprised 87 percent of the prey
items found in scats collected from the den site.
4 1
Table 4. Incidence of food remains in wolf scats collected at the
Brushkana Creek wolf den occupied during late spring and
early summer 1975 in GMU 13 of southcentral Alaska.
Items in 40 Items in 144 Items in 184
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 2 5.0 12 8.2 14 7.5
Calf moose 17 42.5 77 52.7 94 50.5
Adult caribou 12 30.0 22 15.1 34 18.3
Calf caribou 2 5.0 9 6.2 11 5.9
Dall sheep 2 5.0 6 4.1 8 4.3
Beaver 3 7.5 9 6.2 12 6.5
Muskrat 0 0 1 • 7 1 .5
Unid. furbearer 0 0 2 1.4 2 1.1
Wolf 0 0 1 .7 1 .5
Vegetation 0 0 3 2.1 3 1.6
Unidentified I) 5.0 /, 2.7 6 3.2 "" ..,.
Total .40 100.0 146 100.1 186 99.9
Grou12ed Data for 184 Scats
Food item Number of items Percent occurrence
Ungulate 161 86.6
Small mammals 13 7.0
Other 12 6.5
Total 186 100.1
42
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Moose were the most numerous year-round large prey species
within the Brushkana terri tory. Studies of moose movements in
this area indicated that moose were nonmigratory with movements
consisting of short altitudinal movements from summer to winter
range (Ballard and Taylor 1980). Caribou, however, appeared to
be very migratory except for a small subpopulation which
appeared to be year-round residents of Monahan Flats, wintering
on the flats and spending summer through fall at higher
elevations within the pack's territory .
Secondary prey species within this pack's area included
beaver (Castor canadensis) muskrat (Ondatra zibethica), snowshoe
hare (Lepus americanus), ptarmigan (Lagopus sp.) and numerous
microtine rodents. Both beaver and muskrat appeared to be
numerous within the northern half of this pack's territory and
along Brushkana Creek. Ptarmigan appeared numerous along
Brushkana Cree)t and Butte Lake during this period of study.
Dall sheep (Ovis dalli}, however, were not numerous.
Butte Lake Pack -Food habits information oertaininq to
this pack consisted solely of radio location data because the
pack consisted of males only and did not den in 1975. From
April 1975 through January 1976 the pack was observed on 5
kills: 3 adult moose, unidentified 1 moose and 1 caribou.
This pack's territory encompassed the Clearwater Mountains.
Moose appeared to be the most numerous large prey item followed
closely by caribou. Caribou were year-round residents within
the Butte Lake territory while moose in the Clearwater Mountains
were highly migratory (Ballard. and Taylor 1980). Based upon
movement studies, moose would only have been available to this
pack on a year-round basis in the western half of the territory.
Caribou were also available on a year-round basis within the
western half of the territory.
Information pertaining to population levels of secondary
prey species is quite limited. Both beaver and muskrat were
numerous along the Susitna River and its tributaries. Dall
sheep were occasionally observed on the southern slopes of the
Alaska Range near the middle fork of the Susitna River. Sheep
populations in this area have been quite low for a number of
years and only within recent years have they begun to increase.
Ptarmigan could also constitute an important prey i tern during
some years. Snowshoe hares were not numerous according to
aerial observations.
Clearwater Pack -Contact with this pack was not
established until June 1976 when wolf number 001, which
previously had been a loner, was found at the Clearwater de2
site. Pzior to this, wolf 001 ranged over a 1,400 mi
( 3, 626 km ) area but was never observed at a fresh kill. our
aerial observations suggested that wolf 001 fed on an occasional
caribou and often hunted small game.
43
During June and July the two adult members of this pack
were observed on one caribou and one adult moose kill which was
contested by a brown bear. The limited number of scats (n = 33)
collected at the den site indicated that calf moose (86%) were
an important summer food for this pack (Table 5) .
Deadman Pack -Although this pack denned in 1975, no den
site was located. However, two rendezvous sites were located
and scats were collected (Table 6) at one of these. The 30
scats suggested high dependence on moose: 34 percent occurrence
of adult moose and 47 percent occurrence of calf moose. The
small number of kills located while radio-tracking during winter
(3 moose) also indicated that moose were the primary source of
food. Moose in this area were relatively numerous on a
year-round basis and were relatively sedentary. Caribou were
available in limited numbers during much of 1975-76 and may also
have been taken but not detected during the study.
Deep Lake Pack -From 1975 through spring 1980 we located
only one den site ( 1975) even though t~e pack also dew'"'led in
1976 and 1977. Scats from the 1975 site indicated that calf and
adult moose were the most important prey items (53% occurrence)
but that snowshoe hare ( 13 .1%) and beaver ( 15. 6%) were also
important prey items (Table 7) during summer. Caribou comprised
less than 5 percent of the food items.
Aerial observations made primarily during winter months
from 1976 to early 1979, indicated that moose and caribou were
important prey during winter. Of 23 observed kills, 11 were
moose, 12 were caribou, and 3 appeared to be small game species.
Caribou were most numerous within this pack's terri tory during
winter months and least common during summer months.
Fall moose sex and age composition count data indicate that
moose densities within this pack's territory were ~ite low. ~n
1976, only 158 moose were counted in the 697 mi (1,805 km )
Lake Louise Moose Management area. Undoubtedly mid-winter moose
densities were higher than those in fall because of moose
migration from the Alphabet Hills. Despite this, year-round
moose densities within this pack's large terri tory, were the
lowest of any of those pack's studied except for the Ewan and
St. Anne wolf packs. Therefore, it appeared that the Deep Lake
wolves were favoring moose over caribou. ·
Information pertaining to snowshoe hare, muskrat and beaver
population levels was scanty but reports from trappers and our
own observations suggest that the latter two species were quite
abundant during the study period. Snowshoe hare numbe:r-s were
low in this area from 1975-1978 and only in recent years have
they.appeared to increase. ·
Delta Pack -The summer diet of this pack in 1975, as
indicated by scats collected at both den and rendezvous site
(Tables 8 and 9), was co.mprised primarily of small mammals.
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Table 5. Incidence of food remains in wolf scats collected at the
Clearwater wolf den occupied during June 1976 in GMU 13
of southcentral Alaska.
Items in 2 Items in 31
adult scats :eu:e scats
Food item No. % occ. No. % occ.
Adult moose 0 0 1 3.0
Calf moose 2 100 28 84.8
Micro tine 0 0 1 3.0
Vegetation 0 0 3 9.1
Total 2 100.0 33 99.9
Grou:eed Data for 33 Scats
Food item Number of items Percent occurrence
Ungulate 31 88.6
Smal1 1 ~ammals 1 2.9
Other-3 8.6
Total 35 100.1
}) Includes birds, invertebrates, unidentified, wolf,
fish, vegetation.
Items in 33
combined scats
No. % occ.
1 2.9
30 85.7
1 2.9
3 8.6
35 100.1
45
Table 6. Incidence of food remains in wolf scats collected at the
Deadman Lake wolf rendezvous site occupied during mid-summer
1975 in GMU 13 of southcentral Alaska.
Food item
Adult moose
Calf moose
Beaver
Micro tine
Lynx
Total
Food item
Ungulate
Small mammals
Lynx
Total
Items in 1 Items in 29
adult scat EUE scats
No. % occ. No. % occ.
0 0 11 35.5
1 100 14 45.2
0 0 2 6.5
0 0 1 3.2
0 0 3 9.7 -
1 100.0 31 100.1
GrouEed Data for 30 Scats
Number of items
26
3
3
32
Percent occurrence
81.3
9.4
9.4
100.1
Items in 30
combined scats
No. % occ.
11 34.4
15 46.9
2 6.3
1 3.1
3 9.4
32 100.1
46
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Table 7. Incidence of food remains in wolf scats collected at the Deep
Lake wolf den occupied during late spring and summer 1975
in GMU 13 of southcentral Alaska.
Items in 34 Items in 72 Items in 106
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % 0 occ.
Adult moose 5 13.5 14 16.5 19 15.6
Calf moose 15 40.5 31 36.5 46 37.7
Caribou 1 2.7 2 2.4 3 2.5
Beaver 11 29.7 8 9.4 19 15.6
Snowshoe hare 3 8.1 13 15.3 16 13.1
Micro tine 0 0 2 2.4 2 1.6
Lynx 0 0 3 3.5 3 2.5
Vegetation 0 0 8 4.4 8 6.6
Fish 0 0 1 1.2 1 .8
Eggshells 2 5 /, 0 0 " 1.6 o'T "' Unidentified 0 0 3 3.5 3 2.5
Total 37 99.9 85 100.1 122 100.0
GrouEed Data for 106 Scats
Food item Number of items Percent occurrence
Ungulate 68 55.7
Small mammals 40 32.8
Other 14 11.5
Total 122 100.0
Table 8. Incidence of food remains in wolf scats collected at the Delta
River wolf den occupied during late spring and summer 1975 in
GMU 13 of southcentral Alaska.
Items in 24 Items in 113 Items in 137
adult scats :eu:e scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 10 25.1 10 8.1 20 12.3
Calf moose 4 10 35 28.5 39 23.9
Snowshoe hare 16 40.0 51 41.5 67 41.1
Beaver 3 7.5 21 17.1 24 14.7
Muskrat 0 0 1 .8 1 .6
Sciurids 2 5.0 1 .8 3 1.8
Eggshells 0 0 1 .8 1 .6
Bird 4 10.0 2 1.6 6 3.7
Insecta 0 0 1 .8 1 .6
Fish 1 2.5 0 0 1 .6
Total ·40 100.1 123 100.0 163 99.9
Grou:eed Data for 137 Scats
Food item Number of items Percent occurrence
Ungulate 59 36.2
Small mammals 95 58.3
Other 9 5.5
Total 163 100.0
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Table 9. Incidence of food remains in wolf scats collected at the
suspected rendezvous site of the Delta River wolf pack
occupied during summer 1975 in GMU 13 of southcentral Alaska.
Items in 13 Items in 67 Items in 80
adult scats J2UJ2 scats combined scats
Food item No. % occ. No. % occ. No. % o~c:.
Adult moose 2 14.3 2 2.6 4 4.3
Calf moose 4 28.6 5 6.4 9 9.8
Snowshoe hare 2 14.3 3 3.8 5 5.4
Beaver 0 0 45 57.7 45 48.9
Ground squirrel 6 42.9 15 19.2 21 22.8
Eggshells 0 0 1 1.3 1 1.1
Insecta 0 0 1 1.3 1 1.1
Micro tine 0 0 6 7.7 6 6.5
Total 14 100.1 78 100.0 92 99.9
GrouJ2ed Data for 80 Scats
Food item Number of items Percent occurrence
Ungulate 13 14.1
Small mammals 77 83.7
Other 2 2.2
Total 92 100.0
49
Snowshoe hare (41%), calf moose (24%) and beaver (15%) comprised
80 percent of the food items at the den site. At the rendezvous
site, however, beaver (48.9%} and ground squirrel (Spermophilus
undulatus) (22.8%) were the most important foods.
The Delta pack inhabited an area with the lowest ungulate
density of any· of the areas · studied. During radio-tracking
flights, only two ungulate kills were observed during 1975-76
and both were jointly occupied by brown bears. Moose movement
studies in similar habitat in other portions of GMU 13, suggest
that moose only utilize the pack area in late summer and fall,
spending winter and spring farther to the south in. areas with
less snowfall. Similarly, caribou were rarely observed in the
area.
As mentioned in the previous section, after winter 1975-76
this pack area was suspected of being vacant except for an
occasional public observation and our radio contact with wolves
063 and 064, which apparently dispersed shortly after being
radio-collared. Both pups and adults killed in 1975-76
exhibited characteristics of poor nutrition. .1 I 1:ne latter
observations were representative, then it would appear that the
Delta area is not· capable of supporting many wolves. Wolves
which do reproduce and survive are subjected to intense hunting
and trapping under ideal terrain and snow conditions.
Therefore, poor nutrition and hunting pressure will likely
preclude the establishment of a stable population of wolves in
this area in rhe near future.
Ewan Pack -From May 1975 through June 1976 the Ewan pack
was observed on 18 kills: 10 caribou, 7 moose and 1 red fox
(Vulpes vulpes). Following this period, radio contact withthis
pack was highly sporadic and never of sufficient duration to
determine numbers of caribou and moose being taken.
During late spring and early summer 1975, small mammals
appeared to comprise the bulk of the diet ( 65%) of this pack
while it was at the den site (Table 10}. Sixty-three percent of
the food items were either beaver or snowshoe hare. By the end
of July, however, when the pack moved to the first rendezvous
site, scat analysis indicated that calf moose (41% occurrence)
followed by beaver (33% occurrence) were the most important food
items (Table 11).
During 1975-76, there were indications that this pack may
have been nutritionally stressed. Throughout summer 1975 the
Ewan pups appeared smaller than those in other study packs.
Additionally, wolves 991 and 992 were thin and small when
collared, and, at best, were in only fair condition.
The Ewan pack area appeared to have the lowest year-round
moose density of any of the areas studied from 1975 through
. spring 1980. Moose sightings were rare during radio-tracking
flights and moose counts . in the Lake Louise count area also
indicated that moose numbers were quite low in this area.
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Table 10. Incidence of food items in wolf scats collected at the Ewan
Lake wolf den occupied during late spring and summer 1975
in GMU 13 of southcentral Alaska.
Items in 58 Items in 89 Items in 147
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. ~ occ.
Adult moose 9 12.9 4 3.7 13 7.3
Calf moose 15 21.4 22 20.6 37 21.0
Caribou 3 4.3 5 4. 7 8 4.5
Beaver 17 24.3 41 38.3 58 32.5
Snowshoe hare 22 31.4 32 29.9 54 30.5
Micro tine 2 2.8 1 0.9 3 1.7
Wolf 0 0 1 0.9 1 0.6
Eggshells 1 1.4 1 0.9 2 1.1
Bird 1 1.4 0 0 1 0.6
Total 70 99.9 107 99.9 177 99.8
GrouEed Data for 147 Scats
Food item Number of items Percent occurrence
Ungulate 58 32.8
Small mammals 115 65.0
Other 4 2.3
Total 177 100.1
5 1
Table 11. Incidence of food remains in wolf scats collected at the Ewan
Lake wolf rendezvous site occupied during mid-summer 1975
in GMU 13 of southcentral Alaska.
Items in 35 Items in 84 Items in 119
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 19 47.5 38 38.8 57 41.3
Calf moose 5 12.5 10 10.2 15 10.9
Beaver 12 30.0 34 34.7 46 33.3
Snowshoe hare 3 7.5 2 2.1 5 3.6
Wolf 0 0 2 2.1 2 1.4
Vegetation 1 2.5 10 10.4 11 8.0
Unidentified 0 0 2 2.1 2 1.4
Total 40 100.0 98 100.4 138 99.9
Grouoed Data for 119 Scats
-Food item Number of items Percent occurrence
Ungulate 72 52.2
Small mammals 51 37.0
Other 15 10.9
Total 138 100.1
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Following 1975-76 our contact with this pack was limited to
track counts and sporadic observations. These observations
suggested that the pack 1 s nutritional status had not improved
since it only numbered 3 to 5 animals until fall 1979 when it
numbered seven. From 1976 through mid-1980 large numbers of
caribou wintered on the Lake Louise flats. We suspect that
during these winters caribou were the major prey and probably
allowed for the continued existence of the pack. During years
when caribou did not winter in this area, this pack appeared to
dwindle in number, probably due to starvation and dispersal.
Beaver, muskrat and, in recent years, snowshoe hare have
been numerous in this area. Only snowshoe hare, however, could
be viewed as an important alternate winter food source.
Hogan Hill Pack -During 1975-76 and from March 1977
through January 1979, members of the Hogan Hill pack were
observed on 39 kills. Adult or yearling moose appeared to be
the most important year-round food item, comprising 74 percent
of the observed kills. Observations of prey taken during summer
1978 \vere summarized a..11.d compared with those of the Mendel tna
pack in a paper presented at the Portland Wolf Symposium
entitled "Gray wolf-brown bear relationships in the Nelchina
Basin of Southcentral Alaska. 11 A copy of the paper is presented
as Appendix III.
A comparison of food items contained in wolf scats
collected at the 1975 and 1978 den sites is contained in
Tables 12 and 13. In 1975 ungulates, primarily moose, comprised
55 percent of the prey items, whereas in 1978 ungulates
comprised 73 percent of the food items. The main difference
between these years was the percent occurrence of calf moose and
snowshoe hare: 29 percent calf moose and 40 percent snowshoe
hare in 1975 and 56 percent calf moose and 5 percent snowshoe
hare in 1978. These data indicate that more calves were taken
in 1978 than in 1975. By July 1978, the percent of calf moose
in scats from the rendezvous site increased to 81 percent
(Table 14). High summer dependence on newborn moose calves in
1978 was not supported by our aerial observations which
indicated a predominance of adult and yearling moose.
Discrepancies between these data will be discussed in the Food
Habits Summary section.
The Hogan Hill pack area supports one of t~e highest moose
densities in GMU 13; in 1978 1.1 moose per mi were observed
during fall composition counts. Therefore, this pack 1 s high
year-round dependency on moose was not surprising. Caribou were
probably available to this pack on a year-round basis but
usually in very low densities except during spring and fall
migrations to and from the Wrangell Mountains. Beaver and
muskrat populations appeared to be lower than those found in
adjacent pack areas, primarily because there is less aquatic
habitat. No information is available on snowshoe hare
populations but it appears probable that in 1979 and 1980 hare
populations increased as they have elsewhere in GMU 13.
53
Table 12. Incidence of food items in wolf scats collected at the Hogan
Hill wolf den occupied in late spring and early summer 1975
in GMU 13 of southcentral Alaska.
Items in 39 Items in 96 Items in 135 -
adult scats :eu:e scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 11 22.9 9 8.1 20 12.6
Calf moose 15 31.3 31 27.9 46 28.9
Caribou 4 8.3 18 16.2 22 13.8
Snowshoe hare 16 33.3 48 43.2 64 40.3
Wolf 0 0 1 ·I.O 1 0.6
Vegetation 1 2.0 3 2.7 4 2.5
Eggshells 1 2.0 1 1.0 2 1.3
Total 48 99.8 111 100.1 159 100.0
Grou:eed Data for 135 Scats
Food item Number of items Percent occurrence
. Ungulate
Small mammals
Other
Total
88
64
7
159
55.3
40.3
4.4
100.0
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Table 13. Incidence of food remains in wolf scats collected at the
Hogan Hill wolf den occupied during late spring and summer
1978 in GMU 13 of southcentral Alaska.
Items in 126 Items in 77 Items in 203
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 24 16.0 2 2.3 26 10.9
Calf moose 73 48.7 59 67.0 132 55.5
Adult caribou 10 6.7 2 2.3 12 5.0
Calf caribou 1 0.7 2 2.3 3 1.3
Beaver 19 12.7 8 9.1 27 11.3
Muskrat 3 2.0 2 2.3 5 2.1
Snowshoe hare 11 7.3 1 1.1 12 5.0
Parka squirrel 0 0 0 0 0 0
Micro tine 2 1.3 1 1.1 3 1.3
Bird 0 0 1 1.1 1 0.4
Fish 0 0 0 0 0 0
Unidentifiable 3 2.0 5 5.7 8 3.4
Vegetation 4 2.7 5 5.7 9 3.8
Other 0 0 0 0 0 0 --
Total 150 100.1 88 100.0 238 100.0
GrouEed Data for 203 Scats
Food item Number of items Percent occurrence
Ungulate 173 72.7
Small mammals 47 19.7
Other 18 7.6
Total 238 100.0
55
l
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Table 14. Incidence of food remains in wolf scats collected at the n
Hogan Hill wolf rendezvous site occupied during mid-summer
'" 1978 in GMU 13 of southcentral Alaska.
Items in 17 Items in 24 Items in 41
adult scats ]2U]2 scats combined scats
Food item . No. % No. % occ. No. % occ. <=
0 OCC. -
E
Adult moose 3 12.5 1 3.4 4 7.5 . -,.-
Calf moose 19 79.2 24 82.8 43 81.1
Adult caribou 0 0 0 0 0 0 ~
Calf caribou 0 0 0 0 0 0 -· Beaver 0 0 0 0 0 0 = ~
Muskrat 0 0 0 0 0 0
Snowshoe hare 1 4.2 0 0 1 1.9
Parka squirrel 0 0 0 0 0 0 r
Micro tine 0 0 0 0 0 0
Bird 0 0 0 0 0 0 t:
Fish (\ 0 (\ (\ (\ (\ v v v v v
Other 0 0 2 6.9 2 3.8 r Vegetation 1 4.2 1 3.4 2 3.8 t
Unidentifiable 0 0 1 3.4 1 1.9
Total 24 100.1 29
;
99.9 53 100.0
GrouDed Data for 41 Scats
.Food item Number of items Percent occurrence
~
Ungulate 47 88.7 ~
Small mammals 1 1.9 L
Other 1 9.4
' '
Total 53 100.0 r
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·Jay Creek Pack -Limited aerial observations of this pack
suggested a high dependence on adult moose ( 7 of 10 kills)
during winter. During winter months the pack frequented the
riparian areas along the Susi tna River where both resident and
migratory moose wintered (Ballard and Taylor 1980). Although we
only observed one caribou kill, caribou were probably an
important food item in summer and during migrations to and from
the calving grounds on Kosina Creek. Although the pack denned
in 1978, the site was not visited for scat collection because of
logistic problems.
Keg Creek Pack -Members of the Keg Creek pack were
observed on 28 kills during this study. Moose comprised 79
percent of the observed kills and 86 percent of these were adult
moose. Abundance of moose and caribou in this area was similar
to that described for the Hogan Hill pack area and, therefore,
the high predominance of moose kills was not surprising since
moose were the most common ungulate prey.
Summer food habits, as indicated by scat analyses from den
and rendezvous. sites, are summarized in Tables 15 through 18.
Percent occurrence of ungulate food items at the same den site
used in 1975, 1976, and 1978 was 67, 92, and 53 percent,
respectively. In all cases calf moose comprised the predominant
food item (58, 73, and 53%). small mammals, particularly
snowshoe hares, were an important prey item in 1975 (28%) but
not in either 1976 or 1978. The decline in percent occurrence
of snowshoe hare was similar to that exhibited by the Hogan Hill
pack during the same years suggesting that perhaps hare
abundance declined in these pack areas. Also similar was the
increase in percent occurrence of calf moose remains in scats
collected at the den · compared to those collected at the
rendezvous site. This might indicate that movement to the
rendezvous sites was partially related to calf moose abundance.
Regardless, calf moose were an important summer food item during
the 3 years of scat analyses.
Maclaren River Pack -From April 1975 to mid-January 1976,
members of the Maclaren River pack were observed on 12 kills:
10 moose and two caribou, suggesting a winter dependency on
moose. Moose populations within this pack's area were both
sedentary and migratory, with the migratory segment involving
moose from the Clearwater Mountains (Ballard and Taylor 1980).
Densities observed qpring fall composition counts were not high:
0.4 moose per mi were observed in 1976. Caribou were
seasonally abundant and available to this pack primarily during
late winter.
Although this pack denned in 1975, the den site was not
located and thus no scats were collected. A small number of
scats (n = 26) collected from the 1975 rendezvous site suggested
that moose were also the most important prey i tern during summer
(Table 19).
5 7
Table 15. Incidence of food remains in wolf scats collected at the Keg
Creek wolf den occupied during late spring and early summer
1975 in GMU 13 of southcentral Alaska.
Items in 17 Items in 65 Items in 82
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 4 21.1 4 5.8 8 9.1
Calf moose 12 63.2 39 56.5 51 58.0
Beaver 0 0 8 11.6 8 9.1
Snowshoe hare 2 10.5 15 21.7 17 19.3
Wolf 0 0 1 1.4 1 1.1
Eggshells 1 5.3 1 1.4 2 2.3
Unidentified 0 0 1 1.4 1 1.1
Total 19 100.1 69 99.8 88 100.0
Grouoed Data for 82 Scats
Food item .Number of items Percent occurrence
Ungulate 59 67.0
Small mammals 25 28.4
Other 4 4.5
Total 88 99.9
5 8
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Table 16. Incidence of food remains in wolf scats collected at the Keg
Creek wolf rendezvous site occupied in mid-summer 1975 in
GMU 13 of southcentral Alaska.
Items in 12 Items in 114 Items in 126
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 9 64.3 15 12.1 24 17.4
Calf moose 4 28.6 96 77.4 100 72.5
Caribou 0 0 1 1.0 1 1.0
Snowshoe hare 0 0 10 8.1 10 7.2
Vegetation 1 7.1 1 1.0 2 1.4
Bird 0 0 1 1.0 1 1.0
Total 14 100.0 124 100.6 138 100.5
GrouEed Data for 126 Scats
Food item Number of items Percent occurrence
Ungulate 125 90.6
Small mammals 10 7.2
Other 3 2.3
Total 138 100.1
5 9
J
Table 17. Incidence of food remains in wolf scats collected at the
Keg Creek wolf den occupied during late spring and summer
1976 in GMU 13 of southcentral Alaska. J
Items in 36 Items in 45 Items in 81 l
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
' i D
Adult moose 4 10.5 1 2.1 5 5.9
Calf moose 21 55.3 41 87.2 62 72.9
Adult caribou 5 13.2 1 2.1 6 7.1
Calf caribou 4 10.5 1 2.1 5 5.9
Beaver 2 5.3 0 0 2 2.4
Unidentifiable 1 2.6 2 4.3 3 3.5
Vegetation 1 2.6 1 2.1 2 2.4
Total 38 100.0 47 99.9 85 100.1
Grou2ed Data for 81 Scats
F
Food item Number of items Percent occurrence ,
L
Ungulate 78 91.8
Small mammals 2 2.4
Other 5 5.9
Total 85 100.1
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Table 18. Incidence of food remains in wolf scats collected at the
Keg Creek wolf den occupied during late spring and summer
1978 in GMU 13 of southcentral Alaska.
Items in 7 Items in 38 Items in 45
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 1 14.3 0 0 1 2.0
Calf moose 4 57.1 22 52.4 26 53.1
Adult caribou 0 0 1 2.4 1 2.0
Calf caribou 1 14.3 8 19.0 9 18.4
Beaver 1 14.3 0 0 1 . 2 .. 0
Muskrat 0 0 0 0 0 0
Snowshoe hare 0 0 0 0 0 0
Ground squirrel 0 0 0 0 0 0
Micro tine 0 0 1 2.4 1 2.0
Bird 0 0 0 0 0 0
Fish 0 0 0 0 0 0
Unidentifiable 0 0 9 21.4 9 18.4
Vegetation 0 0 0 0 0 0
Other 0 0 1 2.4 1 2.0
Total 7 100.0 42 100.0 49 99.9
Grouped Data for 45 Scats
Food item Number of items Percent occurrence
·Ungulate 37 75.5
Small mammals 2 4.1
Other 10 20.4
Total 49 100.0
6 1
Table 19. Incidence of food remains in wolf scats collected at the
Maclaren River wolf rendezvous site occupied during
mid-summer 1975 in GMU.13 of southcentral Alaska.
Items in 6 Items in 20 Items in 26
adult scats EUE scats combined scats
Food item No. % occ .. No. % occ. No. % occ.
Adult moose 5 83.3 3 11.5 8 25.0
Calf moose 0 0 14 53.8 14 43.8
Beaver 0 0 1 3.8 1 3.1
Snowshoe hare 1 16.7 3 11.5 4 12.4
Micro tine 0 0 2 7.7 2 6.3
Bird 0 0 2 7.7 2 6.3
Snail 0 0 1 3.8 1 3.1 -
Total 6 100.0 26 99.8 32 100.0
Grou:eed Data for. 26 Scats
Food item ·Number of items Percent occurrence
Ungulate 22 68.8
Small mammals 7 21.9
Other 3 n /.
::1o'+
Total 32 100.1
1
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Mendeltna Pack -From August 1976 through November 1977
members of the Mendeltna wolf pack were observed at 36 kills.
Overall, adult and calf moose comprised 67 percent of the kills,
followed by caribou at 18 percent. Beavers, rodents, brown
bears and unknown species comprised the remaining 18 percent of
the observed kills. Of the 36 moose kills, at least nine (25%)
were also shared by one, or more, brown bear.
Moose in moderate densities (0.9 moosejmi 2 observed during
fall composition counts) were year-round inhabitants of the
Mendeltna area. Although no moose movement studies have been
conducted in this area, movements of cows with radio-collared
calves suggested that this population was comprised of both
sedentary and migratory segments. The migratory segment
appeared to migrate from summer to winter range in an east-west
direction. Many of the migratory moose, however, appeared to
summer and winter in this pack's terri tory. Overall/ moose
appeared to be the most abundant year-round residents.
Caribou were also available to this pack on. a year-round
basis but appeared to be most numerous during winter when
portions of the·Nelchina herd wintered on the Lake Louise flats.
They were also quite abundant in late summer in the eastern
Talkeetna foothills following the calving season.
Data pertaining to populations of small mammals were not
collected but it appeared that musJcrats were nu..tnerous in many
ponds while beavers were restricted to creek drainages.
Snowshoe hare numbers have increased since 1978 but hares were
present in low numbers previously ..
A detailed discussion of summer food habits as suggested by
aerial observations is presented in APPendix I I I. These data
indicated that yearling-and adult. moose were the predominant
prey during late May and June rather than newborn calves.
Summer food habits, as indicated by scats collected from
den and rendezvous sites in 1976 and 1977, are depicted in
Tables 20 through 25. In 1976, caribou were the major food item
(58%) while in 1977 moose were the predominant item {44%) at the
main den. In both years calves were the most important age
class represented. At all rendezvous sites, calf moose were the
predominant prey i tern. Small mammals comprised less than 10
percent of the food i terns during all years and at all sites,
except for the 1977 main den and the Kelly Lake rendezvous site
where they occurred in 16 to 19 percent of the food items.
St. Anne Pack -From April 1977 through June 1980, members
of the st. Anne's pack were observed on 20 kills, most of which
were observed during winter. Sixteen (80%) of the kills were
moose, of which 75 percent were adults, suggesting that moose
were by far the most important winter food item.
6 3
Table 20. Incidence of food remains in wolf scats collected at the
Mendeltna wolf den occupied during late spring and summer
1976 in GMU 13 of southcentral Alaska.
Items in 28 Items in 24 Items in 52
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 4 13.3 1 3.7 5 8.8
Calf moose 10 33.3 6 22.2 16 28.1
Adult caribou 4 13.3 6 22.2 10 17.5
Calf caribou 10 33.3 13 48.1 23 40.4
Beaver 1 3.3 0 0 1 1.8
Snowshoe hare 1 3.3 0 0 1 1.8
Vegetation 0 0 1 3.7 1 1.8
Total 30 99.8 27 99.9 57 100.2
Grouped Data for 52 Scats
Food item . Number of items Percent occurrence
Ungulate
Small mammals
Other
Total
54
2
1
57
94.7
3.5
1.8
100.1
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Table 21. Incidence of food remains in wolf scats collected at the
Mendeltna wolf pack rendezvous site near Daisy Creek occupied
during summer 1976 in GMU 13 of southcentral Alaska.
Items in 25 Items in 62 Items in 87
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 4 13.8 3 4.7 7 7.5
Calf moose 15 51.7 36 56.3 51 54.8
Adult caribou 4 13.8 5 7.8 9 9.7
Calf caribou 4 13.8 9 14.1 13 14.0
Muskrat 0 0 2 3.1 2 2.2
Snowshoe· hare 1 3.4 1 1.6 2 2.2
Micro tine 0 0 1 1.6 1 1.1
Red squirrel 0 0 1 1.6 1 1.1
(Tamiasciurus hudsonicus)
Unidentifiable 0 0 3 4.7 3 3.2
Vegetation 1 3.4 2 3.1 3 3.2
Bird 0 0 1 1.6 1 1.1 --
Total 29 99.9 64 100.2 93 100.1
GrouEed Data for 87 Scats
Food item Number of items Percent occurrence
Ungulate 80 86.0
Small mammals 6 6.5
Other 7 7.5
Total 93 100.0
6 5
Table 22. Incidence of food remains in wolf scats collected at the
main Mendeltna wolf den occupied during late spring and early
summer 1977 in GMU 13 of southcentral Alaska.
Items in 40 Items in 72 Items in 112
adult scats :eu:e scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 0 0 2 2.1 2 1.3
Calf moose 26 4.4.8 41 42.3 67 43.2
Adult caribou 4 6.9 2 2. 1 6 3.9
Calf caribou 8 13.8 9 9.3 17 11.0
Beaver 0 0 2 2.1 2 1.3
Muskrat 1 1.7 5 5.2 6 3.9
Snowshoe hare 5 8.6 4 4.1 9 5.8
Parka squirrel 1 1.7 0 0 1 .6
Micro tine 6 10.3 5 5.2 11 7.1
Other 0 0 4 4.1 4 2.6
Vegetation 7 12.1 20 20.6 27 17.4
Unidentifiable 0 0 3 3.1 3 1.9
Total 58 99.9 97 100.2 155 100.0
Grou:eed Data for 112 Scats
,..,..,..A ..;+-om }lumber of items Percent occurrence ..&. VVY. ..... ~--'"
Ungulate 92 59.4
Small mammals 29 18.7
Other 34 21.9
Total 155 100.0
6 6
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Table 23. Incidence of food remains in wolf scats collected at the
Kelly Lake rendezvous site occupied by the Mendeltna wolf
pack during mid-summer 1977 in GMU 13 of southcentral Alaska.
Items in 17 Items in 76 Items in 93
adult scats :eu:e scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 2 9.1 2 2.3 4 3.7
Calf moose 13 59.1 54 62.1 67 61.5
Calf caribou 1 4.5 5 5.7 6 5.5
Beaver 0 0 1 1.1 1 .9
Muskrat 2 9.1 2 2.3 4 3.7
Snowshoe hare 0 0 5 5.7 5 4.6
Micro tine 2 9.1 5 5.7 7 6.4
Bird 0 0 2 2.3 2 1.8
Insect 0 0 1 1.1 1 .9
Other 1 4.5 9 10.3 10 9.2
Vegetation 1 4.5 1 1.1 'l 1.8 .l. "" Unidentifiable 0 0 0 0 0 0
Total 22 99.9 87 99.7 109 100.0
Grou12ed Data for 93 Scats
Food item Number of items Percent occurrence
Ungulate 77 70.6
Small mammals 17 15.6
Other 15 13.8
Total 109 100.0
67
Table 24. Incidence of food remains in wolf scats collected at the
Nickolson Lake wolf den used by the Mendeltna wolf pack in
late spring and summer 1977 in GMU 13 of southcentral Alaska.
Items in 66 Items in·20 Items in 86
adult scats. EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 9 12.9 1 4.5 10 10.9
Calf moose 48 68.6 16 72.7 64 69.6
Calf caribou 4 5.7 2 9.1 6 6.5
Muskrat 0 0 1 4.5 1 1.1
Snowshoe hare 3 4.3 1 4.5 4 4.3
Red squirrel 1 1.4 0 0 1 1.1
Wolf 2 2.9 1 4.5 3 3.3
Vegetation 1 1.4 0 0 1 1.1
Unidentifiable 2 2.9 0 0 2 2.2
Total 70 100.1 22 99.8 92 100.1
Grouped Data for 86 Scats
Food item Number of items Percent occurrence
Ungulate 80 ~7 (\ _, ......
Small mammals 6 6.5
Other 6 6.5
Total 92 100.0
. -
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Table 25. Incidence of food remains in wolf scats collected at the
Nickolson Lake wolf rendezvous site utilized by members of
the Mendeltna wolf pack during summer 1977 in GMU 13 of
southcentral Alaska.
Items in 49 Items in 59 Items in 108
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 3 4.8 2 2.9 5 3.8
Calf moose 40 64.5 49 70.0 89 67.4
Adult caribou 1 1.6 0 0 1 .8
Calf caribou 1 1.6 4 5.7 5 3.8
Muskrat 2 3.2 1 1.4 3 2.3
Snowshoe hare 1 1.6 1 1.4 2 1.5
Parka squirrel 0 0 1 1.4 1 .8
Micro tine 1 1.6 3 4.3 4 3.0
Bird 5 8.1 2 2.9 7 5.3
Veget1Jion 0 0 4 5.7 4 3.0
Other-4 6.5 0 0 4 3.0
Unidentifiable 4 6.5 3 4.3 7 5.3
Total 62 100.0 70 100.0 132 100.0
GrouEed Data for 108 Scats
Food item Number of items Percent occurrence
Ungulate 100 75.8
Small mammals 10 7.6
Other 22 16.7
Total 132 100.1
}j Includes wolf hair, eggshell.
69
Moose numbers within this pack area were similar to those
in the Ewan and Deep Lake areas. In 1979, only 0. 2 moose per
mi 2 were observed in the 517 mi 2 (1,339 km 2 ) moose count area.
This moose population appears to be declining still further,
since calf: 100 cow ratios did not exceed 5 from 1975 through
1978. Reasons for the low calf survival are not known, but in
addition to wolf predation and declining range conditions, the
area probably supports one of the densest bear populations (both
brown and black [Ursus americanus]) in GMU 13. Therefore, bear
predation is probably a significant contributing factor.
During this study, caribou were rarely available to the St.
Anne pack. From 1977 through June 1980, we recorded only three
observations of caribou south of the Tazlina River. Small game,
however, appeared quite abundant. Based upon observations it
appeared that this pack area supported a relatively abundant
hare population during the study. Both beaver and muskrat were
numerous in ponds and along creeks. Ptarmigan and ground
squirrels were numerous in some areas.
Summer food habits, as determined from scats collected at
St. Anne den site·s from 1975 through 1978, are presented in
Tables 26 through 29. During this 4-year period the importance
of ungulate food items varied from 25 percent in 1978 to 50
percent in 1977. In all years, calf moose were the predominant
ungulate food item. The importance of small mammals and birds
to this pack ;~as greater than for any of the other packs
studied. Both snowshoe hare and beaver were important summer
food items.
As mentioned. in the previous section, the St. Anne pack
exhibited considerable annual fluctuations in wolf numbers,
primarily due to dispersal. Because of the limited availability
of ungulate prey and the relatively heavy reliance on small
mammals, we suspect that pack size in this area was governed at
least partially by prey availability. Similar to the Delta and
Ewan packs, which also relied heavily on small mammals, pack
numbers appeared to be unstable. It would appear that areas
with relatively low moose and caribou populations have a low
wolf carrying capacity regardless of the availability of smaller
alternate prey.
Sinona Pack -During this study members of the Sinona wolf
pack were observed at 40 kills. Moose comprised all but two
(95%) of the kills, others were one beaver and one of
unidentified species which may also have been a moose. Ages of
the 38 identified moose kills were aged as follows: 19 adults,
9 unclassified, 8 calves, and 2 yearlings. Moose were clearly
the most important prey species in this area.
Moose were by far the most abundant year-round ungulate
species in this area. Moose densities undoubtedly were greater
in winter than summer due to seasonal migrations of moose from
the upper Gakona River and_the eastern Alphabet Hills
70
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Table 26. Incidence of food remains in wolf scats collected at the
St. Anne wolf den occupied during late spring and early
summer 1975 in GMU 13 of southcentral Alaska.
Items in 106 Items in 126 Items in 232
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No . % occ.
Adult moose 9 6.0 15 8.3 24 7.3
Calf moose 30 20.0 31 17.2 61 18.5
Beaver 1 .7 8 4.4 9 2.7
Muskrat 5 3.3 5 2.8 10 3.0
Snowshoe hare 92 61.3 103 57.2 195 59.1
Red squirrel 2 1.3 0 0 2 .6
Micro tine 3 2.0 13 7.2 16 4.8
Fish 2 1.3 1 .6 3 .9
Bird 6 4.0 1 .6 7 2.1
Vegetation 0 0 3 1.7 3 .9
Total 150 99.9 180 100.0 330 99.9
Grou,Eed Data for 232 Scats
Food item Number of items Percent occurrence
Ungulate 85 25.8
Small mammals 232 70.3
Other 13 3.9
Total 330 100.0
7 1
'-'---
Table 27. Incidence of food remains in wolf scats collected at the
St. Anne wolf den occupied during late spring and summer
1976 in GMU 13 of southcentral Alaska.
Items in 91 Items in 85 Items in 176
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ. F
-
~
..
Adult moose 9 6.8 3 3.1 12 5.2 ·-
Calf moose 38 28.6 44 44.9 82 35.5
Adult caribou 3 2.3 2 2.0 5 2.2 L
Calf caribou 5 3.8 7 7.1 12 5.2 .....
Beaver 14 10.5 3 3.1 17 7.4 ~
Muskrat 1 .8 6 6.1 7 3.0 ·-Snowshoe hare 42 31.6 7 7.1 49 21.2
Parka squirrel 1 .8 1 1.0 2 .9 r
Micro tine 9 6.8 3 3.1 12 5.2 L Bird 1 .8 0 0 1 .4
Other 4 3.0 7 7.1 11 4.8 F Unidentifiable 4 3.0 10 10.2 14 6.1 L Vegetation 2 1.5 5 5.1 7 3.0 --
Total 133 100.3 98 99.9 231 100.1 r
Grou:eed Data for 176 Scats
Food item Number of items Percent occurrence
'---
Ungulate 111 48.1 r
Small mammals 87 37.7 L-
Other 33 14.3
r-
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Total 231 100.1 L.
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Table 28. Incidence of food remains in wolf scats collected at the
St. Anne wolf den occupied during late spring and summer
1977 in GMU 13 of southcentral Alaska.
Food item
Adult moose
Calf moose
Beaver
Muskrat
Snowshoe hare
Micro tine
Unidentifiable
Other
Total
Food item
Ungulate
Small mammals
Other
Total
Items in 26 Items in 23
adult scats pup scats
No. % occ. No. % occ.
2 6.5 0 0
12 38.7 13 56.5
8 25.8 4 17.4
1 3.2 1 4.3
5 16.1 3 13.0
1 3.2 1 4.3
1 3.2 0 0
1 3.2 1 4.3
31 99.9 23 99.8
Grouped Data for 49 Scats
Number of items
27
24
3
54
Percent occurrence
50.0
44.4
5.6
100.0
Items in 49
combined scats
No. % occ.
2 3.7
25 46.3
12 22.2
2 3.7
8 14.8
2 3.7
1 1.9
2 3.7
54 100.0
7 3
,!""=1
Table 29. Incidence of food remains in wolf scats collected at the
St. Anne wolf den occupied during late spring and summer <:"
1978 in GMU 13 of southcentral Alaska.
Items in 68 Items in 62 Items in 130
adult scats J2UJ2 scats combined scats = !C Food item No. % occ. No. % occ. No. % occ.
E . ' ·~ Adult moose 3 3.5 0 0 3 1.8 . '
Calf moose 21 24.4 16 20.5 37 22.6 ~-
Adult caribou 1 1.2 0 0 1 0.6
Calf caribou 0 0 0 0 0 0 -~
Beaver 23 26.7 12 15.4 35 21.3
Muskrat 7 8.1 10 12.8 17 10.4 L
Snowshoe hare 16 18.6 14 17.9 30 18.3
Parka squirrel 0 0 0 0 0 0 r
Red Squirrel 0 0 2 2.6 2 1.2 ~
Micro tine 2 2.3 3 3.8 5 3.0
Bird 6 7.0 9 11.5 15 9.1 f Fish 0 0 1 1.3 1 0.6 L Vegetation 4 4.7 3 3.8 7 4.3
Other 0 0 3 3.8 3 1.8 r Unidentifiable 3 3.5 5 6.4 8 4.9 '
'-
Total 86 100.0 78 99.8 164 99.9
Grou:12ed Data for 130 Scats
'-
Food item Number of items Percent occurrence r
L
Ungulate 41 25.0
Small mammals 89 54.3 r-
~-
Other 34 20.7 ' E:;
Total 164 100.0
7 4
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(VanBallenberghe 1978). Caribou were probably also available on
a · year-round basis but at very low densities except during
caribou migration to arid from the Wrangell Mountains each fall
and spring. Beaver and muskrat appeared to be fairly abundant
in· the area, while snowshoe hares were abundant in scattered
pockets of habitat in the southern half of the territory.
Summer food habits as determined from scats collected at
den and rendezvous sites f.rom 1975 through 1978 are presented in
Tables 30 through 35. As suggested by data obtained from aerial
observations, moose of all ages were by far the most prevalent
food item at all sites and in all years, ranging from 54 to 85
percent occurrence. Calf moose were the most prevalent age
class. Beavers were the next most important prey item of this
pack.
Sus i tna Pack -From February 19 7 9 through June 198 0 , the
Susi tna pack was observed on 43 kills which were comprised of
the following: 16 calf moose, 12 adult moose, 7 adult caribou,
5 unclassified caribou, 1 unclassified moose, 1 bear, and 1
unidentified species. Moose were the most important prey
species during both years of study; however, in 1979-80 caribou
were more important than in the previous year.
Moose were available to this pack in low numbers on a
year-round basis except during winter when large numbers of
migratory moose overwintered in the area (Ballard and Taylor
1980). Numbers of moose in relation to the rates of predation
by this pack will be discussed further in the Predation Rate
section of this report. Caribou were available to this pack
primarily' during late winter when large numbers wintered on the
Lake Louise flats. They were also available during spring and
fall migration and during summer in the southwestern extremes of
the territory following the calving season. Beaver and muskrat
were available during summer but snowshoe hare numbers were low
relative to other areas in GMU 13.
No scats were collected at the 1979 den site.
Tolsona Pack -During this study the Tolsona Pack was
observed at 29 kills, 48 percent of which were caribou. Moose
comprised 41 percent of the total kills (calves 33%), suggesting
that the Tolsona pack relied on moose less than most of the
other packs studied.
Similar to the Deep Lake terri tory, the area occupied by
this pack from 1977 to early 1979 had one of the lowzr moose
densities in GMU 13 : 0. 2 moose were observed per mi during
1976 composition counts. . Consequently this pack relied to a
large extent on prey other than moose. Caribou were undoubtedly
the most abundant ungulate during winter when large numbers of
caribou wintered on the Lake Louise flats. Beaver and muskrat
were available during warmer months in relatively high numbers.
Snowshoe hares were also. present and probably increased in
number after 1977.
7 5
Table 30. Incidence of food remains in wolf scats collected 6 September
1975 at the Sinona Creek wolf den occupied in late spring and
summer 1975 in GMU 13 of southcentral Alaska.
Items in 18 Items in 110 Items in 128
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 7 31.8 32 27.6 39 28.3
Calf moose 10 45.5 67 57.8 77 55.8
Caribou 0 0 1 0.9 1 0.7
Beaver 3 13.6 10 8.6 13 9.4
Snowshoe hare 1 4.5 1 0.9 2 1.4
Micro tine 0 0 1 ·o. 9 1 0.7
Vegetation 0 0 1 0.9 1 0.7
Eggshells 1 4.5 1 0.9 2 1.4
Bird 0 0 1 0.9 1 0.7
Unidentified 0 0 1 0.9 1 0.7
Total 22 99.9 116 100.3 138 99.8
GrouEed Data for 128 Scats
Food item Number of items Percent occurrence
Ungulate 117 84.2
Small mammals 16 11.5
Other 6 4.3
Total 139 100.0
7 6
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Table 31. Incidence of food remains in wolf scats collected at the
Sinona Creek wolf pack den occupied during late spring
and summer 1976 in GMU 13 of southcentral Alaska.
Items in 34 Items in 79 Items in 113
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 5 11.6 14 15.1 19 14.0
Calf moose 19 44.2 47 50.5 66 48.5
Calf caribou 1 2.3 0 0 1 . 7
Beaver 5 11.6 5 5.4 10 7.4
Muskrat 1 2.3 5 5.4 6 4.4
Snowshoe hare 4 9.3 1 1.1 5 3.7
Parka squirrel 1 2.3 1 1.1 2 1.5
Micro tine 1 2.3 1 1.1 2 1.5
Fox 1 2.3 0 0 1 .7
Unidentifiable 0 0 10 10.8 10 7.4
Vegetation 5 11.6 9 9.7 14 10.3
Total 43 99.8 93 100.2 136 100.1
GrouEed Data for 113 Scats
Food item Number of items Percent occurrence
Ungulate 86 63.2
Small mammals 25 18.4
Other 25 18.4
Total 136 100.0
7 7
Table 32. Incidence of food remains in wolf scats collected at the
Sinona Creek wolf pack rendezvous site occupied in 1976
in GMU 13 of southcentral Alaska.
Items in 17 Items in 57 Items in 74
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 1 4.8 0 0 1 1.2
Calf moose 16 76.2 47 78.3 63 77.8
Beaver 3 14.3 2 3.3 5 6.2
Muskrat 0 0 3 5.0 3 3.7
Snowshoe hare 0 0 1 1.7 1 1.2
Micro tine 0 0 1 1.7 1 1.2
Bird 1 4.8 0 0 1 1.2
Unidentifiable 0 0 3 5.0 3 3.7
Vegetation 0 0 3 5.0 3 3.7
Total 21 100.1 60 100.0 81 99.9
. GrouEed Data for 74 Scats
Food item Number of items Percent occurrence
Ungulate 64 -.n 1'\
/-:7.U
Small mammals 10 12.3
Other 7 8.6
Total 81 99.9
7 8
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Table 33. Incidence of food remains in wolf scats collected at a second
Sinona Creek wolf pack den site occupied during summer 1976
in GMU 13 of southcentral Alaska.
Items in 24 Items in 1 Items in 25
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 21 84.0 1 100.0 22 84.6
Beaver 1 4.0 0 0 1 3.8
Unidentifiable 3 12.0 0 0 3 11.5 -
Total 25 100.0 1 100.0 26 99.9
GrouEed Data for 25 Scats
Food item Number of items Percent occurrence
Ungulate 22 84.6
Small mammals 1 3.8
Other 3 11.5
Total 26 99.9
7 9
Table 34. Incidence of food remains in wolf scats collected at the
Sinona wolf den occupied in late spring and summer 1977
in GMU 13 of southcentral Alaska.
Items in 16 Items in 136 Items in 152
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 2 10.5 7 4.3 9 4.9
Calf moose 11 57.9 94 57.3 105 57.4
Beaver 3 15.8 20 12.2 23 12.6
Muskrat 1 5.3 10 6.1 11 6.0
Snowshoe hare 0 0 7 4.3 7 3.8
Micro tine 1 5.3 1 .6 2 1.1
Bird 0 0 1 .6 1 .5
Porcupine 0 0 1 .6 1 .5
Other 1 5.3 3 1.8 4 2.2
Vegetation 0 0 12 7.3 12 6.6
Unidentifiable 0 0 8 4.9 8 4.4
Total 19 100.1 164 100.0 183 100.0
Grou:eed Data for 152 Scats
Food item Number of items Percent occurrence
Ungulate 114 62.3
Small mammals 43 23.5
Other 26 14.2
Total 183 100.0
80
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Table 35. Incidence of food remains in wolf scats collected at th~
Sinona wolf den occupied during late spring and summer
1978 in GMU 13 of southcentral Alaska.
Items in 54 Items in 105 Items in 159
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 3 4.3 5 4.2 8 4.3
Calf moose 32 45.7 55 46.6 87 46.3
Adult caribou 1 1.4 0 0 1 0.5
Calf caribou 2 2.9 3 2.5 5 2.7
Beaver 11 15.7 13 11.0 24 12.8
Muskrat 5 7.1 11 9.3 16 8.5
Snowshoe hare 6 8.6 4 3.4 10 5.3
Porcupine 0 0 0 0 0 0
(Erithiazan idorsatum)
Ground squirrel 0 0 1 0.8 1 0.5
Microtine 0 0 1 0.8 1 0.5
Bird 2 2.9 3 2.5 5 2.7
Insect 0 0 0 0 0 0
Fish 0 0 0 0 0 0
Other 0 0 5 4.2 5 2.7
Vegetation 5 7.1 7 5.9 12 6.4
Unidentifiable 3 4.3 10 8.5 13 6.9
Total 70 100.0 118 99.7 188 100.1
Grou12ed Data for 159 Scats
Food item Number of items Percent occurrence
Ungulate 101 53.7
Small mammals 52 27.7
Other 35 18.6
Total 188 100.0
8 1
As would be predicted, scats collected from the den site in
1978 and 1979 suggested that moose, particularly calves, were
the most important prey item during summer (Tables 36 and 37).
Caribou were also an important food i tern. Both beaver and
snowshoe hare also appeared to be important prey items
accounting for 25 percent of identified food remains in 1978 and
10 percent in 1979.
Tyone Pack -From November 1977 through June 1980, members
of· the Tyone pack were observed on 33 kills. Adult and calf
moose were the most important prey i terns . The observed kills
included 14 adult moose, 12 calf moose, 4 of unidentified
species, 2 adult caribou and 1 unclassified moose. Because most
of our observations were made during winter months when caribou
were not available, our aerial observations probably do not
accurately reflect the importance of caribou during summer
months. Since a majority of the Nelchina caribou herd calves in
the Kosina Creek drainages, we suspect that large numbers of
caribou b-ecome available in mid to late summer following
calving.
Moose were available to this pack on a year-round basis,
although densities were greater in winter than summer due to the
presence of migratory moose from the Maclaren and upper susitna
Rivers (Ballard and Taylor 1980). We suspected that perhaps in
response to winter snow conditions, more moose and caribou were
available to this pack during winter 1979-80 than during winter
1978-79. Moose densities and pack predation rates will be
discussed further in the Predator Rate section of this report.
Although this pack's den site was located in 1979, we were
unable to examine it and thus no scat data are available. We
suspect that calf moose and caribou comprised the bulk of the
diet with small mammals being of secondary importance.
FOOD HABITS SUMMARY
Aerial Observations
Table 38 summarizes the kills at which radio-collared
wolves were observed by month of observation from April 1975
through June 1980. Radio-collared wolves were observed at 360
kills, 38 ( 10.6%) of which were either occupied or had been
previously visited by one, or more, brown bear. Adult moose
(n = 144) were the most common prey item recorded, comprising 40
percent of the total number of kills. However, heavily consumed
yearling moose may have been classified as adults and therefore
yearlings are probably underrepresented in the kill data. For
purposes of this report we define calves as those individuals
born in May or June (assumed birthdate of 1 June) and surviving
to 30 May of the following calendar year. In some cases we
refer to short yearlings which we define as calves 6 through 12
months of age. Long yearlings are defined as those animals from
13 to 24 months of age. . Thereafter they are referred to as
adults.
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Table 36. Incidence of food remains in wolf scats collected at the
Tolsona wolf den occupied during late spring and swnmer
1978 in GMU 13 of southcentral Alaska.
Items in 36 Items in 89 Items in 125
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 0 0 0 0 0 0
Calf moose 21 50 48 45.7 69 46.9
Adult caribou 0 0 0 0 0 0
Calf caribou 11 26.2 6 5.7 17 11.6
Beaver 7 16.7 24 22.9 31 21.1
Muskrat 0 0 4 3.8 4 2.7
Snowshoe hare 1 2.4 5 4.8 6 4.1
Parka squirrel 0 0 0 0 0 0
Micro tine 0 0 6 5.7 6 4.1
Bird 0 0 0 0 0 0
Fish 0 0 0 0 0 (\ v
Vegetation 1 2.4 6 5.7 7 4.8
Unidentifiable 1 2.4 3 2.9 4 2.7
Other 0 0 3 2.9 3 2.0
Total 42 100.1 105 100.1 147 100.0
Grouned Data for 125 Scats
Food item Number of items Percent occurrence
Ungulate 86 58.5
Small mammals 47 32.0
Other 14 9.5
Total 147 100.0
8 3
Table 37. Summary of incidence of food remains in wolf scats collected
at the Tolsona wolf den occupied during late spring and
early summer 1979 in GMU 13 of southcentral Alaska.
Items in 111 Items in 37
adult scats EUE scats
Food item No. % occ. No. % occ.
Adult moose 9 6.2 2 4.9
Calf moose 54 37.2 26 63.4
Adult caribou 12 8.3 0 0
Calf caribou 29 20.0 6 14.6
Beaver 2 1.4 0 0
Muskrat 4 2.8 0 0
Snowshoe hare 16 11.0 1 2.4
Parka squirrel 0 0 0 0
MicroHne 3 2.1 1 2.4
Other-8 5.5 4 9.8
Vegetation 5 3.4 1 2.4
Unidentifiable 3 2.1 0 0
Total 145 100.0 41 99.9
Grou:eed Data for 148 Scats
Food item Number of items Percent occurrence
Ungulate 138 74.2
Small mammals 27 14.5
Other .... ~ .:..L 11.3
Total 186 100.0
Other includes birds, insects, wolf, red squirrel,
unidentified ungulates.
Items in 148
combined scats
No. % occ.
11 5.9
80 43.0
12 6.5
35 18.8
2 1.1
4 2.2
17 9. 1
0 0
4 2.2
12 6.5
6 3.2
3 1.6
186 100.1
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Table 38.
Month of
observation
May
June
July
August
September
October
November
Decemher
January
February
March
April
Total
Percent
00
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Summary by month of prey species discovered while tracking radio-collared wolf packs
in GMU 13 of southcentral Alaska from April 1975 through June 1980.
Calf moose Yearling moose
Est. Est. Unclass-Unclass-
age age Adult ified Calf Adult ified
n (mos.) n (mos.) moose moose caribou caribou caribou Beaver Misc. prey Total
4 0 2 12 8 5 1 2 1 1 2 25
6 1 6 13 15 1 1 1 5 35
2 2 4 14 13 2 1 2 1 25
3 3 15 8 1 5 1 2 20
1 4 16 9 3 13
1 5 2 17 11 5 2 1 22
4 6 18 8 5 6 3 5 31
3 7 19 11 3 2 2 1 1 23
10 8 20 18 13 41
10 9 21 16 4 3 1 4 38
13 10 22 14 4 6 2 2 41
16 11 1 23 13 2 1 8 4 45 -
73 15 144 26 6 48 16 7 25 360
20.3 4.2 40.0 7.2 1.7 13.3 4.4 1.9 6.9 99.9
Only 5.8 percent of the 360 kills at which radio-collared
wolves were observed were calf moose. Short yearling moose,
however, comprised 14.2 percent of the kills. Moose of all ages
comprised 71.7 percent of the kills. Other prey items, in order
of their importance, were adult caribou ( 13 . 3%) , miscellaneous
and unidentified species (6.9%), unclassified caribou (4.4%),
beaver (1.9%) and calf caribou (1.7%). Obviously these data
underrepresent the importance of small mammals and birds because
of th~ difficulty involved in observing these small carcasses
from fixed-wing aircraft. They probably also underrepresent the
importance of calf caribou to one or two wolf packs which
overlap the Kosina Creek caribou calving grounds because these
packs were not intensively studied during the calving season.
Also, observability of kills varied seasonally with kills being
least observable during summer and most observable during
winter.
Wolf kill data obtained from aerial observation were
subjected to an analysis of residuals (Everith 1977) to
determine which cells of a Chi-square table were significant in
rejection of the null hypothesis. Kill data were compared
between numbers of adult and calf moose and caribou by month of
observation from . April 1975 through June 1980. Significant
(P<O. 05) deviations were observed for short yearling moose
during April, long yearling moose during June and July, adult
moose in January, and adult caribou in December and January.
Wolves appeared to select adult moose during most months of the
year except during January through July, when short and long
yearling moose appeared to comprise a disproportionate
percentage of the kill. These particular moose cohorts
comprised 12 to 19 percent of the moose counted during November
composition counts for 1975 through 1979 when they were 6 months
old. However, these cohorts comprised 42 percent (a much larger
percent if potential bear kills were excluded) of the moose
killed by wolves during this time period when 7 to 14 months of
age. We conclude fr0m this analysis that wolves were killing
short and long yearling moose from January through July
disproportionately to their presence in the moose population.
Further, we conclude that, overall, moose of all age classes
were the most important prey for wolves in the study areas.
The data presented in Table 38 do not support the
hypothesis that wolves were actively selecting newborn moose
calves in the Nelchina Basin. Intensive studies of selected
wolf packs during late and early summer support this conclusion
(Appendix I I I) . However, the results of food habit studies
based upon percent occurrence of food items in scat collections
at den and rendezvous sites suggest that calf moose were more
important.
Scat Analyses--Summer Food Habits
Scat analyses from den and rendezvous sites were pooled for
each year of study and are presented in Tables 39 through 42.
8 6
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Table 39. Summary of incidence of food remains in wolf scats collected
at dens and rendezvous sites used during late spring and
summer 1975 in GMU 13 of southcentral Alaska.
Items in 403 Items in 1,129 Items in 1,532
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 92 18.4 169 13.1 261 14.5
Calf moose 132 26.3 472 36.5 604 33.6
Caribou 22 4.4 58 4.5 80 4.5
Dall sheep 2 .4 6 .5 8 .4
Beaver 50 10.0 187 14.5 237 13.2
Snowshoe hare 158 31.5 281 21.7 439 24.5
Ground squirrel 8 1.6 16 1.2 24 1.3
Red squirrel 2 .4 0 0 2 .1
Muskrat 5 1.0 7 .5 12 . 7
Mi~roHne 5 1.0 26 2.0 31 1.7
20 4.0 "" 2.4 52 2.9 Otner-.)~
Vegetation 3 .6 29 2.2 32 1.8
Unidentifiable 2 .4 11 .9 13 . 7
Total 501 100.0 1,294 100.0 1,795 99.9
Grouped Data for 1,532 Scats
Food item Number of items Percent occurrence
Ungulate 953 53.1
Sma n 11amma l s 745 41.5
Other-97 5.4
Total 1,795 100.0
]j Includes birds, invertebrates, unidentified, wolf,
fish, vegetation.
8 7
Table 40. Summary of incidence of food remains in wolf scats collected
at wolf den and rendezvous sites used during late spring and
summer 1976 in GMU 13 of southcentral Alaska.
Items in 257 Items in 384 Items in 641
adult scats EUE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 27 8.4 23 5.4 50 6.7
Calf moose 142 44.2 250 59.1 392 52.7
Adult caribou 16 5.0 14 3.3 30 4.0
Calf caribou 24 7.5 30 7.1 54 7.3
Beaver 26 8.1 10 2.4 36 4.8
Muskrat 2 .6 16 3.8 18 2.4
Snowshoe hare 48 15.0 10 2.4 58 7.8
Parka squirrel 2 .6 2 .5 4 .5
Microune 10 3.1 7 1.7 17 2.3
Other-7 2.2 8 1.9 16 2.2
Vegetation (\ 2.8 24 5.7 33 4.4 :7
Unidentifiable 8 2.5 28 6.6 36 4.8
Total 321 100.0 423 100.1 744 99.9
GrouEed Data for 641 Scats
Food item Number of items Percent occurrence
Ungulate 526 70.7
Small mammals 133 17.9
Other 85 11.4
Total 744 100.0
±' _, Other includes birds, insects, eggshells, wolf,
red squirrel, unidentified ungulates.
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Table 41. Summary of incidence of food remains in wolf scats collected
at wolf den and rendezvous sites used during late spring and
summer 1977 in GMU 13 of southcent.ral Alaska.
Items in 214 Items in 386 Items in 600
adult scats I!UE scats combined scats
Food item No. % occ. No. % occ. No. % occ.
Adult moose 18 6.9 14 3.0 32 4.4
Calf moose 150 57.3 267 57.5 417 57.4
Adult caribou 5 1.9 2 .4 7 1.0
Calf caribou 14 5.3 20 4.3 34 4.7
Beaver 11 4.2 27 5.8 38 5.2
Muskrat 7 2.7 20 4:3 27 3.7
Snowshoe hare 14 5.3 21 4.5 35 4.8
Parka squirrel 1 .4 1 .2 2 .3
MicroHne 11 4.2 15 3.2 26 3.6
Other-15 5.7 26 5.6 41 5.6
Vegetation 9 3.4 37 8.0 46 6.3
Unidentifiable 7 2.7 14 3.0 21 2.9
Total 262 100.0 464 99.8 726 99.9
Grou12ed Data for 600 Scats
Food item Number of items Percent occurrence
Ungulate 490 67.5
Small mammals 128 17.6
Other 108 14.9
Total 726 100.0
y Other includes birds, insects, wolf, red squirrel,
unidentified ungulates.
8 9
Table 42. Summary of incidence of food remains in wolf scats collected
at the den and rendezvous sites occupied during late spring
and early summer 1978 in GMU 13 of southcentral Alaska.
Items in 308 Items in 395 Items in 703
adult scats pup scats combined scats
% Food item No. % occ. No. % occ. No. 0 occ.
Adult moose 34 9.0 8 1.7 42 5.0
Calf moose 170 44.9 224 48.7 394 47.0
Adult caribou 12 3.2 3 0.7 15 1.8
Calf caribou 15 4.0 19 4.1 34 4.1
Beaver 61 16.1 57 12.4 118 14.1
Muskrat 15 4.0 27 5.9 42 5.0
Snowshoe hare 35 9.2 24 5.2 59 7.0
Parka squirrel 0 0 1 0.2 1 0.1
MicroHne 4 1.1 12 2.6 16 1.9
8 2.1 30 6.5 38 4.5 Other-
Vegetation 15 4.0 22 4.8 37 4.4
Unidentifiable 10 2.6 33 7.2 43 5.1
Total 379 100.2 460 100.0 839 100.0
GrouEed Data for 703 Scats
Food item Number of items Percent occurrence
Ungulate 485 57.8
Small mammals 236 28.1
Other 118 14.1
Total 839 100.0
}j Other includes birds, insects, wolf, red squirrel,
unidentified ungulates.
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Although no statistical tests were performed it appeared that
there were few, if any, differences in the proportions of food
i terns in adult versus pup scats. This suggests that adult
wolves were feeding pups the same items they were feeding upon.
Therefore, there appears to be no justification for analyzing
pup and adult scats separately.
Theberge et al. (1978) determined that the contents of wolf
scats collected at rendezvous sites in Alonquin Park, Ontario
differed from those collected elsewhere in the par}!;. In that
study beaver was a common food item at rendezvous sites
suggesting that local resources were being exploited. In areas
away from rendezvous sites white-tailed deer (Odocoileus
virginianus) remains were more common. In this study moose
appeared to be the most common food i tern at both den and
rendezvous sites as well as being the most important prey during
winter. However, three packs (Delta, Ewan, and St. Anne) did
not follow this pattern. In all cases, the ungulate densities
in these areas were quite low indicating that dependence on
small mammals was of necessity rather than selection.
Interestingly, these three packs exhibited more nutritional
deficiencies · than in any of the other pack areas and also
appeared to fluctuate in numbers more drastically, remaining at
low numbers for longer periods of time. This indicates that
areas with low ungulate density also have a low carrying
capacity for wolves regardless of the abundance of small mammals
and birds. However, our study did not encompass a period when
snowshoe hare numbers were at peak levels over large areas. It
is possible that high densities of small mammals under some
circumstances could be an important food supplement.
There were differences in the percent occurrence of food
items in scats collected at den sites versus rendezvous sites.
Three packs had an increasing reliance on calf moose following
movement to the rendezvous site. These increases were as
follows: Hogan Hill in 1978--56 to 81 percent; Keg Creek in
1975--58 to 73 percent; Mendeltna in 1976--28 to 55 percent and
again in 1977--43 to 62 percent; Sinona in 1976--49 to 78
percent. These data may reflect wolf movement into areas of
higher ungulate abundance.
Of possible significance, were the differences in food
habits from den to rendezvous sites for the Delta and Ewan
packs. In 1975 the Delta pack had a decline in use of both calf
moose (24 to 10%) and snowshoe hare (41 to 5%) following their
movement from den to rendezvous site, but an increase in both
beaver (1 5 to 49%) and squirrel ( 2 to 23%) . In the same year,
the Ewan pack exhibited a decline in use of calf moose (21 to
10%) and snowshoe hare (31 to 4%), while beaver remained
constant at 33 percent, and the percent of adult moose rose from
7 to 41 percent. These changes in food habits could reflect
wolf movement into areas of different prey abundance as well as
seasonal differences in availability and/or vulnerability of
prey.
9 ~
A combined summary of food items identified in wolf scats
by year of study for all packs and sites is presented in
Table 43. During the 5 years of study, 4,290 food items were
identified in 3, 624 adult and pup wolf scats. The combined
results were, in general, similar to most of the analyses for
individual packs in that calf moose was the predominant food
i tern comprising 44 percent of the total i terns. Adult moose,
however, ranked only fourth, comprising 9.2 percent. Snowshoe
hare and beaver ranked second (14.2%) and third (10.0%),
respectively. Adult and calf caribou combined accounted for 7
percent of the total food items. This latter percentage was not
surprising, however, because caribou were largely absent from
many of the wolf pack territories during late spring and summer.
Perhaps coincidentally, the lowest percent occurrence of
calf moose was in 1975 when calf: 100 cow ratios were at their
lowest level ever recorded in the Basin. To determine if there
was a relationship between calf moose abundance and occurrence
in wolf s~ats,, .we c9mpared annual fall moose count data for the
Basin with EreqUe:ilcy · of occurrence (arcsine square root
transformation) of calf moose in wolf scats collected from 1975
through 1979 (Tab~e 43). If wolf predation were a significant
factor limiting calf moose survival, we would anticipate an
inverse relationship between percent calf moose occurrence and
subsequent fall indices of calf abundance. Instead, there
appeared to be a positive but nonsignificant relationship
(P>0.05, D.F. = 3) between calf moose occur2ence in scats and
moose ~undance (calves observed I2er hr, r = .46; calf % of
herd, r-= .54; calves/100 cows, r = .57). This relationship
was investigated further by pooling den and rendezvous site
collections for individual packs each year and then comparing
percent calf moose occurrence in scats (Table 44) with calf
abundance indices obtained in specific moose count areas where
each pack resided. Significant (P<0.05) curvilinear
relationships existed for percent cal~ ~9Pse occurrence versus
calves observed per hour (y = 2.02X · , r = .57), percent
calves in herd (y = 1 4~7 13 + 14.49 ln (X), r = .65) and calf:100
cow ratios (y = .OlX · , r = .68). These analyses suggest that
the occurrence of calf moose in scats was related to the number
of calves available. Therefore, the number of moose calves
· eaten, did not determine subsequent calf moose numbers observed
in fall. Further, this analysis suggests that wolves were
taking calf moose in proportion to their annual abundance rather
than continuing to $elect calves when they were not abundant.
To determine the degree of variability in occurrence of
-food i terns in scats for individual packs between years, den and
rendezvous site scat data were pooled by pack per year ,13.nd
compared for packs for which more than 1 year of data were
available (Table 44). Eventually percentages will be
transformed to arcsine square-root percentages and analyzed with
analysis of variance for comparison with the analyses presented
by Voigt et al. ( 1976). Our simple comparison suggests that
there was considerable variation in the occurrence of food items
9 2
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Table 43. Summary by year of incidence of food remains in wolf scats collected at wolf den and rendezvous sites from 1975 through 1979 in
Game Management Unit 13 of southcentral Alaska.
1975 1976 1977 1978 1979 Total
1,532 scats 1>41 scats bOO scats 703 scats 148 scats 3,624 scats
Number Percent Number Percent Number Percent Number Percent Number Percent Number Percent
O . ......:..J
Food items of items occurrence of items occurrence of items occurrence of items ·occurrence of items occurrence of items occurrence
Adult moose 261 14.5 50 6.7 32 4.4 42 5.0 11 5.9 396 9.2
Calf moose 604 33.6 392 52.7 417 57.4 394 47.0 80 43.0 1,887 44.0
Adult caribou -~o.?.l 30 4.0 7 l.O 15 1.8 12 6.5 64 1.5
Calf caribou 4.5 54 7.3 34 4.7 34 4 .I 35 18.8 237 5.5
Beaver 237 13.2 36 4.8 38 5.2 118 14.1 2 1.1 431 10.0
Muskrat 12 0.7 18 2.4 27 3.7 42 5.0 4 2.2 103 2.4
Snowshoe hare 439 24.5 58 7.8 35 4.8 59 7.0 17 9.1 608 14.2
Parka squirrel 24 1.3 4 .5 2 0.3 l 0.1 0 0 31 0.7
MicroHne 31 1.7 17 2.3 26 3.6 16 1.9 4 2.2 94 2.2
Other-62 3.5 16 2.2 41 5.6 38 4.5 12 6.5 169 3.9
Vegetation 32 1.8 33 4.4 46 6.3 37 4.4 6 3.2 154 3.6
Unidentified 13 0.7 36 4.8 21 2.9 43 5.1 3 1.6 116 2.7
Totals 1,795 100.0 744 99.9 726 99.9 839 100.0 186 100.1 4,290 99.9
!I Other includes birds, insects, egg shells, wolf, red squirrel, unidentified ungulate.
.?.I Adult and calf caribou combined.
Table 44. Comparison between years by individual wolf pack of incidence of food remains in wolf scats collected at
dens and rendezvous sites used from 1975 through 1979 in Game Management Unit 13 of southcentral Alaska.
Hogan Hill Keg Creek Mendeltna
1975 1978 1975 1976 1978 1976 1977
135 scats 244 scats 208 scats 81 scats 45 scats 139 scats 399 scats
% of % % of % % of % % of % %of % % of % % of %
Food items items occ. items occ. items occ. items occ. items occ. items occ. items occ.
Adult moose 20 12.6 30 10.3 32 14.2 5 5.9 1 2.0 12 8.0 21 4.3
Calf moose 46 28.9 175 60.1 151 66.8 62 72.9 26 53.1 67 44.7 287 58.8
Adult caribou ;;y 12 4.1 -~~/ 6 7.1 1 2.0 19 12.7 7 1.4
Calf caribou 13.8 3 1.0 0.4 5 5.9 9 18.4 36 24.0 34 7.0
Beaver 27 9.3 8 3.5 2 2.4 1 2.0 I 0.7 3 0.6
Muskrat 5 1.7 0 0 2 1.3 14 2.9
Snowshoe hare 64 40.3 13 4.5 27 11.9 0 0 3 2.0 20 4.1
Parka squirrel 0 0 0 0 0 0 2 0.4
MicroHne 3 1.0 1 2.0 1 0.7 22 4.5
Other-3 1.9 3 1.0 4 1.8 1 2.0 2 1.3 32 6.6
Vegetation 4 2.5 11 3.8 2 0.9 2 2.4 0 0 4 2.7 34 7.0
Unidentified 9 3.1 1 0.4 3 3.5 9 18.4 3 2.0 12 2.5
Totals 159 100.0 291 99.9 226 99.9 85 100.1 49 99.9 150 100.1 488 100.1
1/ Other includes birds, insects, egg shells, wolf, red squirrel, unidentified ungulates. ~I Adult and calf caribou combined.
. ... . ~
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Table 44 (cont.). Comparison between years by individual wolf pack of incidence of food remains in wolf scats collected at dens and rendezvous sites used from
1975 through 1979 in Game Management Unit 13 of southcentral Alaska.
Sinona St. Anne's Tolsona
1975 I97o 1977 1978 1975 197o 1977 1978 1978 1979
128 scats 212 scats 152 scats 159 scats 232 scats 17o scats 49 scats 130 .scats 125 scats 148 scats;
%of % %of % % of % % of % % of % %of % % of % %of % % of % % of %
Food items items occ. items occ. items occ. items occ. items occ. items occ. items occ. items occ. items occ. items occ
·Adult moose 39 28.3 20 8.2 9 4.9 8 4.3 24 7.3 12 5.2 2 3.7 3 1.8 0 0 11 5.9
Calf moose 77 55.8 151 62.1 105 57.4 87 46.3 61 18.5 82 35.5 25 46.3 37 22.6 69 46.9 80 43.0
Adult caribou -iY 0 0 0 0 1 0.5 0 0 5 2.2 0 0 1 0.6 0 0 12 6.5
Calf caribou 0.7 1 0.4 0 0 5 2.7 0 0 12 5.2 0 0 0 0 17 11.6 35 18.8
Beaver 13 9.4 16 6.6 23 12.6 24 12.8 9 2.7 17 7.4 12 22.2 35 21.3 31 21.1 2 1.1
Muskrat 9 3.7 11 6.0 16 8.5 10 3.0 7 3.0 2 3.7 17 10.4 4 2.7 4 2.2
Snowshoe hare 2 1.4 6 2.5 7 3.8 10 5.3 195 59.1 49 21.2 8 14.8 30 18.3 6 4.1 17 9.1
Parka squirrel 2 0.8 0 0 1 0.5 0 0 2 0.9 0 0 0 0 0 0 0 0
MicroHne 1 0.7 3 1.2 2 1.1 1 0.5 16 4.8 12 5.2 2 3.7 5 3.0 6 4.1 4 2.2
Other-3 2.2 2 0.8 6 3.3 10 5.4 12 3.6 12 5.2 2 3.7 21 12.8 3 2.0 12 6.5
Vegetation 1 0.7 17 7.0 12 6.6 12 6.4 3 0.9 7 3.0 0 0 7 4.3 7 4.8 6 3.2
Unidentified 1 0.7 16 6.6 8 4.4 13 6.9 0 0 14 6.1 1 1.9 8 4.9 4 2.7 3 1.6
Totals 138 99.9 243 99.9 183 100.1 188 100.1 330 99.9 231 100.1 54 100.0 164 100.0 147 100.0 186 100.1
1/ Other includes birds, insects, egg shells, wolf, red squirrel, unidentified ungulates. ~I
<D
U1
Adult and calf caribou combined.
for individual packs between years. This appeared to be
particularly true for adult and calf moose, adult and calf
caribou, beaver and snowshoe hare. Whether these differences
were due to shifts in prey distribution or abundance is un~~o·N.n,
but it appears likely.
Floyd et al. ( 1978) discussed some of the problems
associated with using frequency of occurrence of food i terns as
indicators of wolf food habits. Mech (1970) suspected, and
Floyd et al. (1978) confirmed, that percent occurrence of small
animals was overrepresented in relation to large animals because
small animals have a relatively higher surface-volume ratio and
are covered with relatively more hair (most predominant
identifiers in wolf scats) than large animals. To correct for
this discrepancy they determined the relationship between
collectable scats and weight of prey consumed for several animal
species. The relationship between these two variables allows
calculation of kilograms of prey consumed and number of prey
eaten as represented by the "collectable 11 scats.
Data in Table 43 were analyzed using the method derived by
Floyd et al. (1978) to provide kilograms of prey consumed and
number of individuals consumed per prey species per year
(Table 45). To simplify calculations we used number of prey
items rather than number of scats. Although this method is less
accurate, it purportedly does not produce too great an error
(Floyd et al. 1978). Conversion of the scat data to numbers of
kilograms consumed indicated that, as Floyd et al. (1978)
suggested, small animals were overrepresented when percent
occurrence was used. The importance of calf moose, however,
changed very little while L~e importance of adult moose
increased substantially.
Mech (cited in Peterson 1977) determined that one Minnesota
wolf pack declined after a winter when only 3 . 0 to
3.4 kg/wolf/day of food was available, increased at
5.8 kg/wolf/day, and remained stable at 3.6 kg/wolf/day. Based
upon this Minnesota study, we estimated the number of days that
adult wolves from the den and rendezvous sites could be
sustained based upon collected scats at the above mentioned
consumption rates. We assumed that the packs represented by the
scat collections were capable of increasing or at least
remaining stable based upon available prey. The n~ber of days
that wolves could be sustained based on th.e scats collected
ranged from 46 days, at a consumption rate of 5.8 kg/wolf/day in
1979, to 956 days in 1975, based upon a consumption rate of
3. 6 kg/wolf/day. In an effort to determine approximately how
many individual days per adult wolf were represented, we divided
the total days at each consumption rate per year by ~~e number
of adult wolves thought to be present. This analysis indicated
that the scats represented from 6 to 17 days of consumption.
These figures may be inflated, however, because some wolves are
more sedentary than others, particularly mothers of pups {Murie
1944; Theberge et al. 1978; Foster and Ballard unpub. data) and,
therefore, scat collections from these sites would be weighted
in favor of these sedentary individuals.
9 6
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Table 45. Summary of estimated numbers of kilograms and individual prey consumed by wolves as determined from wolf scats collected at den and rendezvous
____ sites in Game Management Unit 13 of southcentral Alaska from 1975 through 1979.
Prey
Adult moose
Calf moose
Adult caribou
Calf caribou
Beaver
Muskrat
Assumed
weight
(kg)
of prey
(X)
Snowshoe hare
Squirrel
Microtine rodent
71 # of packs it
Kgs/scat-# of adult wolves-
(y) 11 of pups =
8.93
1.16
3.28
.62
.63
.41
.42
.39
.38
Number of scat food items
1975
10
55
37
1976 -5
31
29
261 50
604 392
-30
so'!../ 54
237 36
12 18
439 58
24 4
31 17
1977
-3
19
22
32
417
7
34
38
27
35
2
26
1978
-5
25
27
42
394
15
34
118
42
59
1
16
1979 -1-
11
80
12
35
.2
4
17
0
4
7
9
1975
10
55
37
2330.73
700.64
49.6
149.31
4.92
184.38
9.36
11.78
1976 -5
31
29'
kgs
446.50
454.72
98.40
33.48
22.68
7.38
24.36
1.56
6.46
consumed
1977
-3
19
22
285.76
483.72
22.96
21.08
23.94
11.07
14.70
0. 78
9.88
1978 -5-
25
27
375.06
457.04
49.20
21.88
74.34
17.22
24.78
0.39
6.08
1979 -1-
7
9
98.23
92.80
39.36
21.70
1.26
1.64
7.14
0.00
1.52
Number of individuals/
prey s~ecies represented
1975 197 1977 1978 1979 10 -5---3- --5---1-
55 31 19 25 7
37 29 22 27 9
5.5 1.0 0.7
18.0 11.7 12.4
0.7 0.2
4.1 2.79 1.8
11.9 1.8 1.9
3.5 5.3 7.9
102.4 13.5 8.2
18.7 3.1 1.6
117.8 ~ 98.8
0.9
11.7
0.3
1.8
5.9
12.3
13.8
0.8
60.8
0.2
2.4
0.3
1.8
0.1
1.2
4.0
0.0
15.2
Totals 1688 659 618 721 165 3440.72 1095.54 873.89 1025.19 263.65 281.9 104.49 133.5 108.2 25.2
Total number days represented based upon 5.8 kg available food/day/adult wolf for
population to increase (Mech cited in Peterson 1977)
Number days/wolf represented by consumption rate of 5.8 kg food/day/adult wolf
Total number days represented based upon 3.6 kg available food/day/adult wolf for
popul~tion to remain stable (from Mech cited in Peterson 1977).
Number days/wolf represented by consumption rate of 3.6 kg food/day/adult wolf
593.2 188.9
10.8 6.1
955.8 304.3
17.4 9.8
150.1 176.8 45.5
7.9 7.1 6.5
242.7 284.8 73.2
12.8 11.4 10.5
!I From Franzmann and Bailey 1977. ~/ Weight of 1 month old moose calf computed as follows: newborn weight equals 18.0 kg (from Franzmann et al. 1980) with assumed weight gain of 1.0 kg/day
31 for 21 days equals 39.0 kg.
T. From Skoog 1968. ~/ From Floyd et al. 1978. 51 From Burt and Grossenheider 1952. ~/ From Stephenson 1978. 71 From Floyd et al. 1978 calculated by equation y = 0.38+0.02(x). =-~~ Number in some packs in 1975 and 1976 were estimated.
Calf and adult caribou combined.
We used percent kilograms of each prey species consumed as
indicated by scat analyses to extrapolate the possible impact of
wolf predation on individual prey species in GMU 13. This •,vas
done by calculating two estimates for spring GMU 13 wolf
population estimates from 1975 through 1979 based upon
consumption rates of 3. 6 and 5. 8 kg of food/wolf/day (Tables 46
and 47). These calculations indicated that at a consumption
rate of 5. 8 kgjwolf/day from mid-May through mid-July for 1975
through 1979, GMU 13 wolves were annually consuming the
following numbers of ungulates: 42 to 132 adult or yearling
moose, 434 to 1, 013 calf moose, 11 to 59 adult caribou and from
75 to 329 calf caribou. These estimates were considerably lower
when the consumption rate of 3.6 kg/wolf/day was used: 23 to 82
adult moose, 270 to 515 calf moose, 7 to 37 adult caribou, and
73 to 204 calf caribou. The implications of these estimates to
GMU 13 moose populations will be presented in the section of
this report entitled Evaluation of Wolf Predation on Moose.
Age and Condition of Prey
When possible, we examined kills to aid in determining the
age, sex and physical condition of prey taken by wolves. During
this study we examined 125 moose (Table 48) and 25 caribou
(Table 49) which had died of various causes. Portions of the
caribou data have been prepared for publication in an article
entitled "Surplus Killing of Caribou by Wolves"· authored by
Sterling Eide and Warren Ballard. A draft copy of the
manuscript submitted to the Canadian Field Naturalist is
included as Appendix IV. Cause of death of kills exami:ned on
the ground were placed into one of five categories: · ( 1) \volf
predation, ( 2) accidental which included road kills, tagging
mortalities, nuisance kills, potlatch kills (moose killed for
religious purposes) and unknown causes, ( 3) winter kill
(starvation), { 4) bear (both brown and black) predation, and
(5) unknown species predation.
In an effort to obtain an index of the physical conditon of
moose and caribou at the time of death we collected long bones
and mandibles (ramuses) at each kill to determine percent fat in
bone marrow (Neiland 1970). Mandibles were collected in
addition to long bones because at heavily consu.llled carcasses
wolves had frequently chewed the ends of long bones and licked
out the bone marrow and/or carried these bones away from the
site. At these kills the only remaining items •,.yere visceral
material, hair, and mandibles. Mandibles contain small volumes
of bone marrow and, if found useful for determining percent fat,
c9uld greatly increase the sample of condition data on
wolf-killed moose. Therefore, we began comparing percent fat of
bone marrow between long bones and mandibles to determine what,
if any, correlations existed. A preliminary analysis suggests a
good correlation between percent fat values obtained from the
two bones for adult moose and caribou but not for calf and short
yearling moose and caribou. These data will be fur~"ler analyzed
in a future publication. . The condition data exarnined in t..1-).is
report . only pertain to percent fat values obtained from long
bones (femurs, metatarsals).
9 8
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Table 46. Estimated nurrilier of kilograms of prey consumed by Game Management Unit 13 wolves for a 61 day period ranging from
approximately mid-May through mid-July 1975 through 1979 as extrapolated from scat analyses and wolf population
estimates.
Percent kg Estimated kgs prey Estimated kgs prey
of prey consumed by wolves consumed by GMU 13 wolves at consumed by GMU 13 wolves at
according to scat analysis consumption rate of 5.8 kg/day{/ consumption rate of 3.6 kg/dav_f
(derived from Table 44) wolf from aEErox. 5/1 -6/30-wolf from aEErox. 5/1 -6/30-
Prey 1975 1976 1977 1978 1979 1975 1976 1977 ·1978 1979 1975 1976 1977 1978 1979
Adult moose 67.7 40.2 32.7 36.6 37.3 56288 38259 19089 15669 17948 34937 23747 11849 9725 11140
Calf moose 20.4 41.5 55.4 44.6 35.2 16961 391.96 32340 19093 16937 10528 24515 20074 11851 10513
Adult caribou ~~4?:_/ 9.0 2.6 4.8 14.9 --8566 1518 2055 7169 --5316 942 1275 4450
Calf caribou 3.1 2.4 2.1 8.2 1164?:_/ 2950 1401 899 3946 122?:_1 1831 870 558 2449
Beaver 4.3 2.1 2.7 7.3 .5 3575 1999 1576 3125 241 2219 1241 978 1940 149
Muskrat 0.1 .7 1.3 1.7 .6 83 666 759 728 289 52 414 471 452 179
Snowshoe hare 5.4 2.2 1.7 2.4 2.7 4490 2094 992 1027 1299 2787 1300 616 638 806
Squirrel 0.3 .1 0.1 0.03 249 95 58 13 155 59 36 8
Micro tine rodents 0,3 .6 1.1 .6 .6 249 571 642 257 289 155 354 399 159 179
Totals 99.9 99.5 100.0 100.1 100.0 83059 94696 58375 42866 48118 51555 58777 36235 26606 29865
!I Estimated spring GMU 13 wolf population was as follows (see Table 59): 1975 -235, 1976-269, 1977 -165, 1978-121,
1979 -136.
?:_I Adult and calf caribou combined.
Table 47. Estimated number of prey individuals consumed by Game Management 13 wolves for a 61 day period ranging from
approximately mid-May through mid-July 1975 through 1979 as extrapolated from wolf scat analysis and wolf
population estimates.
Prey
Assumed weight
(kg) of prey
(from Table 44)
Adult moose
Calf moose
Adult caribou
Calf caribou
Beaver
Muskrat
Snowshoe·hare
Squirrel
Microtine rodents
Totals
427.5
39.0
145.0
12.0
12.5
1.4
1.8
.5
. 1
Estimated number
of prey consumed by GMU 13 wolves based 1 upon consumption rate of 5.8 kg/wolf/day-/
1975 1976 1977 1978 1979 Subtotal
132
435
-;7y
286
59
2494
498
2490
90
1013
59
246
160
476
1163
190
5710
45
829
11
117
126
542
551
116
6420
37
490
14
75
250
520
571
26
2570
42
434
49
329
19
206
722
2890
346
3201
133
864
841
1803
5501
830
20080
6491 9107 8757 4553 4691 33599
Estimated number
of prey consumed by •GMU 13 wolves based 1 upon consumption rate of 3.6 kg/wolf/day-/
1975 1976 1977 1978 1979 Subtotal
82
270
6~?:/
178
37
1548
310
1550
56
629
37
153
99
296
722
118
3540
4035 5650
28
515
7
73
78
336
342
72
3990
23
304
9
47
155
323
354
16
1590
26
270
31
204
12
128
448
1790
215
1988
84
537
522
1120
3414
516
12460
5441 3221 2909 20856
--~~ Estimated spring GMU 13 wolf population was as follows: 1975 -235, 1976 -269, 1977 -165, 1978 -121, 1979 -136.
Adult and calf caribou combined.
0
C>
r "" 1 1'7"' ..... .
..
r·'!1 r 'l· r r ll r ''l r l j I L.J i ] r· [_J LLL:J . C......:..J
L:..:J t:::..:....J [__J C'' '''!i c__J r ······ 1
'• ~
Table 48. Age, sex, condition as determined by percent fat and cause of mortality of moose kills examined from
June 1975 through June 1980 in GMU 13 of southcentral· Alaska.
Date of Approximate Percent fat Marrow
Accession II Age Sex collection location longbone Ramus color Cause of death
120000A 11 ~ 06/06/75 Hogan Hill 56 Wolf predation
120000B Calf 11/24/75 Sinona Lodge Wolf predation
120000C 4 ~ 05/05/75 Lake Louise 26 Wolf predation
120000D Yearling cJ 06/21/76 Spring Creek 60 Wolf predation
120001 8 ~ 01/03/77 Lower Sinona Ck. Wolf predation
120002 17 ~ 01/03/77 Lower Sinona Ck. 78.0 Light pink with Wolf predation
red spots
120003 15 ~ 12/09/76 Grayling Lake 45.2 Wolf predation
120004 18 ~ 12/ ? /76 Tonsina River 91.3 Wolf predation
120005 . 9 ~ 09/25/76 Mendeltna Creek 78.0 White Unknown predation
120006 Adult ~ 12/27/76 Gakona River 46.4 White with pink Wolf predation
spots
120007 Yearling ~ 10/29/76 Grayling Lake 43.9 Outer -red Wolf predation
Inner -white
120008 15 ~ 01/ ? /77 Copper River 75.6 Light pink Winter kill? (legs
tucked under body)
120008B Calf ~ 01/25/77 Marsh Lake 32.6 Red -semi Wolf predation
gelatinous
120009 Calf ? 01 Mendeltna Wolf predation
120010 Calf ? 01/30/77 Gulkana Road kill
120011 Calf ? 02/08/77 Susitna take Drowned
120012 Calf ~ 02/10/77 Tok Highway 10.1 Red -gelatinous Winter kill
120014 12 ~ 02/21/77 Grayling Lake 88.0 White Wolf predation
120015 Calf ? 03/03/77 Susitna River at 74.2 67.6 Red Wolf Predation
Jay Creek
120016 16 ~ 02/16/77 Susitna River at Red -gelatinous Wolf predation
Clarence Lake Stream
120017 Calf ? 02/16/77 W. Fork Gulkana River Red -gelatinous Wolf predation
120018 Calf ? 03/14/77 W. Fork Gulkana River Red -gelatinous Winter kill
120019A 13 ~ 03/21/77 W. Fork Gulkana River Wolf predation
120019B Calf ? 03/21/77 W. Fork Gulkana River Pink spotted Wolf predation
120020 Calf ? 04/06/77 Gakona River 12.2 Red Wolf predation
·~ 120021 Yearling ~ 04/13/77 Richardson Highway 78.4 66.7 Pink Road kill
0 120022 Adult ~ 04/13/77 Edgerton Highway 91.5 Light pink Road kill
Table 48 (cont.).
Date of Approximate Percent fat Marrow
Accession fl Age Sex collection location longbone Ramus color Cause of death
120023 Adult ? 04/13/77 Unknown -Unit 13 90.0 Light pink Poached or road kill
120025 Calf ~ 04/14/77 Richardson Highway 29.9 34.3 Red crystals Nusiance kill
near wall
120026 Yearling ~ 04/28/77 Roundtop Mountain 93.3 White Brown bear predation
120062 Calf ~ 07/25/77 Glenn Highway 65.6 Red Road kill
120085 Yearling cJ 01/17/77 Rat Lake ~;.7!1 White Wolf predation
120089 Adult ~ 01/26/78 Tok cutoff Dry cavity Wolf predation
120090 Calf ? 02/03/77 Gulkana Wolf predation
120091 Adult ~ 01/22/78 Copper River Wolf predation
120092 Adult !j? 03/03/78 Richardson Highway Road kill
120093 Adult-10 yr. ~ 04/10/78 W. Fork Gulkana River 83.0 Pink Wolf predation
tooth wear
120094 Calf ~ 04/09/78 W. Fork Gulkana River l7. 7 Pink Wolf predation
120095 Calf cJ 04/09/78 Dog Creek 7.0 Red Wolf predation
120096 ???? ~ 04/10/78 Nickel Creek 84.2 Pink Wolf predation
120097 Calf ~ 04/10/78 Nickel Creek 7.5 Red Wolf predation
120184 Yearling ~ 05/12/78 Simpson Hill 36.5 Auto
120194 7 ~ 07/17/78 Cat Lake 45.3 Brown bear predation
120195 Calf 07/17/80 Fish Lake Brown bear predation
120196 Adult ~ 06/08/77 Middle Lake 8.5 Yellow Black bear predation
120197 2 ~ 01/23/79 Nelchina River 78.9 65.3 Wolf predation
120198 Yearling ~ 09/05/78 Haggard Creek 89.1 Pink Road kill
120200 9 cJ 03/08/79 Mile 17 Tok Road 67.0 32.1 Pink Road kill
120201 2 ~ 03/17/79 Chickaloon River 7.0 Probable road kill
120202 Yearling ? 03/12/79 Tyone Mountains 89 88.9 Pink Wolf predation
120203 Adult ? 02/20/79 Dog Lake 96 White Wolf predation
120204 Calf ? 02/22/79 Dog Lake 30 Pink Wolf predation
120205 Calf ? 02/26/79 Lily lake 73.2 Pink Wolf predation
120206 15 ~ 02/27/79 Lily lake 81.7 73.3 Pink Wolf predation
120207 Calf ? 02/27/79 Lily lake 68.4 Pink Wolf predation
120208 14 ~ 02/20/79 Tyone-Goose Creek 86.9 78.0. White Wolf predation
120209 6 ~ 02/26/79 Tyone River 78.2 Pink Wolf predation
120210 Calf ? 02/28/79 Oshetna River 56.2 Pink Wolf predation
120211 Calf ? 03/20/79 Tyone River lO.O Red Wolf predation
0 120212 Calf ? 02/25j78 Glenn Highway 18.4 8.2 Red Winter kill
'"' 120213 13 ~ 03/30/79 Tyone Creek 91.3 68.8 Pink Wolf predation
,. ~ ·~ '· \ r .~ ... "1 rr· l r· l r···· .. ·n r· 1 r 1 r" ·~ r!'' l r. n r ., r 1 r· .~, r··· l r _'] r~ [___] c.....:..J c.....J c.:_:] I
C' .,!1 (.__] q, ..
Table 48 (cont.).
Accession 11 Age
120350 Calf
120358 Adult
120371 Adult
120372 Adult
120373 Adult
120374 Adult
120409 Calf
120601 2
120602 Calf
120603 Adult calf
120604
120605
120606
120607
120608
120609
120610
120611
120612
120613
120614
120615
120616
120620
120657
120658
120659
120660
0
("'
Calf
Adult
Calf
2
Calf
10
Adult
Adult
Adult
14
Calf
Calf
12
Adult
Calf
16
Calf
Adult
Date of
Sex collection
~ 06/05/79
~ 05/17/79
~ 04/27/79
~ 10/-/78
? 08/17/79
~ 09/23/79
~ 12/16/79
~ 01/28/80
? 02/21/80
? 02/21/80
? 01/12/80
02/20/80
? 02/21/80
? 02/21/80
? 02/21/80
~ 02/21/80
~ 02/04/80
~ 04/01/80
~ 03/18/80
~ 03/13/80
? 03/25/80
? 03/26/80
~ 03/26/80
~ 04/22/80
~ 04/30/80
~ 03/08/80
~ 03/08/80
~ 05/21/80
Approximate Percent fat
location longbone Ramus
West Fork Gulkana 25.5
Richardson Highway 85.6
Sanona Creek
West Fork Maclaren R. 91.6 76.5
Twin Lakes
Kenny Lake
Tons ina 11.6 27.7
Kenny Lake 90.4 68.1
Susitna Lake 24.0
Oshetna River 46.5
Sanona Creek 89.7
Black River 84.5 61.6
Little Nelchina River 53.3 41.0
Old Man Lake 84.1 65.2
Little Nelchina River
Grayling Lake 88.1 72.6
Eureka 60.8
Kenny Lake 98.1
Tyone Mountains 81.2
Black River 83.7 67.2
Squaw Lake 57.1 44.9
Black River 61.2 60.0
Oshetna River 87.3 65.8
Watana Creek 83.9 54.6
Glenn Highway 5.9 9.1
Sanona Creek 85.6 77.1
Sanona Creek 33.9
Glennallen 15.0
[_:J' ~ c...:J . ..... . l:
Marrow
color Cause of death
Pink Unknown
Pink Winter kill
Wolf predation
White Unknown predation
Brown hear predation
Road kill
Fink Road kill
Pink Accident
Pink Wolf predation
Pink Wolf predation
Pink Wolf predation
Pink Wolf predation
Pink Wolf predation
Pink Wolf predation
Wolf predation
Red Unknown
Pink Potlach kill
White Shot
Pink Wolf predation
\Yhite-red Wolf predation
Red Wolf predation
Pink Wolf predation
Pink-red Wolf predation
Pink Tagging
Pink Winter kill
Pink-red Wolf predation
Pink Wolf predation
Pink Winter kill
r•: .. 1
"----.1
Table 48 (cont.).
Date of Approximate Percent fat Marrow
Accession fl Age Sex collection location longbone Ramus color Cause of death
120214 Calf d 04/02/79 Richardson Highway 10.3 Pink Auto
120215 9 9 04/01/79 Tolsona Lake 87.2 75.9 Pink Accident
120216 Adult 9 03/30/79 Boulder Creek 91.5 Pink Winter kill
120217 Calf ? 03/23/79 Tyone Creek 16.8 Pink Wolf predation
120218 Calf ? 03/25/79 Tyone River 6.9 Pink Wolf predation
120219 Calf ? 03/25/79 Oshetna River 7.6 Pink Wolf predation
120220 2 9 03/24/79 Brushkana Creek 68.1 45.5 Pink Prob. wolf predation
120221 Adult ? 03/25/79 Lone Butte 74.1 61.9 Pink Wolf predation
120222 Calf ? 04/18/79 Tyone Creek 7.5 12.1 Pink Wolf predation
120239 Calf 9 03/28/79 Mendeltna 24.6 Tagging mortality
120270 Calf d 03/30/79 Tyone Mountains 15.6 Red Prob. tagging mort.
120277 Calf d 03/20/79 Tyone River Bear predation
120286 Adult 9 04/14/79 Valdez Creek 8.6 Pink Winter kill
120188 Calf 9 04/10/79 Glenn Highway 12.8 31.0 Red Tagging mortality
120289 Calf d 04/23/79 Nabesna Road 42.4 17.7 Red Probable road kill
120292 Calf d 05/18/79 Square Lake 21.5 Light pink Winter kill
120309 Adult ? 04/27/79 Oshetna River 59.5 White Wolf predation
120310 14 ? 04/27/79 Moore Lake 71.6 26.3 White Wolf predation
120311 Calf ? 04/27/79 Oshetna River 53.4 60.5 Pink Wolf predation
120312 Calf ? 04/27/79 Moore Lake 8.1 Red Wolf predation
120313 Calf ? 04/27/79 Tyone Mountains 7.7 Red Winter kill
120314 Calf 04/27/79 Tyone Mountains ' 7. 7 Red Winter kill
120315 Calf d 04/27/79 Tyone Mountains 8.1 9.9 Red Winter kill
120316 Calf 04/.27/79 Tyone Mountains 7.6 11.8 Pink Winter kill
120317 Calf 04/27/79 Moore Lake 9.3 Red Winter kill
120318 2 9 05/01/79 Richardson Highway 92.6 Pink White Unknown
120319 14 9 04/27/79 Moore Lake 87.7 84.6 White Wolf predation
120320 Calf ? 04/27/79 Moore Lake Wolf predation
120321 Calf d 04/27/79 Moore Lake 11.4 14.7 Pink Winter kill
120322 Calf ? 04/27/79 Sanona Creek 28.9 12.9 Red Winter kill
120323 Calf ? 04/27/79 Tyone Creek 47.1 9.9 Red Winter kill
120324 Calf ? 04/27/79 Lily Lake 9.0 7.8 Red Winter kill
120325 Calf 04/27/79 Sanona Creek 31.7 Red Wolf predation
120326 Calf ? 04/28/79 Nickolson Lake 20.4 23.3 Pink Unidentified predator
-' 120343 Calf 9 06/08/79 Middle Fork Susitna R. 35/3 Pink Pneumonia
~ 120344 Calf d 07/03/79 Susitna River 88.7 Pink Bear predation
•' r r·c liT r· "l r
!"'" " -,
ii.--.1 LJ
Table 49.
Accession /1
55100A
55100B
55100C
55100D
55101
55102
55103
55104
55105
55106 .
55107
55108
55109
55110
55111
55112
55113
55115
55116
55117
55118
55119
55121
55122
55123
., .
Age, sex condition as determined by percent marrow fat, and cause of mortality of
in GMU 13 of southcentral Alaska from November 1975 through June 1980.
Date of Approximate Percent fat Marrow
Age Sex collection location longbone Ramus color
Adult cJ 11/12/75 Tyone Creek 8
Adult 12/05/75 Keg Creek
Adult ~ 04/21/76 West Fork Gakona River 8
Adult ~ 05/03/76 Grayling Lake 60
caribou kills examined
Cause of death
Wolf predation
Wolf predation
Wolf predation
Wolf predation
Adult cJ ll/03/76 Marie Lake 10.4 Red-gelatinous Possible wolf predation
4-7 cJ 01/18/77 Tolsona Creek 81.5 83.8 Pink Wolf predation
10 ~ 01/25/77 Moose Lake 87.5 Light pink Wolf predation
Calf ? 04/08/77 Lake Louise 59.3 Red Winter kill?
9 !;? 03/28/79 Nickoli Lake 82.4 67.7 Pink Wolf predation
10+ ~ 04/01/79 Moore Lake 8.6 28.9 Red-pink Winter kill
3-5 ~ 03/31/79 Copper River 86.1 69.4 Pink Wolf predation
3-5 ~ 03/31/79 Copper River 87.6 66.9 Pink Wolf predation
2+ ~ 03/31/79 Copper River 63.1 48.4 Red Wolf predation
3-5 !;? 03/31/79 Copper River 85.7 68.8 Pink Wolf predation
2+ !;? 04/01/79 Copper River 57.3 49.2 Pink Wolf predation
2+ !;? 04/01/79 Copper River 52.2 52.4 Pink Wolf predation
3-5 cJ 04/01/79 Copper River 83.4 66.7 Pink Wolf predation
6-9 cJ 04/18/79 Glenn Highway 79.2 17.1 Pink-white Wolf predation
Adult !;? 04/23/79 Lake Louise Road 90.4 84.8 Pink Wolf predation
Old adult !;? 04/30/79 Bell Lake 61.6 40.9 Pink Wolf predation
Calf ? 04/30/79 Bell Lake 36.6 Pink Wolf predation
Yearling ? 04/? /79 Unit 13 69.5 62.2 Pink Unknown
Adult !;? 01/27/80 Susitna Lake 90.5 Pink Wolf predation
Calf ? ? 02/03/80 Minnesota Lake 75.6 Pink Wolf predation
Adult cJ 02/21/80 Tolsona Creek 29.4 Pink Wolf predation
The moose carcasses examined (Table 48) consisted of 68
wolf kills, 29 accidental deaths, 19 winter kills, 7 bear kills,
and 2 killed by unknown predators. Wolf kills were comprised of
32 calves (27 of unknown sex) and 36 adults. Of the adult moose
killed by wolves, 2 were bulls, 27 were cows, and 8 were of
unidentified sex. Accidental moose kills included 2 adult
bulls, 13 adult cows, 1 adult of unknown sex, 3 bull calves, 7
cow calves, and 3 calves of unknown sex. Winter kills consisted
of 5 adult cows and 14 calves, and bear kills were comprised of
4 adults and 3 calves. Two kills were by unknown predator
species.
Franzmann and Arneson (1976), on the Kenai Peninsula, found
that percent fat varied by month of study because of a lag in
deposition and mobilization of marrow fat. High fat values were
observed in summer and fall but these values declined as winter
progressed, with a definite decrease occurring between December
and January. We expected a similar pattern for Nelchina moose
so we compared percent marrow fat for calves and adults for the
five classified causes of mortality by man~~ of kill. None of
these comparisons were significant (P>0.05), however, the data
did reveal a tren!i of declining fat as winter progressed and,
thus, did not contradict the hypothesis.
Aerial observations during this study and pr:;vious studies
in GMU 13 (Stephenson and Johnson 1972, 1973) have indicated
that wolves were preying upon calf and short yearling moose in
excess of their presence in the moose population. Of particular
importance, however, was the physical condition of the
individual moose wolves were preying upon. If wolves prey upon
calves and yearlings in weak physical condition then it might be
surmised that these animals would die anyway due to starvation
(winter kill). Thus, the impact of wolves on calves and
yearlings would be diminished. If, however, they were preying
upon healthy calves which could or would otherwise survive, then
wolf predation would hamper recruitment into the population and
possibly could prevent it from growing.
Marrow fat levels of calf and short yearling moose killed
by wolves (X = 35.7%) were significantly (P<0.01) higher than
those dying from accidental causes (X= 25.9%). There were also
significant differences (P< 0. 01) between marrow fat of
winter-killed calves and short yearlings (X = 14.8%) and those
killed by accidental causes (25. 9%). On the Kenai Peninsula,
Alaska, Franzmann and Arneson (1976) suggested ~~at marrow fat
values below 10 percent were indicative of winter-killed
(starved) moose. In this study, six calves dying from
starvation had percent fat values in excess of 10 percent (range
of 10.1 to 47.1). Perhaps these latter calves died as a result
of other complicating factors besides malnutrition. During this
study we noted several calf carcasses with malformed (swollen)
leg bones in the vicinity of the carpus and tarsus.
Unfortunately we were unable to document the occurrence of this
abnormality in the moos~ population nor do we know what
influence it may have on a calf's ability to survive.
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We· were unable to explain the differences in percent fat
between accidental and wolf-killed. calves. In other studies,
there were no apparent differences in marrow fat levels between
wolf-caused and accidental deaths among calves (Stephenson and
Johnson 1973; Franzmann and Arneson 1976). Perhaps the
differences recorded here were attributable in part to sample
size, especially because most of t.~e samples of accidentally
killed calves were collected in late March and April when
percent fat values were declining. Also, it appears possible
that the opportunistic manner in which samples of moose killed
in accidents were collected may have biased our sample toward
moose which may have been close to succumbing to winter kill.
There probably are few if any differences between wolf and
accidentally killed calves. In any case these comparisons
indicate that wolves were preying upon relatively healthy calf
and short yearling moose during this study.
In an effort to better assess age and physical status of
adult moose taken by wolves, data from this study were combined
with those collected from 1970-1972 (Stephenson and Johnson
1972; 1973).-In addition, the age composition of wolf-killed
adult moose was compared with that of moose live-captured and
tagged from 1976 through spring 1980 (Table 50). Wolf-killed
adult moose averaged 9.3 years of age (S.D. = 5.4) while tagged
moose averaged 7.1 years of age (S.D. = 3.7). Significant
differences in age were detected for the following: wolf-killed
versus tagged moose (P<O.OOS), wolf-killed versus winter-killed
moose (P<0.10), wolf versus accidentally killed moose (P<O.OS),
winter versus accidentally killed moose (P<O. 01), and tagged
versus winter-killed moose (P<0.001). There was not a
significant difference, however, between tagged and accidentally
killed moose {P>0.10).
Our findings that wolves were preying upon moose which were
older than those tagged from 1976-1980 suggest that wolves
selected for relatively old adults. This comparison includes
wolf kills from the period 1970-72, during which time no moose
were tagged. As a result, the comparisons are not directly
comparable due to the assumption that the tagged animals also
represented the age structure of moose during t..llis earlier
period. Therefore, we compared the age structure of tagged
moose to wolf kills by individual year from 1976-1980.
Significant (P<0.10) differences existed for 1976, 1977, and
1980. Sample sizes were too small (df = 3) for 1978; however,
no significant differences (P>O.OS) were evident for 1979.
Winter 1978-79 was the second most severe winter in the Nelchina
Basin (Eide and Ballard, in review). Thus, during a severe
winter wolves preyed upon adult moose of various ages in
proportion to their presence in the population, while during
mild winters older moose were preyed upon more heavily.
Winters 1970-71, 1971-72 and 1978-79 are considered the
most severe winters in the study area in terms of total snowfall
and its impact on the moo_se population (Stephenson and Johnson
1 0 7
Table SO. Summary of average age of adult cow moose suffering mortality
in comparison to average age of tagged moose in GMU 13 of
southcentral Alaska from 1971 through spring 1980.
Wolf kills Accidental kill Winter kills Tagged samEle
--Year x age n x age n x age n x age n
1971 6.60 5
1972 7.75 16 9.67 3 14.50 2
1975 7.50 2
1976 11.33 3 6.54 39
1977 13.20 5 1.00 1 15.0 1 6.02 49
1978 10.50 2 1.00 2 9.67 3
1979 9.43 7 5.50 2 8.58 12
1980 14.00 3 2.00 1 8.76 32
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1972, 1973; Bishop and Rausch 1974, Ballard et al. 1980).
Although we could detect no significant correlations (P> 0.10)
between total snowfall and average age of adult moose preyed
upon by wolves, it does appear that during "mild" winters wolves
were selecting older moose, whereas during "severe" winters they
were preying upon younger moose, indicating that younger moose
became more vulnerable to wolf predation because of deep and
perhaps crusted snow.
There were no significant (P> 0. 05) differences in mean
marrow percent fat of adult moose killed accidentally and those
preyed upon by wolves for winters of 1970 through 1980
(Table 51). Winter-killed adult moose had a significantly lower
(P<O.OOl) percent marrow fat (X = 24.4%) than wolf-killed moose
(X= 64.3%), while winter-killed adult moose had lower (P<0.05)
marrow percent fat than did adult moose dying from accidental
causes (X = 58.1). These data suggest that from 1970 through
winter 1979-80, wolves were preying upon moose that were in
better physical condition than those dying from winter-kill but
similar physical condition to those dying from accidental causes
(which were assumed to be in normal physical condition).
Because of small sample sizes it was difficult to make
valid comparisons of percent marrow fat levels in different age
classes of adult moose killed by wolves. There did not appear
to be any relationship between average age of adult moose and
average marrow percent fat. Therefore, we conclude that during
winter, wolves preyed upon adult, short yearling, and calf moose
which were in better condition than those succumbing to
winter-kill. During winter, wolves were preying upon calves out
of proportion to their presence in the moose population. In
severe winters young, relatively heal thy adults, were being
preyed upon while in relatively mild winters relatively healthy
older adults were being killed. The latter classification,
except for physical condition, appears to correspond with the
traditional predator-prey scheme elaborated upon by Mech (1970).
Markgren (1969) suggested that even though twinning rates
decline with age, most moose continue producing calves and,
therefore, removal of these productive members of the moose·
population must be of concern to game managers.
Twenty-two of 25 caribou carcasses examined during this
study were attributed to wolf predation (Table 49). Two of the
remaining kills (one calf and one 10+ yr old cow) were
classified as winter kills, and cause of death for one was
unknown. Wolf caused mortalities were comprised of 20 adults (6
bulls, 13 cows and 2 of unknown sex) and 2 calves (sex unknown).
Ages of adUlt caribou killed by wolves ranged from 1.5 to 10 or
more years. The overall aae structure of wolf-killed adults
appeared to be fairly low (xJ= 4.91, S.D. = 2.80), however, 7 of
these caribou were killed by one wolf pack (Appendix IV) and,
therefore, may not be representative of caribou normally preyed
upon by wolves.
1 0 9
Table 51. Average bone marrow percent fat of moose kills
examined from 1971 thro~~h spring 1980 in GMU 13
of southcentral Alaska.-
Wolf kills Accidental kill Winter kills
Year x% fat n x% fat n x% fat n
1971 81.18 5
1972 55.39 13 52.60 2 6.78 5
1975 41.00 2
1976 56.70 4
1977 66.20 3 64.53 4 42.85 2
1978 57.78 5 62.80 2
1979 76.40 6 35.83 3 50.05 2
1980 85.53 3 79.45 2
y 1971 and 1972 data from Stephenson and Johnson
(1972;1973).·
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The physical condition of wolf-killed adult caribou as
indicated by percent fat marrow was quite variable, ranging from
8 to 80.5 percent but averaging 63.7 (n = 20, S.D. = 28.4).
Neiland {pers. cornrn.) considered caribou with less than 25
percent fat to be in poor physical condition. Based upon that
criterion 4 of 20 (20%) of the adult caribou killed by wolves
were in poor physical condition (<30% fat). Greer (cited in
Franzmann and Arneson 1976) classified elk (Cervus canadensis)
as in excellent condition when fat exceded 80 percent. Using
Greer's general criteria for elk, 11 of 20 (55%) wolf-killed
caribou had percent fat in excess of 75 percent and would be
considered in good physical condition. The remaining five
wolf-killed caribou had fat values varying from 52.2 to 63.1 and
averaged 58.8 percent (S.D. = 4.29). The condition of these
animals was probably declining as winter progressed.
Because our sample of caribou killed by factors other than
wolf predation was so small, it is impossible to compare the
physical condition of wolf kills with other mortality factors.
It is also difficult to generalize about the condition ·Of
caribou take~ by wolves during this 5-year study. Fifty-six
percent of the wolf-killed caribou were examined during winter
1978-79 which, in terms of total snowfall, was the second most
severe winter since 1952. The relatively large number of
caribou carcasses examined during that winter reflect the
increased vulnerability of caribou due to excessive snow
accumulation. Caribou taken by wolves during that severe winter
were, for the most part, in relatively good physical condition.
Whether caribou taken later in spring 1979 were also in good
condition is unknown.
Wolf Predation Rates
We calculated predation rates for selected study packs with
which we had the most radio contact (Table 52) by dividing the
number of kills we observed into the number of pack days.
Fuller and Keith (1980) suggested that this method tends to
overestimate the kill rate because wolf packs comprised of less
than 12-15 animals (includes nearly all of our packs) often
remain at kills longer than 1 day, and, therefore, ~~e chance of
locating them at a kill is greater than the actual daily kill.
Nevertheless, the average rate listed in Table 52 provides
a fair approximation of the kill rate for a GMU 13 wolf pack.
It also reveals the range of predation estimates which can be
generated by this method of calculation. According to these
data, the year-round kill rate of various packs was from
1 kill/3.1 days to 1 kill/12.7 days and averaged
1 kill/4.9 days. This method of estimating kill rates assumes
that all kills were observed. In this study there was large
variation in observability by pack area, and for some packs we
probably were unable to locate a significant number of ungulate
kills. Also it was possible to miss one kill when a pack had
two kills but radio-collared wolves were all at one. We
1 1 1
Table 52. Rates of predation of radio-collared wolf packs
studied in GMU 13 of southcentral Alaska from
April 1975 through June 1980.
Wolf pack
Brushkana
Butte Lake
Deadman
Deep Lake
Ewan
Hogan Hill
Jay Creek
Keg Creek
Maclaren River
Mendeltna
Middle Fork
Saint Anne
Sinona
Susitna
Tolsona
Tyone
Watana
Number
pack days
34
31
38
145
151
176
43
240
52
128
27
76
170
135
89
170
30
1735
Number of
kills observed Days/kill
5 6.8
5 6.2
3 12.7
23 6.3
18 8.4
39 4.5
10 4.3
28 8.6
12 4.3
36 3.6
5 5.4
20 3.8
40 4.3
43 3.1
29 3.1
33 5.2
3 10.0
352 x4.9
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subjectively believe that we missed a significant number of
kills for the following packs: Deadman, Ewan, Keg Creek, Middle
Fork, and Watana. Also, these estimates include those kills
which were also occupied by brown bears (10.6%) and, thus, some
of.the kills were not made by wolves even though they fed upon
them. Further these kill rates include observations of prey
species other than ungulates and do not take into account the
large variations in individual pack numbers that occurred for
some packs.
During late spring and early summer in 1977 and 1978, we
intensively monitored three wolf packs (1 in 1977 and 2 in 1978)
in an effort to better document food habits and determine
predation rates for that season of the year. During this
portion of the study we also had an opportunity to witness some
interactions between wolves and brown bears. Predation rates
and brown bear-wolf interactions for the Mendeltna pack in 1977
and the Hogan Hill pack in 1978 were combined and presented in a
paper entitled 11 Gray Wolf-brown Bear Relationships in the
Nelchina Basin of Southcentral Alaska 11 which was presented by
Ballard at the Portland Wolf Symposium in August 1979. A copy
of the paper is presented as Appendix III and, therefore, no
further discussion of those data are warranted at this point in
the report. One pack studied in 1978, which was not reported
upon in the aforementioned paper, deserves mention because it
provides data on the activity of a single pack member.
Between 29 May and 24 June 1978, the two radio-collared
members (#'s 009 and 204) of the Deep Lake pack were monitored
either once or twice daily. From 30 May through 24 June wolf
number 009 ( 3-or 4-year-old nonbreeding female) was observed
traveling the Deep Lake territory alone except on two occasions.
During this period she was observed alone at a total of four
kills: one calf moose which was also occupied by a brown bear
sow with a yearling cub, one adult moose, and two kills which
could not be identified. We do not know if this single wolf
actually made any of these kills. If we assume that she did,
the indicated rate of kill would be one every 6.5 days.
However, we suspected that she also fed on small mammals and
birds and thus this rate may pertain only to ungulates.
During winters 1979 and 1980 we began radio-tracking two to
three wolf packs every 2 days in an effort to determine
predation rates according to sex, age and number of individuals
per pack.
1979 Predation Rate Study
Susitna Pack -From 20 February through 22 April 1979, this
pack of two adults and seven pups was observed on 18 kills, 17
of which were made during 't.J."le study period (Table 53). Kills
were comprised entirely of moose: 10 calves, 7 adults, and 1
long yearling. Due to deep snow which provided excellent
tracking conditions we were able to back track these wolves to
1 1 3
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Table 53. Chronological record of kills made by the Susitna wolf pack in
Game Management Unit 13 of southcentral Alaska from 20 February
through 22 April 1979.
Estimated
Date of Kill Percent date of kill
observation Time made Species and age consumed or comments
2/20/79 2:30 p.m. Yes Moose -adult 100% 2/19/79 --~ 2/22/79 9:30 a.m. Yes Moose -calf 100% 2/21/79
2/24/79 10:00 a.m. Yes Moose -calf 100% 2/23/79 0
L
2/26/79 8:20 a.m. Yes Moose -calf 100% 2/25/79 ~ 2/27/79 12:30 p.m. Yes Moose -adult cow 10-20% 2/27/79 r
Moose -calf
2/28/79 11:40 a.m. No still on ..
. 2/27 kills r 3/02/79 9:00 a.m. No close to
2/27 kills b._
3/05/79 8:45 a.m. No
3/07/79 11:30 a.m. Yes Moose -calf 95-100% 3/06/79 F
3/09/79 3:15p.m. Yes Moose -adult 90-100%
3/12/79 1:50 p.m. Yes Moose -adult 85-95% 3/11/79 L
3/14/79 4:30 p.m. No
3/16/79 1:50 p.m. No r
3/18/79 4:00 p.m. No i~
3/20/79 2:45 p.m. Yes Moose -calf 100% 3/19/79
3/23/79 2:05 p.m. Yes Moose -calf 100% 3/23/79 a.m. ..---
3/25/79 9:00 a.m. Yes Moose -calf 100% 3/24/79 p.m.
3/27/79 5:45 p.m. Yes Moose -adult 75% 3/26/79 L
3/29/79 8:10 a.m. No still on
3/27 kill <=
3/30/79 11:10 a.m. No still on L
3/27 kill
3/31/79 1:15 p.m. No still on r
3/27 kill
4/02/79 12:30 p.m. Yes Moose adult cow 25% ~
Moose -calf
4/04/79 3:35 p.m. No 75% still on r
4/2 kills ' '-· 4/06/79 10:55 a.m. No 100% still on
4/2 kills
4/08/79 12:00 noon No ~ mile from ..
4/2 kill L
4/10/79 2:30 p.m. No
4/12/79 10:00 Yes Moose adult cow 30% 4/12/79 -_r a.m.
Moose -long yrlg.
4/14/79 5:50 p.m. No 60% still
on 4/2 kills F
4/16/79 7:05 a.m. No
4/18/79 2:30 p.m. No traveling SW -L
2 mi. from den
4/20/79 8:20 a.m. No
4/22/79 6:50 p.m. Yes Moose -calf assume 75
to 100%?
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their previous locations and, therefore, felt confident that no
kills were missed. During 62 pack days of observation the kill
rate was one per/3.6 days. Calf moose (short yearlings)
comprised 55.6 percent of the kills. Of the 17 kills, six were
cows with calves or long yearlings. All kills were heavily
consumed. Assuming that short yearlings weighed an average of
19 7 . 5 kg ( from Fr anzmann and Arneson 19 7 3 ) and that adults
(includes long yearlings) weighed 427.5 kg, and also that 75
percent (Peterson 1977) of the live weight was consumable, then
6.7 kg of food was potentially available to each pack member per
day.
Tyone Pack -From 20 February to 24 April 1979 1 this pack
was comprised of an adult female and an adult male. We were
able to observe these wolves every other day from 20 February
through 14 March and from 18 March through 24 April 1979
(Table 54). These observations represent 58 pack days. During
this period we observed eight moose kills, seven of which were
made during the study period yielding a kill rate of 1/8.3 days.
No cow-calf pairs were killed. In contrast to the Susitna pack,
75 percent of the kills from the Tyone pack were adult moose.
Using the same conversion factors used for the Susitna pack to
determine kgs of available prey 1 each adult wolf had 16.4 kgs of
food available per day. They did not, however, consume this
amount. These wolves lingered around adult moose kills
considerably longer than did the Susi tna pack and we noticed
that scavengers, particularly ravens (on one kill 30 were
observed) and wolverines ( Gulo gulo) were taking advantage of
the time required for these two wolves to consume a kill.
1980 Predation Rate Study
Susitna Pack -From 23 January through 27 March 1980,
members of the Susitna pack were observed on nine kills
{1able 55). These data were divided into two periods because of
changes in pack numbers described in the pack history section.
The first period extended from 23 January through 12 February
1980 during which time the pack numbered seven :3 adults, 2
yearlings, and 2 pups). During this interval they preyed upon
four caribou and one adult moose for a kill rate of 1/4.2 days.
Car~bou comprised 80 percent of the kills in 1980 while in 1979 1
all of the observed prey were moose. Differences appeared to be
related to the availability of prey because in 1979 few, if any/
caribou were available to this pack while in 1980 relatively
large numbers of caribou overwintered in this pack's area.
In 1980, changes in prey availability and abundance and
perhaps in pack numbers also appeared to alter the movement
patterns of this pack compared to 1979. In 1979, they had
frequented the drainages of the Tyone River while in 1980 they
occupied the area near Susi tna Lake. Based upon the kills
observed during this time, this pack of seven wolves had 5.3 kg
of available food/wolf/day.
1 1 5
Table 54.
Date of
observation
2/20/79
2/22/79
2/24/79
Chronological summary of kills the Tyone wolf pack were observed at
from 20 February through 24 April 1979 in Game Management Unit 13
of southcentral Alaska.
Estimated
Kill
made
Percent date of kill
Time Species and age consumed or comments
1:00 p.m. Yes Moose -adult 50%
12:00 noon No
10:03 a.m. No
9:15 a.m. Yes Moose -adult 10%
1:45 p.m. Yes Moose -calf 75%
2/19/79(est.)
little movement
since 2/22
observation
2/25/79 (est.)
2/27/79(est.)
2/26/79
2/28/79
3/02/79
3/05/79
3/07/79
3/09/79
10:30 a.m.(est.) No
3/12/79
3/14/79
3/16/79
3/18/79
3/20/79
3/23/79
3/25/79
3/27/79
3/29/79
3/31/79
4/02/79
4/04/79
4/06/79
4/08/79
4/10/79
4/12/79
4/14/79
4/16/79
4/18/79
4/20/79
4/22/79
4/24/79
9:45 a.m. No
11:50 a.m. No
3:50 p.m. Yes Moose -calf 50%
3:30 p.m. No
5:00 p.m. No
No observation due to poor weather
4:35 p.m. Yes Moose adult 10%
3:05 p.m. No SO%
2:25 p.m. No
No observation due to wolf collaring operation
6:15 p.m. No
8:35 a.m. No
1:05 p.m. No
12:05 p.m. No
4:00 p.m. No
11:10 a.m. No
1:00 p.m. Yes
3:00 p.m. No
11:00 a.m. No
6:15p.m. No
7:45 a.m. No
3:05 p.m. No
7:35 a.m. No
6:20 p.m. Yes
7:50 a.m. No
Moose -adult
Moose -adult
40%
80%
90%
100%
25%
3/8/79 late or
3/9/79 early
morning of 3/18
still on 3/18 kill
tracks lead by old
kill of 2/20/79
have been visiting
very old moose kill
on ridge top
still on 3/29 location
observed digging den
observed at old moose
kill site of 2/28/79
still at old kill site
of 2/28/79
4/7 /79(est.)
still at 4/8/79 location
still at 4/8/79 location
had visited 4/8 kill
female observed at den
site
at kill site of 4/8
4/22/79
had visited 4/22/79
kill
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Table 55. Chronological summary of kills at which the Susitna wolf pack was
observed from 23 January through 12 February and from 12 March £?rough
27 March 1980 in Game Management Unit 13 of southcentral Alaska-.
Date of
observation
1/23/80
1/25/80
1/27/80
1/28/80
1/29/80
2/01/80
2/03/80
2/05/80
2/07/80
2/10/80
2/12/80
3/12/80
3/14/80
3/16/80
3/18/80
3/20/80
3/22/80
3/25/80
3/27/80
Time
1:00 pm
11:08 am
12:23 pm
9:23 am
11:05 am
2:32 pm
12:03 pm
10:30 am
2:48 pm
11:50 pm
2:00 pm
9:35 am .
12:20 pm
12:30 pm
10:45 am
10:45 am
3:40 pm
11:30 am
8:45 am
Kill
made
Yes
No
Yes
No
Yes
No
Yes
No
No
No
Yes
No
Yes
No
No
Yes
Yes
Yes
No
Species and age
Moose -adult
Caribou -adult 9
Caribou -adult
Caribou -adult
Caribou -assumed
adult
Caribou -adult 9
Caribou -adult
Moose -assumed calf
Moose -calf
Percent
consumed
75%
90%
100%
100%
100%
90%
95%
100%
100%
90%
Estimated
date of kill or comments
1/22 or 1/23/-7 wolves
1/26
1/28
2/2
- 7 wolves
- 7 wolves
- 7 wolves
2/11 or 2/12 - 7 wolves
- 4 wolves
3/13 - 4 wolves
Still on kill of 3/14 - 4 wolves
3/18 -4 wolves
3/20 or 3/21 - 4 wolves
3/23 or 3/24 - 4 wolves
From 23 January through 12 February 1980, pack was comprised of 3 adults,
2 yearlings and 2 pups while from 12 March through 27 March, pack was comprised
of 2 adults, 1 yearling and 1 pup or yearling.
1 1 7
During the second sampling period from 12 March through
27 March 1980, this pack numbered four wolves providing an
opportunity to compare kill rates of the pack when at a lower
number. Kills were comprised of 1 adult moose, 1 calf moose,
and 2 adult caribou which provided 5.7 kgs of available
food/wolf/day. The kill rate was 1/4.0 days, which was fairly
close to the rate of kill observed when the pack included seven
members.
In an effort to determine possible impacts of this wolf
pack on moose during early March we conducted a moose survey in
this pack area2 Four and one-half hours of ~ight time
(0.59 minutes/mi ) were spent surveying this 462 mi area, and
51 moose were counted: 43 adults and 8 calves (15. 7%). The
observer subjectively estimated that he may have observed 25
percent of the moose present. Assuming moose were being taken
at the rate indicated {caribou comprised 66.7% of kills), this
pack killed eight adults and four calves from December through
April. These kills represented 19 percent of the adult moose
and 50 percent of the calf moose counted in March after most of
the predation had occurred. If we assumed the observer had
indeed counted only 25 percent of the moose, and if we include
the projected kills as part of the base population, the
projected predation loss would have been 4 percent of the adult
moose and 11 percent of the calf (short yearling) moose in the
area. Wolf predation appears to be contributing to high
mortality of short yearling moose.
Tyone Pack -During early 1980 weather and tracking
conditions were excellent, allowing this pack of two adults and
six pups to be moni tared during a 54-day period ( 23 January
through 16 March 1980). The pack was observed on 11 kills: 3
adult moose, 7 calf moose, and 1 adult caribou (Table 56). The
prey used by this pack was similar to that observed in 1979 when
it was comprised of two adults. However t in 1979 calf moose
(short yearlings) comprised only 29 percent of ~~e kills while
in 1980 they comprised 64 percent of the kill indicating a
change in prey selectivity based on pack composition. This pack
was observed on a fresh kill at the rate of 1/4.9 days, with an
estimated 4.9 kgs of food available/wolf/day.
Similar to that reported for the susi tna pad~, we also
counted moose in this pack area dur~ng early March 1980. Four
and o~-half hours (0.89 minutes/mi ) were spent surveying the
302 mi pack area, and 266 moose were counted: 221 adults and
45 calves (17%). The observer subjectively estimated that he
had observed 50 percent of the moose present. We extrapolated
the observed moose kill rate to the months of December through
April which yielded an estimated kill of 8 adult moose and 20
calf moose. These projections comprised 4 percent of the adult
moose and 44 percent of the calf moose observed during the
survey, which was conducted after most of ~"le predation had
occurred. If we assumed the moose survey had only observed 50
percent of the moose and if we include the projected kills as
1 1 8
1
,_j
'! ~ I u
.. .-
~
L
r
L
L
L
L
L
L
L
L
L
L
r
' L
J
J
n
J
1
o~
0
u
J
J
J
~' u'
J
Table 56. Chronological summary of kills at which the Tyone wolf pack (2 adults,
Date of
observation
1/23/80
1/25/80
1/27/80
1/29/80
2/01/80
2/03/80
2/05/80
2/07/80
2/09/80
2/12/80
2/14/80
2/16/80
2/17/80
2/18/80
2/20/80
2/22/80
2/24/80
2/25/80
2/27/80
2/29/80
3/02/80
3/04/80
3/06/80
3/08/80
3/09/80
3/10/80
3/12/80
3/14/80
3/16/80
6 pups) was observed from 23 January through 16 March 1980 in Game Management
Unit 13 of southcentral Alaska.
Time
11:30 am
10:35 am
11:51 am
11:25 am
2:06 pm
10:56 am
10:00 am
3:05 pm
10:50 am
2:20 pm,
10:30 am
5:22 pm
1:00 pm
10:10 am
8:50 am
12:00 m
2:30 pm
9:15 am
10:15 am
4:30 pm
11:00 am
10:30 am
11:45 am
9:45 am
10:00 am
11:00 am
1:00 pm
Kill
made
No
Yes
No
Yes
No
No
No
No
No
Yes
No
No
No
No
Yes
No
Yes
No
No
No
No
No
No
Yes
Yes
No
No
No
Yes
Yes
Percent
Species and age consumed
Caribou -adult d
Moose -calf 95%
Moose -calf 50-70%
100%
Moose·-adult~ 75%
Moose -calf 60%
95%
100%
100%
100%
Moose -calf 5%
Moose -adult ~ 25%
Moose -calf <5%
60%
Moose -adult 60%
Moose -calf
Estimated
date of kill or comments
1/28
Dug up old kill
Visited caribou kill
of 1/25/80
Visited old unidentified
kill
2/11 or 2/12
still on kill of 2/12
2/19
2/23/or 2/24
Still on kill of 2/24
Still on kill of 2/24
Still on kill of 2/24
Still on kill of 2/24
3/6
3/8 killed 2 Susitna pack
members and consumed
3/8 two ptarmigan
Still at kill of 3/8
Still at kill of 3/8
Still at kill of 3/8
3/13
3/16
1 1 9
part of the base population, the percentages would have been 2
percent of the adult moose and 18 percent of the calf (short
yearling) moose in the area. In either case, it appears that
wolf predation on short yearlings in this area was a significant
mortality factor. -
Predation Rates Summary
Pe-terson (1977) suggested that wolf packs preyed upon
ungulates at relatively constant rates regardless of differences
in pack size. Table 57 summarizes predation statistics acquired
from studying the Susitna and Tyone wolf packs in 1979 and 1980.
To determine if a relationship existed between pack size and
predation rates in the Nelchina Basin, we performed a series of
correlation analyses (Snedecor and Cochran 1973). Pack size and
kgs of available food/day/pack were not significantly 2 {P<O. 05,
df 4) correlated even though positive relationships (r = 0.55)
appeared to exist. There was, however, a significant (P>O.OS)
~~~~)!~~fJdea~at~~n:h~~. ~~;we~. ~acfo<f¥,e :fld= ~~~2 )f i~~f~;~~~
that smaller packs have more food available per kill per
individual wolf than do larger packs. This relates only to
available food, however, and not to actual consumption which at
smaller pack sizes must be far less than the amount of food
available.
Fig. 5 illustrates the relationship between pack size and
predation rates calculated for the Susitna and Tyone pack during
winter 1978-79 and 1979-80. Pack size and days/kill were
positively correlated (P<O.OS, r = 0.83) suggesting that, for
the range of pack sizes studied, large packs were killing moose
or caribou more often than were smaller packs. These findings
do not necessarily refute those of Peterson (1977) because there
may not be differences in predation rates for packs larger than
nine individuals, which was the smallest pack studied by
Peterson {1977).
Summer Activity Patterns
Data pertaining to summer activity patterns of wolf packs
were collected in 1977 and 1978 concurrent with summer predation
studies. Wolf observations were classified into five general
categories: ( 1) resting at den, ( 2) traveling, {3) bedded away
from den, ( 4) at kill site and ( 5) stalking. The first four
categories are self explanatory; we classified an observation as
stalking-when we subjectively believed there was a relationship
between the wolf's presence and the presence of prey.
Tables 58 and 59 summarize the activity patterns of six
radio-collared members of the Mendeltna (1977) and Hogan Hill
{1978) wolf packs during late May and June of each respective
year.
1 2 ;;
J
j
r
' L
L
L
L
L
L
L
L
,--
r
~
L
J
J
J
]'•
o:
J
0
0
J
J
D
J
D
u
Table 57. Summary of predation statistics derived from intensive
radio-monitoring of the Susitna and Tyone wolf packs
during winters 1978-79 and 1979-80 in Game Management
Unit 13 of southcentral Alaska.
Size of Kgs of available Kgs of available Days/ungulate
wolf pack food/day/pack food/day/pack kill
9 60.1 6.7 3.6
8 38.9 4.9 4.9
7 36.9 5.3 4.2
4 22.9 5.7 4.0
2 32.8 16.4 8.3
1 2 1
N
lv
Figure 5. Relationship of wolf pack size to predation rates of two wolf
packs studied during winter 1978-79 and 1979-80 in Game
Management Unit 13 of southcentral Alaska.
10
y a + b lnX
9 9.20 + -2.53(lnX)
0.83
8
::.: 7 ~
p::
I'Ll 6 p..
Ul
I'Ll ~ 5 ~
gs
p:: 4 I'Ll ~ 3
2 )(
1
1 2 3 4 5 6 7 8 9 10
DAYS PER KILL
'
•' I r 'l r l rr· l
N
w
.,.. ..... 1
L....J
Table 58.
Individual
wolf
Adult cJ
122083
Yrl. cJ
122008
Yrl. 9
122007
Totals
c.....J r ~~
t·~ ~
'
Activity and association of three radio-collared members of the Mendeltna pack between late May
and July, 1977 in Game Management Unit 13.
Activit
Resting at den or Bedded away At kill
rendezvous site Traveling from den site Stalking Total
0 1 1+ 0 1 1+ 0 1 1+ 0 1 1:t 0 1 1+ Not found relocations
16 9 7 2 7 10 2 l 4 1 7 1 1 3 4 75
10 8 6 14 4 7 6 2 --4 6 2 --1 3 73
60 8 5 2 l ------l 1 0 0 78
8b 25 18 18 IT 18 8 3 9 I I4 3 I 4 7 226
Table 59. Activity and association of three radio-collared wolves from the Hogan Hill wolf pack
Management Unit 13 which were intensively monitored during late May and June 1978.
Individual
wolf
Adult gray cJ
122202
Yrl. gray d
122205
Yrl. gray 9
12"2206
Totals
Resting at den or
rendezvous site
2
1
4
7
3
2
2
7
10
12
8
30
' ,•
1'-1 r , ~
Traveling
5 3
2 1 2
5 1
12 I 6
,..., "1 l
Activit
Bedded away At kill Not
from den site Stalking found
3 1 3 2 2 4
8 1 3 --1 5
8 1 l 8
19 2 0 7 0 0 3 3 0 17
.~ .. n l 1
in Game
Total
38
38
38
114
J
J
D
J
oo·
j'
0
0
D
J
0
D
u--
q .. · :-u-
J
iJ
D
During 1977, we located the three Mendeltna wolves (#007,
008 and 083) on 219 (97%) of 226 occasions on which we searched
for them. During these 219 observations they were resting at
the den or rendezvous site on 129 (59%) occasions. Wolf 007,
the dam of one litter of pups, was responsible for 57 percent of
the den-rendezvous site radio-locations. The two adult males
were located at these sites on 38 percent of the searches. When
these wolves were either stalking or at a kill site, they were
. (ilene. on only 12 (38%) of :32 occasions. We examined the
association of male wolves with their kill activities and found
that whenever a major kill (calf moose or larger) was made, wolf
083, (believed to be the alpha male), was present alone or in
the company of other pack members.
Activity data for the Hogan Hill pack (Table 59} in 1978
revealed a slightly different pattern than that show-n for the
Mendeltna pack. This may have been due, in part, to the fact
that the alpha female of the Hogan Hill pack was not
radio-collared. Of the 114 occasions we attempted to locate the
three radio-collared Hogan Hill wolves (#202, 205 and 206) we
were only able to locate them on 97 ( 85%) occasions. Of those
97 locations the three wolves were located at the den or
rendezvous site on 44 (45%) occasions. These data suggest that
the Hogan Hill wolves were more solitary than the Mendel tna
wolves when stalking prey or visiting kills. On 10 (77%) of 13
of these occasions the wolves were observed alone.
Nevertheless, examination of these data, in relation to kills
made, indicates that the adult male ( #202) was present on every
new kill of calf moose or larger prey. When other
radio-collared pack members were observed alone at kills the
prey was either a small unidentified kill or a kill previously
made by wolf 202 and his associates.
Activity data collected during summer 1977 and 1978 support
the conclusion of Theberge et al. (1978) that adult wolf scats
collected at den and rendezvous sites are predominantly those of
the lactating female. These wolves spend more time at these
sites than do other pack members at least until the pups are
weaned and moved to rendezvous sites.
Although wolf pack members are more solitary in their
hunting habits during summer than in winter, our findings
demonstrate the importance of adult males (probably alpha males)
in food gathering activities. Adult males were involved in
nearly all, if not all, of the major kills made by the pack.
Other pack members, particularly yearling wolves, often appeared
to be hunting, but rarely were they successful on their own;
when they were, the kill was usually a small game species. We
recommend that if summer predation data.are to be collected, an
attempt be made to radio-collar all adult members of each study
pack.
1 2 5
GMU 13 Wolf Densities
Spring and fall wolf densities within a portion of GMU 13
were determined from spring 1975 through spring 1980 (Table 60).
Density estimates were based on only those wolf packs for which
accurate pack numbers were available within the Nelchina study
area. Because radio contact with some packs was intermittent,
the number of packs from which estimates were derived varied by
season. These estimates differ slightly from earlier
preliminary estimates given by Stephenson (1978) and Ballard and
Spraker (1979) because for this report we used those packs with
which we had the most contact during the study period. We
believe this method provides a better comparison of the annual
fluctuations in density occurring in the Unit.
Wolf densities in GMU 13, ex~uding the2 susitna River study
area, varied f~om 1 wolf/!7 .6 mi (97 .3 km ) in fall 1975 to
1 wolf/121.7 mi (315.2 km ) in spring 1978. Spring densities
reflected losses due to trapping and hunting, dispersal and
natural mortality, while fall densities reflected subsequent pup
production and survival and immigration. Both spring and fall
densities showed a progressive decline from spring 1975 through
spring 1978. The decline could be attributed primarily to
hunting and trapping mortality, although natural mortality and
dispersal, particularly for the St. Anne pack, also accounted
for some of these losses.
From fall 1978 through spring 1980, wolf densities in
portions of the Nelchina Study Area increased. The increase was
the result of complete or partial closures of some study wolf
pack areas to hunting and trapping. Boundaries of the closed
areas were described by Eide (1979) and Tobey (1980).
Wolf densities in the Nelchina Study Area from spring 1975
through spring 1980 were used to derive wolf population
estimates for GMU 13 (Table 60). Two sets of estimates were
made, one for all of Unit 13 excluding the Susitna River study
area and the other for wolf habitat only, where glaciers and all
areas lying above 4,000 ft (1,219 m) elevation were excluded.
We believe the second method provides a more realistic estimate
of wolf numbers, since in this region neither wolves nor their
prey regularly inhabit areas above 4,000 ft elevation. The main
potential problem with this method was that some Dall sheep
habitat is excluded from the estimate. Since our observations
of study packs where sheep occur indicate that this prey species
constitutes a very small percentage of the diet, the resulting
error is small. Based upon this latter method the numbers of
wolves in GMU 13 during this study have· fluctuated from 390
wolves in fall 1975 to a low of 121 wolves in spring 1978.
Because the 1979 and 1980 density estimates were partially based
upon wolf numbers in protected areas, the resulting estimates
were probably inflated and the actual population was undoubtedly
lower than reported.
1 2 s
l u
J
L
L..
LJ
L
=
L
-·_r
~~ ,_,
L..J r ··.·· 1
., ~
Table 60. Spring and fall wolf numbers from selected study packs utilized to calculate wolf density and population estimates for GMU-13 in Southcentral Alaska
from spring 1975 through spring 1980.
Spring E"all Spring Fall Spring Fall Spring Fall Spring Fall Spring
Pack Name 1975 1975 1976 1976 1977 1977 1978 1978 1979 1979 1980
Deep Lake ? ? 8 5 2 8 3 3 1-2 2 ?
Ewan 7 ll 5? 5 3 5 3 3? 4 7 ?
Hogan Hill 7 9 7 5 5 8 8 12 1? ? ?
Keg Creek 7 l3 5 8 1 l 2 8 ? ? ?
Mendeltna ? ? 5 9 7 15 0
Middle Fork ? ll 9 9 3 3 0 1 ? 5 ?
Saint Anne ? 11 8 8 8 14 8 7 4 8 2-3
Sinona 7 ll ll 20 2 8 4 10 1-2 1-2 4
Susitna ? ? ? ? ? ? 2 10 7 13 4
Tolsona ? ? ? 4? ? 7+ 3 10 6-7 16 9
Tyone ? ? 6 6 7-8 12 1 2 2 9 4
Subtotal 28 66 64 79 39 81 34 66 29 62 24
II lone wolves (10%) 3 7 6 8 4 8 3 7 3 6 2
Total 31 73 70 87 43 89 37 73 32 68 26
Sample area 2 1932 2743 3818 4281 3818 4281 4503 4302 3455 3957 2266
Sq. mi. (km ) (5004) (7104) (9889) (11088) (9889) (11088) (11663) (lll42) (8948) (10249) (5869)
Sq. mi.
(km)/wolf 62.3(161.4) 37.6(97 .3) 54.5 ( 141. 3) 49.2(127.4) 88.8(230.0) 48.1(124.6) 121. 7(315.2) 58.9(152.6) 108.0(279.6) 58.2(150.7) 87 .2(225. 7)
Population2?stimate
for GMU 1~-=
20,978 mi
194 (54,333 km 2) 337 558 385 426 236 436 172 356 360 241
population estimate
for wo~f habitat i2
GMU 13-1~,666 mi
235 390 269 298 165 305 121 249 136 151 168 (37,985 km)
1/ Keg Creek pack re~uced by ex~erimental removal. ~I Excludes 2,800 mi (7,252 km) wolf removal area.
Tables 61 through 64 summarize wolf sightings made by the
public and Department personnel from 1976 through 1980. Based
upon these observations and our radio-telemetry data we had
estimated that from 1975 to 1978 between 40 and 50 wolf packs
inhabited GMU 13 (Ballard and Spraker 1979). We estimated that
by June 1980 between 20 and 30 packs inhabited the Unit.
GMU 13 wolf densities were compared with those reported
elsewhere in North America (Table 65).2 In Alaska, reported wolf
densities range from 1 wolf/25 mi 2 in southeast Alaska
(Atwell et al. 1963) to 1 wolf/124 mi in the Brooks Range
(Stephenson 1975). Unit 13 densities fall in the middle of this
range but in 1975 the density was close to that reported on the
Kenai Peninsula, Alaska (Peterson 1980). Unit 13 densities fall
roughly in the middle of densities reported in North America and
appear quite similar to densities reported in other areas of
southcentral Alaska (Murie 1944; Rausch 1967).
Trapping and Hunting Mortality
Table 66 summarizes wolf harvests for GMU 13 from the
1971-72 through 1979-80 seasons. During this period annual
harvests ranged from 57 to 128 wolves. Method of harvest was
classified into four categories: (1) trapped (snared or caught
by leg-hold trap), {2) ground shot {which usually involves
landing aircraft and shooting same day airborne)
(3) experimental removal by ADF&G personnel, which involves
shooting from helicopter or fixed-wing aircraft and, (4) other,
which includes miscellaneous forms of mortality such as
automobile collisions and natural mortality. Aerial hunting was
legal only in 1971-72. Any subsequent illegal aerial harvest
was lumped with ground shooting.
From 1972 through 1975-76, trapping was the most com.111on
method of harvesting wolves in GMU 13, accounting for 59 percent
of the total harvest. Beginning with the 1976-77 season,
however, ground shooting became the most common method of
harvest, accounting for 52 percent.
Harvests for 1977-78 were the highest recorded for GMU 13
since 1968. A portion of the increase was the result of the
Department experimental removal program. Most of the increase
could be attributed to the increased efficiency of two or three
aerial trappers; from 1916 through 1980 their harvest ranged
from 12.2 percent to 60.9 percent of the total GWJ 13 harvest.
Figs. 6 through 9 depict the approximate distribution of the
wolf harvests for both GMU 13 and adjacent GMU 11 from 1976-77
through 1979-80. In most cases, the reported kills and public
observations of numbers of wolves per pack corresponded with our
observations of declines in numbers of wolves in radio-collared
packs, indicating that information provided from hunters,
trappers, and the general public was generally accurate.
1 2 8
fl,
L
L
L
F
L
L
r
r
L
L
r
L
L
t __
b
L
J
J
J
J~
[f
J
D
J
J
}.
J;'
D
J
D
J
Table 61. Summary of reported Department and public wolf observations within the Nelchina,
Susitna and Copper River Basins from 1 July 1976 through 30 June 1977.
Date of
observations
7/11/76
7/16/76
8/26/76
8/29/76
8/31/76
9/1/76
9/76
9/76
9/5/76
9/5/76
9/5/76
9/6/76
9/10/76
9/15/76
9/18/76
9/26/76
l/8/76
10/8/76
10/8/76
10/17/76
10/21/76
10/21/76
10/21/76
10/21/76
10/23/76
10/23/76
10/29/76
10/28/76
10/30/76
11/2/76
ll/3/76
ll/5/76
11/5/76
11/5/76
il/5/76
11/8/76
11/3/76
11/3/76
11/13/76
Number and type
of observations
l wolf
1 wolf,
2 wolves (black-gray)
2 wolves (white)
1 wolf (gray)
1 wolf ("dark")
7 wolves (2 black-S Gray)
5 wolves (gray)
2 wolves (1 black-1 gray)
1 wolf (white)
2 wolves ·(black-gray)
12 wolves (1/2 blk-l/2 gray)
10-15 wolves (hunter got 2-3)
l wolf
S-6 (howls)
3 wolves (2 black-1 gray)
. 3 tracks
6 tracks
16 wolves
l wolf (gray)
2-4 tracks
2 tracks
2-3 tracks
4-6 tracks
2 tracks
l track
1 wolf (gray)
9 tracks
3 tracks
3 tracks (l wolf taken)
1 track on caribou kill
6 wolves (4 gray-2 black)
2 wolves (gray-black)
2 wolves (gray)
3-4 tracks
1 wolf (gray)
1 wolf (gray)
1 wolf (gray)
5 tracks
Location Source
Clarence Lake ?
Dump at Glenn-Rich Russel
15 mile Denali Highway Gardner
Sikonsina Pass McMahan
Athna Lodge Potterville
Mile 34 Denali Highway Johnson
Between Tazlina Lake and Klutina Ronning
Stuver Creek ? (Ellis)
Opposite Tyone River on west Hunter to ADF&G
side Sue
Chistochina
So. fork of Coal Creek
Head of Daisy Creek
Between Monsoon Lk. & Maclaren
Chis to china
?
Hunter to ADF&G
M. Haggstran
Hunter to ADF&G
?
Between Boulder and Drop Creek ADF&G
N. Roundtop Mtn. on gut pile
Between Goose Creek and Oshetna
Between Sanona-Oshetna
White River
Tolsona Lake
Big bend of Maclaren
On Denali between lodge &
Clearwater
McMahan
ADF&G
ADF&G
Vaden
Roberson
ADF&G
ADF&G
East Fork of Sue ADF&G
7 miles above hwy. to 15 below ADF&G
Sue River down to bridge
Between lodge and Clearwater
Kelly Lake
Dickey Lake
West of Gulkana on pipe
Warm Sp. Camp
Marie Lake
3 miles west of K.C. den on
West Fork
Across from Maclaren Lake
2 miles N. of Tyone Butte
on Tyone River
5-6 mi. below M. Brushkana
3-4 miles from H. Hill
Island Lake on caribou kill
SW of Island Lake on caribou
kill
W. Fork to Sourdough
McMahan
McMahan
ADF&G
McMahan
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
1 2 9
Table 61 (cont.)
Date of
observations
11/13/76
11/17/76
11/14/76
11/23/76
1/2/76
10/76 or
12/27/76
12/6/76
12/18/76
12/23/76
l/5/77
l/5/77
l/10/77
1/10/77
l/10/77
l/21/77
l/21/77.
l/24/77
Number and type
of observations
3 tracks
Fresh caribou kill
2 wolves (gray 1 small
female or pup
4 tracks (2A-2 pup)
2 tracks
6 wolves
6-8 wolves
2 tracks
1 wolf (gray)
1 wolf (gray)
1 wolf (gray)
3-4 tracks
7 tracks
4-5 tracks
l track
6-8 tracks
3 tracks (one trapped and
lost)
Location
Little Tonsina
Lone Butte Lake
Butte Creek
2 mi. S. of Lake Louise
On pipe near Sourdough
Sanford River
Lower Tonsina
Chistochina, 2-3 mi. from road
Sheep Mountain
Round Lake
N.E. of Round Lake
S. of Ewan Lake & Glenn Hwy.
W. Fork Gulkana
Klutina Lake
Between Sue Lodge-Valdez Creek
lower Tyone Creek
Fish Lake
l/24/77
l/24/77
3/18/77
3/24/77
4-5 tracks (one trapped and Fish Lake
3/77
Winter
4/5/77
4/8/77
4/8/77
4/19/77
4/19/77
4/25/77
5/10/77
5/10/77
5/20/77
lost)
l wolf (black)
15 tracks on moose kill
12 wolves (gray) on 2
moose kills
3-5 tracks
6-7 tracks
6 wolves on moose kill
3 wolves (gray)
4 wolves
1+ tracks on caribou kill
4 tracks on caribou kill
3 tracks on moose kill
4 wolves (gray)
1+ wolf (gray) on collared
moose kill
1 wolf (gray)
Simpson Hill
Tazlina River
Mankomen Lake
East Fork of Susitna
Indian River
Woods Canyon on Copper River
Ewan Lake
Ewan Lake
South of Lake Louise
Copper Lake, Unit 11
Natat Creek on Slana, Unit 11
5 miles west Mankomen Lake
West of the Copper River near
Gulkana Airport
West of Sheep Mountain
Source
McLaughlin
ADF&G
L. Steele
L. Steele
ADF&G
ADF&G
ADF.&G
Steele
Schmidt
Steele
Steele
Hunt
Hunt
Hunt
Shorty
ADF&G
D. Hansen
D. Hansen
E. McKenzie
H. Billum
C. Farnham
Hardy
C. Farnham
A. Fejes
ADF&G
J. Smolen
ADF&G
ADF&G
ADF&G
ADF&G
K. Bunch
ADF&G
1 3 G
,.=.-..---
L
,.
r
l._
r
i_
.,-
L
r
L
r
L
c
L.
r
L
L
L
. -.. r
L
L
L
L
l u
J
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Table 62. Summary of reported Department and public wolf observations within the Nelchina,
Susitna and Copper River Basins from l July 1977 through 30 June 1978.
Date of
observations
8/ll/77
9/l-9/20/77
9/l/77
9/l/77
9/7/77
9/26/77
10/20/77
10/24/77
10/27/77
10/28/77
10/28/77
10/28/77
10/30/77
10/31/77
10/31/77
10/31/77
10/31/77
ll/2/77
ll/3/77
ll/3/77
11/3/77
11/7/77
11/9/77
11/11/77
11/15/77
11/17/77
ll/22/77.
Early winter
Early winter
Early winter
Early winter
Early winter
Number and type
of observations
4-6 tracks
2 wolves
1 wolf (gray)
l wolf (black)
7 tracks
7 wolves (all gray)
l track
7 wolves
20 approx. wolves?
1 wolf (black)
2 wolves (gray)
2 wolves (gray)(harvested)
1 track
14 wolves on moose kill
6 tracks
5 tracks
2-3 tracks
3-4 tracks
l+ tracks on yearling moose
kill with bear and cub
l track
5 tracks
3 tracks
5 tracks
2 tracks
2-5 tracks
1+ tracks on caribou kill
3-4 tracks
4 wolves (all grays)
2-4 tracks on moose kill
2 tracks
3 tracks
5 tracks
5 wolves (all gray)
4-5 tracks
1 wolf
3-5 tracks
3-5 tracks
6-7 wolves
7 wolves
18 wolves
Location Source
Dan Creek
Lone Butte
R. Johnson
ADF&G hunter
check station
Tanawa Lake
Little Oshetna River
Confluence Tyone Creek-River
Goose Creek and Susitna River
Middle Fork Chistochina River
Little Nelchina River
Head of Chistochina River
II
II
II
II
II
II
II
II
II
II
II
II
II
It
II
II 8-10 miles N. of Eureka
Swiss Lake
West Fork Susitna River
Lower Clearwater Creek
W. bank Big Nelchina River
Trappers Den Lake
Hunter to R. Halfor(
Hunter to ADF&l
Hunter to ADF&l
Northright
Tyone River above village
3 miles S. Sourdough
Upper Gakona River
Upper Gakona River
Mouth Windy Creek
l mile N. Sourdough
Minnesota Lake
Tyone River '
Across from Tyone River
East Fork Watana Creek
South side Ewan Lake
Mouth Oshetna River
On Susitna River between Jay
and Watana Creek
5 miles So. of confluence of
Maclaren and Susitna R. on Sue
5 miles So. of confluence of
Maclaren and Susitna R. on Sue
R. Carter
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
On pipeline at Roundtop Mtn ADF&G
Meiers Lake ADF&G
On Lake Louise Road J. Dimarco
Between Sourdough and Glennallen ADF&G
On pipeline W. of Gulkana Airport R. Armstrong
Mae West Lake H. Billum
Ewan Lake H. Billum
Drop Creek to Boulder Creek McMahan
Boulder Creek to Sanford River McMahan
Sanford River to Nadina River McMahan
.1 3 1
Table 62 (cont.)
Date of
observations
Early winter
Early winter
Early winter
Early winter
Early winter
l/10/78
1/22/78
l/24/78
l/27 /78
l/27 /78
l/27 /78
2/9/78
2/9/78
2/9/78
2/ll/78
2/ll/78
2/ll/78
3/7/78
3/7/78
3/78
4/6/78
4/6/78
Number and type
of observations
12 wolves
5•6 wolves
12+ tracks
4 tracks
5 tracks
6 tracks (2 harvested)
8-9 tracks on adult and
calf moose kill
7 tracks
4-6 tracks
2-4 tracks on moose kill
4 wolves (all grays)
3-4 tracks on 2 adult moose
kills
· • 5 ·old tracks
l-2 tracks
3 wolves (all grays) on
adult moose kill
l wolf (gray) and 2 tracks
1-2 track
2 tracks
4-5 tracks
2 wolves
5 tracks
2 tracks
Location Source
Nadina River Chesnina River McMahan
Chesnina River to Kuskulana River McMahan
Gold Creek (Susitna) ADF&G
Upper Talkeetna River ADF&G
Indian River E. Ross
Between Butte Creek and K. Bunch
Deadman Lake
Mouth of Gulkana River ADF&G
Matanuska Glacier
Head of Moose Creek
Maclaren River Bend
Lower Watana Creek
On Susitna River near Jay Creek
Moose Creek (Alphabets)
Maclaren River cabin
Jay Creek and Susitna River
3 miles up Watana Creek
Kelley
15 miles So. Susitna River Br.
W. Fork Gulkana River
Mile 10 Denali Highway
W. Fork Gulkana River
3 miles So. of Birch Lake
R. Giger
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
J. Wilson
J. Wilson
C. Gardner
ADF&G
ADF&G
1 3 2
'l
!
LJ
J
n c I
L
L
L
L.
L
L
~
L
L
-
L
L
J
~ d
J
1
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D
d
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];"
D
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0
D
Table 63. Summary of Department and public wolf sightings 1 July 1978
through 30 June 1979 in the Nelchina, Susitna and Copper River
Basins of southcentral Alaska.
Date
8/17/78
8/20/78
8/23/78
8/23/78
9/78
9/78
9/78
9/78
9/78
9/2/78
9/2/78
9/1/78
9/1/78
9/1/78
9/1/78
9/3/78
9/5/78
9/6/78
9/10/78
9/13/78
9/15/78
10/78
11/14/78
11/14/78
11/15/78
11/15/78
11/15/78
12/78
12/15/78
12/15/78
12/15/78
12/16/78
12/21/78
1/2/79
4/2/79
4/2/79
5/9/79
1 wolf
Z wolves
1 wolf
10 wolves
1 black-1 gray
17 wolves
6-7 wolves
Tracks of 1
2 wolves
1 black
1 black-1 gray
1 set tracks
1 wolf track
11 _wolves
1 gray
1 white
1 black
1 wolf
Tracks of 2
2 gray
Tracks
1 wolf
4 wolves
Tracks 6-8
Tracks of 3
Tracks 2-4
Tracks of 2?
12 wolves
Pack of 9-10, 3 taken
Pack of 11, 4 left
Pack of 8 or 9
Tracks of 7
Tracks 3-4 So.
1 black
Tracks of 6
1 track
1 track
Location
Crater Lake
Nabesna River
Clarence Lake
Capital Mtn.-Mt. Sanford
Roundtop Mountain
Bremner
Daisy Creek
West Fork Susitna
Monahan Flats
Twelvemile Creek
Upper Chistochina River
Big Oshetna River
West Fork Gulkana
Upper Slana
Upper Kuskalana River
4 mi east Keg Creek
Upper Twin Lakes
Tyone River
Susitna River-Fog Lakes
Bone Creek on Slana
E. Fork Chistochina River
Tons ina
Mankomen Lake
East of Hogan Hill
Watana-Butte Creek
Tyone-Jay Creek
Upper Clearwater Creek
Boulder Creek-Copper
Sinona Creek
Fox Lake
Sanford River
Lower Gulkana
Highway-Little Nelchina River
Richardson Highway
Ewan to Crosswinds
Little Tulsona Creek
Gulkana
Sightings by
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
Public
ADF&G
ADF&G
ADF&G
ADF&G
ADF&G
Public
Public
Public
Public
Public
ADF&G
Public
Public
ADF&G
ADF&G
1 3 3
l
Li
Table 64. Summary of Department and public wolf sightings 1 July 1978 J through 30 June 1979 in the Nelchina, Susitna and Copper River
Basins of southcentral Alaska.
Date Location Sightings by
9/79 5 grays W. Fork Maclaren River Public
11/79 6 wolves Dadina River Public J
12/79 Tracks 2-3 Chistochina River Public
12/79 5 wolves Boulder Creek-Copper River Public
12/1/79 2 wolves tracks of 5 South end Paxson Lake Public
12/9/79 1 wolf pup Between Richardson & Gakona River Public
12/19/79 Tracks 3-4 South of Lake Louise Public
1/5/80 5 wolves Mouth Maclaren River Public
1/6/80 Track 2-3 Gakona River ADF&G
1/17/80 1 gray Tolsona Public
1/20/80 20 wolves-7 caught Nabesna Glacier Public
2/80 Tracks of 2+ Barnet Creek Public
2/10/80 7 grays West of Fish Lake ADF&G
2/18/80 Tracks of 10, 4 taken Mouth Maclaren River Public
2/18/80 4 wolves Northeast of Fish Lake Public
2/18/80 Tracks 4-5 Between Gakona and Gulkana River Public L
2/20/80 1 wolf Dog Lake Public
2/25/80 Tracks of 6 5 mi northeast of Bell Lake Public
3/80 7 wolves Mouth Watana Creek Public
3/6/80 4 grays Susitna River opp. Clarence Lake ADF&G
3/12/80 7 wolves Hudson Lake Public
3/13/80 Tracks 4-6 West Fork Gulkana ADF&G
5/22/80 2 blacks East Fork Susitna River Public L
6/80 Tracks of 3 Mouth Gulkana River Public
L
L
L
'
L
L
'·
L
1 3 4
L
1 u
J
J
Dr,
0
D
j
}"
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D
Table 65. Summary of reported wolf densities in North America.
Wolf density
mi 2/wolf
31:./
4.62/
6.9-
7-10 7.8-13.8~/
9.2
10
10
10.6
12.9-55.2
17
24-42
25-40
25-29
35
37.6-121.7
40-83
50
50
58
2 km /wolf
. 7.8
11.9
17.9
18-26
20.2-35.7
23.8
26
26
27.5
33.3-142.9
44
62-109
65-104
65-75
91
97-315.2
104-215
130
130
151
60-120 155-311
65-124 3/ 168-321
87-111(10-) 225-287
88 228
120 311
200 518
Size of study area
.2 ml.
30
210
384
210
1,274
717
1,000
2,490
4,203
121-998
4,100
1,500
7,500
1150-1845
7,000
1932-4503
2,000
20,000
9,653
48,0000
3,600
4,200
593
1,800
109,000
78
544
995
544
3,300
1,857
2,590
6,449
10,886
313-2584
10,619
3,885
19,425
2979-4779
18,130
5004-11663
5,180
51,800
25,000
1,243,200
9,324
10,878
1,536
4,662
282,310
Location of study area
Coronation Isl., Alaska
Isle Royale, Michigan
NW Territories
Isle Royale, Michigan
Manitoba-Saskatchewan
Minnesota
Algonquin Park, Ontario
Minnesota
Minnesota
Beltram, Isl. St. For.,
Minnesota
Minnesota
Mt. McKinley Nat. Park
Alaska
SE Alaska
Kenai Peninsula, Alaska
Tanana Flats, Alaska
Nelchina Basin, Alaska
Saskatchewan
NcKinley Nat. Park, Alaska
Nelchina Basin Study area,
Alaska
NE Alberta
NW Territories
NC Brooks Range, Alaska
W. Canada
W. Canada
Baffin Isl., Canada
Manitoba-Sasketchewan-
NW Territories
1/
2;
~I
Artificial situation -four wolves transplanted to Island.
Wolves .concentrated on caribou winter range.
Maximum abundance on winter range.
Source
Merriam 1964
Peterson 1976
Kuyt 1972
Mech 1966
Parker 1973
VanBallenberghe
et al. 1975
Pimlott et al.
1969
Olson 1938
Mech 1973
Frttts & Mech
In Press
Stenlund 1955
Haber 1968
Atwell et al.
1963
Peterson and
Wolvington 1978
Stephenson 1977
This study
Banfield 1951
Murie 1944
Rausch 1967
Fuller & Keith
1980
Kelsall 1957
Stephenson 197~
Cowan 1947
Carbyn 1974
Clark 1971
Parker 1973
1 3 5
1 3 5
L
Figure 6. Diagram of hunting-trapping location of wolf harvests in GMU-11 and 13 of Southcentral Alaska
for 1976-77 regulatory year~ ..
f
w
00
r· '"'l
Figure 7. Diagram of hunting-trapping location of wolf harvests in GMU-11 and 13 of Southcentral Alaska for
1977-78 regulatory year.
~ ~;~~~v~~R§~~
r
. t ,,, '1 r ',1 r· I 1 T ' ~ l f''" 1 r,--·n r I l [, _ _]
c:J CLJ c..J c::J
Figure 8.
CJ LJ r...LJ E....J c......J LJ t..:..J c..::..J c.J c.:...J c:..:..::J [__J '·• . j ·. r--' )'I
of hunting-trapping location of wolf harvests in Game Management Units 11 and 13 of
Alaska for 1978-1979 regulatory year.
l..l.J L.....J [_J
Figure 9.
C.J
r , ,. ,., r·
Diagram of hunting-trapping location of wolf harvests in Game Management Units ll and 13 of
Southcentral Alaska for 1979-1980 regulatory year. 1
..
1 r r .. rl' n 1 '1 'l r '
I
.,.
J L..J [. _ _j
l
;_j
n
J
J~
n{
0
J
]·.
J ... .
.
D
J
Mech (1970) concluded, based upon data from North America,
that wolves can compensate for annual losses of 50 percent and
possibly more to animals aged 5-10 months or older. He further
he suggested that, in order to control wolf numbers, at least 50
percent of the animals of this age must be killed each year. We
compared harvest levels of various radio-collared wolf packs
with subsequent pack numbers in an effort to determine the
effects of various human harvest levels on GMU 13 wolves (Fig.
10}. In this analysis, both dispersal and natural mortality
were included and, thus, the comparison is not related to just
human induced mortality. Nevertheless, because most pack losses
were hunting and trapping related it provides a guideline for
harvest strategy. Harvest level (% loss) and subsequent fall
pack numbers were significantly correlated (P<O.Ol), indicating
that this wolf population was being controlled by human harvest.
Wolf packs remained stable or increased at harvest rates of 22
to 60 percent but declined at harvest levels of 33 to 92 percent
of the preceeding fall's population. Based upon the equation
given in Fig. 10, a harvest level of 41 percent would, in most
cases, allow these wolf packs to remain stable. This would not
always be the. case however, because in this heavily exploited
population, packs are often comprised of two to three adults and
five to six pups. If annual harvests remove one or two of the
adult members, the pack will probably not reproduce even if only
30-40 percent were harvested. Also, the 41 percent figure
includes dispersal and natural mortality so the actual level of
human harvest that would usually stabilize pack numbers probably
lies between 30 and 40 percent.
Hunting and trapping closures in portions of the Nelchina
study area in 1978 and 1979 resulted in population increases in
the closed areas. Since these packs were used in estimating the
Unit's population they inflated this estimate and gave the false
impression that the Unit population had increased. Since 1978,
the number of public sightings of wolves appears to have
declined (Tables 61 through 64). Examination of Figs. 6 through
9 indicates that in many cases packs are being harvested well in
excess of 41 percent.
The GMU 13 wolf population has recovered from low densities
when restrictive hunting and trapping regulations have bee~
implemented. The Unit wolf population grew from 12 to 300
individuals in 17 years when given relief from human harvest
(Rausch 1967}. An equally impressive rate of increase occurred
during this study when portions of the Nelchina study area were
closed to hunting and trapping over a 2. 5-year period .
Management of GMU 13 wolves has been guided by policies which
state that the Department will manage the resource on the basis
of: (a) maximum overall recreational opportunity, (b) maximum
sustained harvest, and (c) maximum aesthetic appeal to the user
(A.D.F.&G. 1963). A series of hunting-trapping season openings
and closures allowing great increases and decreases in wolf
numbers may not be consistent with this policy. Alaska's game
managers need to determi~e what constitutes a desirable wolf
1 4 1
lv
Figure 10 .. Relationship of percent annual reduction per pack to subsequent wolf numbers per pack
as expressed by previous fall populations for radio-collared wolf packs studied from
April 1975 through June 1980 in Game Management Unit 13 of southcentral Alaska.
100
)(
90-X
80
::.4 70 ~ p..
rz..
....:1 60 ~ -6
H 50 H u
:;:l
Q
~ 40 H
f;i
~
J:il 30 p..
20
10
X
20 40 60 80 100 120 140 160 180 200 220 240 260 280 300
FALL WOLF NUMBERS/PACK (Percent of previous fall's population)
.. . r· ·l r : 1 r · 1 [ _ _]
1 t.J
J
0
0
1_·_----~-J
o~
0
J
d
~ J .
•. J ..
D
J
D .
J
density which does .not create adverse predation on moose and
caribou populations which also are to be managed in most cases
on a maximum sustained yield basis. To date no such guidelines
have been proposed.
Based upon the downward trend exhibited by this wolf
population under the reported harvest levels, we propose that
management strive to maint~in a post harvest wolf density of
approximately 1 wolf/100 mi of habitat, a population of -150
wolves in GMU 13. To achieve this objective wit..'lout unduly
restricting wolf harvests some minor restrictions on hunting and
trapping may be necessary. Examination of wolf sealing
documents from 1976-80 reveals that successful ground trappers
took an average of 1.5 wolves/season (range of 1 to 6). Aerial
trappers, however, took an aver age of 3 . 4 wolves per season
(range of 1 to 23). The upper limit of wolf harvest per trapper
could be reduced, preventing further wolf population declines.
Wolf numbers and numbers of trappers should continue to be
monitored to maintain desired wolf densities.
Table 67 lists the fate of the 103 wolves radio-collared
from April 1975 through June 1980. Of that total, 14 (14%) were
known to be alive on 30 June 1980 while 26 (25%) were known to
have dispersed and were either dead or missing by the same date.
Excluding experimental removal ( 12) and collaring mortality ( 1
plus 1 not listed), the leading known cause of mortality was
human induced (37 of 48, 77%). Ground shooting which involves
the use of aircraft, and suspected illegal aerial hunting,
accounted for 76 and at least 11 percent, respectively, of L~e
human-caused mortality. Four (11%) wolves were trapped and one
( 3%) was hit by an auto. Natural forms of mortality (see
Appendix V) accounted for 23 percent of the total known wolf
mortality and were attributable to the following: 4 killed by
other wolves, l stepped on by a moose and 5 due to miscellaneous
factors such as bear predation, drowning or unknown causes.
Illegal aerial hunting was probably underrepresented; we
observed buckshot holes in several wolf hides and carcasses
presented for sealing which were purportedly taken by ground
shooting.
During this study we lost radio contact with 16 of the 103
radio-collared wolves (16%). We suspect that most of the
missing radio-collared wolves were either: (1) taken illegally,
( 2) if taken legally reported as taken elsewhere, ( 3) properly
reported, but no mention made of the radio-collar, ( 4) lost due
to radio failure, or (5) dispersed. Since 26 percent of all
radio-collared wolves were known to have dispersed, undoubtedly
some of the missing radio-collared wolves are in this category.
Toward the latter portion of this study we began receiving
unconfirmed reports that radio-collared wolves were being taken
but that the radio-collars were being destroyed and not
reported. We are certain that some of the rumors were true and
some radio-collars were destroyed. Since these animals
represent a large financia.l investment by the State and because
1 4 3
Table 67. Summary of the status of all radio-collared wolves in GMU-13 from 1975 through June 1980.
Month
Date Number contact or Status
Original suspected Accession radio-radio-mortality as of Cause of
pack affiliation number collared Age Color Sex Weight locations occurred 6/30/80 mortality
Brushkana 121983 04/20/75 Adult Black ·~ 90 31 1/76 Dead Exp. removal
Butte Lake 121984 04/20/75 Adult Black cJ 102 23 1/76 Dead Exp. removal
Deadman 121981 04/20/75 Adult Gray cJ 92 35 3/76 Dead Exp. removal
121982 04/20/75 Adult Gray ~ 108 28 3/76 Dead Exp. removal
Deep Lake 122067 03/22/76 Adult Gray cJ 102 46 3/77 Dead .Exp. removal
121993 03/22/76 Adult Gray ~ 94 58 10/76 Missing Prob. dispersed
122009 10/29/76 Adult Gray ~ 86 87 2/79 Dead Killed by
Susitna wolves
11/13/77 90
11/19/78 92
122096 11/13/77 Adult Gray cJ 110 11 1/78 Dead Ground shot
01/05/78
122204 04/10/78 Pup Gray cJ 99 18 7/78 Missing Dispersed
122212 11/19/78 Adult Black cJ 92 10 2/80 Dead Ground shot
Delta 121997 06/06/75 Adult Gray cJ 98 21 11/76 Dead Nat. mortality-
cause unknown
121998 06/06/75 Pup Gray cJ so Unk Dead Trapped
122063 03/21/77 Adult Gray cJ 72 4 3/79 Missing
122064 03/21/77 Adult Gray ~ 104 2 4/77 Dead Exp. removal
in Unit 20A
Ewan 121990 04/15/75 Adult Gray ~ 87 148 10/76 Missing ? dispersed
121991 11/04/75 Adult Gray cJ 85 11 1/76 Dead Killed by St. Anne
wolves
121992 11/04/75 Pup Gray cJ 55 17 12/75 Dead Trapped
122151 03/16/78 Pup Gray ~ 65 6 3/78 Dead Killed by moose
122219 11/17/78 Pup Gray cJ 52 4 3/79 Dead Trapped
03/15/79
Gakona 121999 03/18/76 Adult Gray cJ 110 2 8/76 Dead Auto collision
122217 11/16/78 Adult Gray 9 100 3 12/78 Dead Ground shot
122218 11/16/78 Adult Black cJ 6 2/79 Dead Ground shot
):-.
p..
Table 67 (cont.). Summary of the status of all radio-collared wolves in GMU-13 from 1975 through June 1980.
Month
Date Number contact or Status
Original suspected Accession radio-radio-mortality as of Cause of
pack affiliation number collared Age Color Sex Weight locations occurred 6/30/80 mortality
Hogan Hill 121987 06/06/75 Yrlg. Gray !j? 69 104 1/79 Dead Ground shot
121988 11/05/75 Adult Gray Cf 88 62 4/77 Missing Dispersed
122202 04/08/78 Adult Gray Cf 108 52 1/79 Dead Ground shot
122205 04/09/78 Pup Gray Cf 82 51 1/79 Dead Ground shot
122206 04/09/78 Pup Gray !j? 79 37 9/78 Missing ? Prob. dispersed.
Keg Creek 121986 04/20/75 Adult Black Cf 100 101 3/76 Possibly Illegal aerial
dead hunting
122083 04/20/75 Adult Gray Cf 94 191 2/78 Dead Ground shot
04/27/77
122002 04/20/75 Adult Black !j? 82 147 7/76 Dead Exp. removal
122068 03/21/76 Adult Black !j? 93 74 3/77 Dead Exp. removal
122201 03/21/76 Yrlg. Gray !j? 74 71 2/79 Suspect Illegal aerial
dead shooting
04/08/78 Adult 96
122010 10/29/76 Pup Gray !j? 54 23 3/77 Dead Exp. removal
122011 10/29/76 Pup Gray Cf 52 24 3/77 Dead Exp. removal
122203 04/09/78 Adult Gray Cf 104 53 2/79 Suspect Illegal aerial
dead shooting
Maclaren 121985 04/21/75 Adult Gray !j? 77 51 1/76 Dead Exp. removal
Mendeltna 122007 10/29/76 Adult Black 9 72 124 2/78 Dead Ground shot
122008 10/29/76 Pup Gray Cf 54 119 11/77 Dead Killed by
Tyone wolves
04/27/77 92
122095 10/13/77 Adult Gray 9 86 13 2/78 Dead Ground shot
122090 10/13/77 Pup Gray d 50 13 2/67 Dead Ground shot
Middle Fork 121996 03/21/76 1.5 Gray !j? 80 2 5/16 Missing ?
122062 03/21/76 Pup Gray 9 63 3 3/77 Dead Collaring
03/21/77 Yrlg. mortality
122078 03/21/76 Pup Gray Cf 66 2 2/77 Dead Ground shot
122061 03/21/77 Pup Gray Cf 68 24 1/78 Dead Ground shot
122085 04/28/77 Pup Gray 9 82 18 2/78 Dead Ground shot
........ 11/10/77 Yrlg . 78
01
Table 67 (cont.). Summary of the status of all radio-collared wolves in GMU-13 from 1975 through June 1980.
Month
Date Number contact or Status
Original suspected Accession radio-radio-mortality as of Cause of
pack affiliation number collared Age Color Sex Weight locations occurred 6/30/80 mortality
St. Anne 122086 04/28/77 Pup Gray (j 91 3 5/77 Missing Dispersed?
122088 04/28/77 Adult Gray (j 100 37 6/78~3/79 Dead Drowned
122087 04/28/77 Pup Gray (j 82 4 5/77 Missing Dispersed
122093 10/11/77 Adult Gray (j 88 8 2/78-9/79 Dead Natural
unknown cause
122094 10/12/77 Adult White ~ 72 53 Alive
122118 02/10/78 Adult Gray (j 92 9 4/78-2/79 Missing Dispersed
122209 02/26/78 Pup Gray ~ 26 4/79-12/79 Dead Starvation
122207 04/10/78 Pup Gray (j 106 10 4/78 Missing ?
122208 04/10/78 Pup Gray ~ 84 9 8/78-9/79 Missing Dispersed
122292 03/31/79 Pup Gray (j 75 13 6/80 Missing ?
122293 03/31/79 Adult Gray (j 92 9 1/80 Missing Dispersed
Sinona 121989 04/20/75 Adult Gray ~ 85 85 5/76 Missing ?
122038 11/05/75 Adult Gray cJ 106 70 2/77 Dead Ground shot
122048 11/05/75 Adult Gray (j 87 108 1/79 Missing
03/14/77
10/11/77
122049 03/14/77 Adult Gray (j 2 3/77 Dead Ground shot
122084 04/28/77 Adult Gray ~ 112 32 12/78 Dead Ground shot
11/10/77 102
11/16/78 98
122213 11/16/78 Pup Gray ~ 60 4 12/78 Probably Suspect ground
dead shot
122214 11/16/78 Pup Gray ~ 60 3 12/78 Dead Ground shot
Stephan Lake 122016 02/19/76 Adult Gray ~ 75 24 10/76 Dead Killed by bear
or wolf
'' r"'"''ll r ·l r ., 1
Table ~7 (cont.). Summary of the status of all radio-collared wolves in GMU-13 from 1975 through June 1980.
Month
Date Number contact or Status
Original suspected Accession radio-radio-mortality as of Cause of
pack affiliation number collared Age Color Sex Weight locations occurred 6/30/80 mortality
Susitna 122227 02/06/79 Pup Gray (j 100 24 4/79 Missing Prob. radio
·failure
122228 02/06/79 Pup Gray (j 99 43 4/79 Missing Prob. radio
failure
122229 02/06/79 Pup Gray (j 94 65 Alive
04/16/80 105
122230 02/06/79 Pup Gray (j 100 47 1/80 Missing ?
122294 03/28/79 Pup Gray 9 80 27 4/79 Missing ?
122295 03/25/79 Adult Gray 9 95 103 Alive
02/20/80
04/15/80
122296 03/25/79 Adult Gray (j 108 53 3/80 Dead Killed by
02/20/80 . wolves
122302 02/20/80 Pup Gray 9 40 Alive
122303 02/20/80 Yrlg. Gray (j 80 10 3/80 Dead Killed by
wolves
122305 04/13/80 Adult Gray (j 100 18 Alive
122306 04/13/80 Pup Gray (j 86 lbs est 18 Alive
Tolsona 122210 06/08/78 Adult Gray (j 104 29 10/79 Missing Dispersed
122220 01/22/79 Pup Black 9 70 lbs est 69 Alive
1222289 03/28/79 Pup Gray 9 26 12/79 Missing Dropped collar
1222290 03/28/79 Pup Gray (j 84 24 10/79 Missing Prob. dispersed?
1222291 03/28/79 Pup Gray 9 70 5 12/79 Dead Trapped
Tsusena 121994 02/20/76 Adult Gray (j 85 4 5/76 Missing ?
122199 02/17/76 Adult Gray 9 80 59 7/78 Missing Prob. alive
03/23/78
122056 03/19/77 Adult Gray (j 92 1 3/77 Missing Suspect ground
shot
122057 03/19/77 Pup Gray 9 72 1 3/77 Missing Suspect ground
shot
Table 67 (cont.). Summary of the status of all radio~collared wolves in GMU-13 from 1975 through June 1980.
Month
Date Number contact or Status
Original suspected Accession radio-· radio-mortality as of Cause of
pack affiliation number collared Age Color Sex Weight locations occurred 6/30/80 mortality
Tyone 122001 02/18/76 Adult Gray cj 104 37 7/76 Dead Exp. removal
121995 02/16/76 Adult Gray ~ 80 15 3/76 Dead Exp. removal
122097 11/15/77 Adult Gray cj 112 5 1/78 Dead Ground shot
122098 11/15/77 Pup Gray cj 92 8 2/78 Dead Illegal areial
hunting
122099 11/15/77 Adult Gray cJ 112 5 1/78 Dead Ground shot
122115 02/10/78 Adult Gray ~ 92 2 2/78 Dead Ground shot
122ll6 02/10/78 Adult Gray cj 112 20 10/78 Mis~ing Dispersed
122117 02/10/78 Adult Gray cj 106 2 2/78 Dead Ground shot
122215 11/18/78 Adult Gray cJ 102 81 3/80 Dead Ground shot
02/20/78 111
122216 11/17/78 Adult Gray ~ 90 68 3/80 Dead Ground shot
02/20/80 95
122298 02/20/80 Pup Gray ~ 84 33 Alive
122299 02/20/80 Pup Gray ~ 82 36 Alive
122300 02/20/80 Pup Gray cJ 93 33 Alive
122301 02/20/80 Pup Gray d 100 14 3/.80 Dead Ground shot
Watana 122197 03/20/78 Adult Gray cJ 5 4/78 Missing ?
122308 04/24/80 Adult Gray ~ 91 16 Alive
122309 04/19/80 Pup Gray ~ 79 10 Alive
122310 04/23/80 Adult Gray d 101 13 Alive
122311 04/23/80 Adult Gray cJ 112 13 Alive
'
t't I l'c ·1 r· 'I l r!'''l ... -r······~ ~ [:.::..=::;]
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the ultimate fate of each of them is important to our
understanding of population dynamics, it should be mandatory
that all tags, tattoos, radio-collars, etc. from tagged big game
species be reported. This type of regulation would also aid
enforcement officials in enforcing other regulations.
Wolf Dispersal
Table 68 summarizes pertinent data concerning 26
radio-collared wolves known to have dispersed from their
original pack area during this study period. Sixty-nine percent
of the dispersing wolves were males, of which seven (39%) were
under 2 years of age. Females under 2 years of age comprised 50
percent of the dispersing females. Dispersing males averaged
35 months (range of 6 roo. to 92 mo. ) of age while females
averaged 37 months (range of 15 mos. to 112 mo.).
Wolves dispersed from their original pack territories
during all months of the year except December (Fig. 11).
Dispersal appeared to be most prevalent during April through
June and in · September, with 48 percent of all dispersals
occurring during these months.
Of the 26 known dispersals, at least 11 (42%) were in the
company of one or more wolves when last observed. Five were
associated with one other wolf in apparent vacant areas and six
were in company of two to four wolves, of which three cases may
have been vacant territories. At least three and possibly four
wolves were known to have become members of previously
established packs. The reasons why wolves left established
territories may b~ related to prey abundance and/or attempts by
young wolves to colonize new areas. During the study at least
eight dispersing wolves were observed leaving and returning to
the original pack before making the final movement. Seven of
these wolves were estimated to range in age from 12 to 24
months.
Average distance moved by dispersing wolves prior to loss
of radio contact was 67.7 miles (106 km) [range of 14 (22.4 km)
to 460 mi (736 km)]. The longest documented movement (736 km)
exceeds the longest movement (670 km) recorded in the literature
(Van Camp and Gluckie 1979). Excluding this record movement,
dispersing wolves moved an average of at least 49.9 mi
(76.8 km).
There is a paucity of information in the literature
concerning wolf dispersal in populations elsewhere in North
America. Recently, Fritts and Mech (In Press} documented a
number of instances of wolf dispersal in a newly protected wolf
population in northwestern Minnesota. In their study, movements
of loners, dispersers and newly formed pairs were greatly
influenced by established packs. This appeared to be at least
partially true in this study since many dispersers were last
observed with one or more wolves in apparently vacant wolf
1 !, 9
Table 68. Summary of data pertaining to radio-collared wolf dispersal in GMU 13 of southcentral Alaska
from April 1975 through June 1980.
Distance
Estimated • last observed from Date
Original pack Accession age at Date original territory movement
affiliation number Color/sex capture captured mi km occurred Status 6/80
Deep Lake 121993 Gray ~ 9 yr 03/22/76 20 9/76 Missing 10/78
122067 Gray d 3 yr 03/22/76 30 8/76 Shot on 3/77 with Keg
Creek pack
122204 Gray d 11 mo 04/10/78 62 6-7/78 Missing 11/18/78
122212 Black d 2 yr 11/19/78 35 3/79-2/80 Shot 2/80
Delta 122063 Gray ~ 2-3 yr 03/21/77 120 4/77-3/79 Shot 3/79 on Wood River
Ewan 121990 Gray ~ 2 yr 04/15/75 40 2-4/76 Missing from Mendeltna
pack 10/76
121991 Gray d 7 yr 11/04/75 20+ 1/76 Killed by St. Anne's
wolves
122219 Gray d 6 mo 11/17/78 80 11/78-3/79 Trapped
Hogan Hill 121988 Gray d 2.5 yr 11/05/75 40 9/76-3/77 Missing 3/77
122206 Gray ~ 11 mo 04/10/78 ? 9/78 Missing 9/78
Keg Creek 122002 Black ~ 1-2 yr 04/20/75 25 5/76 Shot 7/76
122083 Gray d 3 yr 04/20/75 45 3-10/76 Shot from Mendeltna
pack 1978
Mendeltna 122008 Gray d 5 mo 10/29/76 14 11/77 Killed by Tyone wolves
St. Anne 122086 Gray d 11 mo 04/28/77 ? 5/77 Missing 5/77
122087 Gray d 11 mo 04/28/77 30 5/77 Missing 5/77
122088 Gray d 4-5 yr 04/18/77 32 6/78-3/79 Drowned 1979
122093 Gray d 2-3 yr 10/11/77 80 2/78-9/79 Dead, unknown causes
122118 Gray d 2 yr 02/10/78 460 4/78-2/79 Missing in Brooks Range
3/79
122209 Gray ~ 9 mo 02/26/78 80 4/79-12/79 Dead, starvation prob.
122208 Gray ~ 11 mo 04/10/78 88 8/78-9/79 Missing 9/79
122293 Gray d 3 yr 03/31/79 28 10/79 Missing 1/80
Susitna 122229 Gray d 9 mo 02/06/79 48 1/80 Alive
Tolsona 122210 Gray d 1 yr 06/08/78 60 10/79 Missing
122290 Gray d 10 mo 03/28/79 ? 10/79 Lost contact
(11 Tsusena 122199 Gray ~ 2 yr 02/17/76 60 7/76 Missing 7/78
("_) Tyone 122116 Gray d 6 yr 02/10/78 60+ 6-10/78 Lost contact 10/78
r· ' r ! j
>.; . 4
11 r ~, r l r . 'TI r 1 !' ] ['' ;, j r_:_:] ['_~~=j c____] [ ,
I ·----'
L:.J CJ L.J ["' T 1 c:__;J LJ ~ c..J L:J L..J c...:.J c...J c:.J L..J C.......::.J L..J . I
' .... : ' ,..
Figure 11. Percent frequency distribution of radio-collared wolf dispersals from pack
territories in Game Management Unit 13 of southcentral Alaska from April 1975
through June 1980.
12
iS 11
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0
8
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1~~+4~~~~~~~4-~+4-rtt~rt~rrti~-tt1-rttl-tlttll
I
Jan Feb Mar Apr May June July Aug Sept
MONTH OF YEAR
I
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Oct Nov Dec
c.:..:JJ c:...:._] [_]
territories. Fritts and Mech (In Press) documented one case,
and suspected three other cases in which a lone wolf was
accepted into an established pack. They suspected that these
"loners" may have previously dispersed from the pack they had
joined. They concluded that acceptance of lone wolves by
established packs helps to maintain pack structure in
populations with h,igh mortality which in turn results in a
stable social system and higher productivity. This statement
seems particularly pertinent to this study since the lone wolves
that were accepted into established packs were sexually mature
(average age 37 mo.). This could be particularly important in a
heavily exploited wolf population where pack size is often very
small and mortality is high. Whether these new pack members
eventually become socially dominant in the pack structure is not
known but our observations suggest 't.t."lat in some cases they do.
Experimental Wolf Removal
From January 1976 through July 1978, 60 wolves were shot
from helicopter or fixed-wing aircraft by Department personnel
as part of the experimental wolf removal program (Table 69).
Thirteen of these were killed outside the Susi tna study area.
These included seven from the Keg Creek pack and six believed to
be from the Tyone Creek pack. Although only small portions of
the territories of these packs overlapped the Susi tna study
area, a substantial number of study area moose migrated out of
the area and wintered and calved in the two pack areas {Ballard
and Taylor 1980). As a consequence, moose originally thought to
be inhabiting areas of low wolf density were spending half the
year zin areas where wolf densities approached one wolf per
SO mi . Therefore, between January 1976 through March 1977,
attempts were made to remove wolves occupying the western
Alphabet Hills and lower Tyone Creek floodplains. In addition
to the wolves harvested by Department personnel, 24 wolves may
have been harvested in the susitna study area by the public from
September 1975 through June 1980 (Table 70).
Based on track counts, wolf removal and radio telemetry, we
were able to estimate wolf population densities within the
Susitna River study area from spring 1975 through spring 1980.
During this period wolf populations were surveyed after fresh
snowfall; once in late fall and once in spring. Attempts to
remove wolves from the southern portion of the study area were
terminated in December 1978 while removal effort in the northern
portions of the area north of the Denali Highway were
discontinued in spring 1979.
Prior to initiation of Department wolf removal in 1976,
wolf densiti~ in the Susitna River study were estimated az
1 wolf/~48 mi ( 383 km) in spring 1975 to 1 wolf/78 mi 2 (202 krn ) in fall 1975 (Table 71). Densities werez"ligher if mi
of actual wolf habitat were utilized ( 98 -52 mi ) instead of
the entire study area. Comparison of these density figures with
those presented for the re~ainder of GMU 13 during the same time
1 5 2
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Table 69. Summary of wolves taken by Department personnel in the Susitna River
-n study. Area as part of an experimental wolf removal program from
January 1976 through June 1980 in GMU 13 of southcentral Alaska.
J Long-
Wolf ID Suspected pack bone Date of
number affiliation Color Sex age harvest Location
]~
61324 Oshetna Gray M A 3/20/76 SE Lone Butte
j~ 121995 Oshetna Gray F A 3/20/76 Clarence Lake
61322 Oshetna Gray F A 3/19/76 Clarence Lake
61321 Oshetna Gray M A 3/19/76 Clarence Lake
61319 Oshetna Gray M A 3/19/76 Clarence Lake
J 61320 Pack association Gray M A 3/19/76 Western Alphabet Hills
unknown
121984 Butte Creek Black M A 1/17/76 Butte Creek
61311 Butte Creek Black M A 1/17/76 Butte Creek
J Butte Creek Gray M A 1/17176 Butte Creek
61300 Deadman pack -Black M p 1/17/76 Watana Creek
61301 Deadman pack Black M p 1/17/76 Watana Creek
J 61316 Deadman pack Gray F p 1/17/76 Watana Creek
121981 Deadman pack Gray M A 3/19/76 Watana Creek
121982 Deadman pack Gray-black F A 3/19/76 Watana Creek
61302 Maclaren pack Gray M p 1/17/76 Maclaren River
0 61303 Maclaren pack Gray M p 1/17/76 Maclaren River
Maclaren pack Gray p 1/17/76 Maclaren River
Maclaren pack Gray M A 1/17/76 Maclaren River
121985 Maclaren pack Gray F A 1/17/76 Maclaren River
D 61304 Possible Black M A 1/17/76 Butte Creek
Brushkana pack
61307 Brushkana pack Black F A 1/17/76 Butte Creek
D 121983 Brushkana pack Black F A 1/17/76 Butte Creek
61312 Brushkana pack Black F Yrlg 1/17/76 Butte Creek
61306 Brushkana pack Black F p 1/17/76 Butte Creek
61308 Brushkana pack Black F p 1/17/76 Butte Creek
[] 61310 Brushkana pack Black F p 1/17/76 Butte Creek
61315 Brushkana pack Black F p 1/17/76 Butte Creek
61309 Brushkana pack Black M p 1/17/76 Butte Creek
]' -
61313 Brushkana pack Black M p 1/17/76 Butte Creek
122001 Clearwater Gray M A 7/29/76 Upper Valdez Creek
122002 Coal Creek Black F A 7/29/76 Upper Coal Creek
122003 Coal Creek Gray M A 7/29/76 Upper Coal Creek o·-122012 Clearwater Black F A 10/28/76 Upper Clearwater Creek
-122013 Clearwater Gray M A 10/28/76 Upper Clearwater Creek
122017 Brushkana Gray M A 12/30/76 Brushkana Creek
D
122018 Brushkana Gray F p 12/30/76 Brushkana Creek
122019 Brushkana Black F p 12/30/76 Brushkana Creek
122020 Brushkana Black F A 12/30/76 Brushkana Creek
122021 Brushkana Black M p 12/30/76 Brushkana Creek
J 122022 Brushkana Black M p 12/30/76 Brushkana Creek
122058 Unknown Gray F A 3/20/77 Middle Maclaren River
D
1 53
D
Table 69
Wolf ID
number
122059
122060
122010
122011
122067
122068
122069
122070
122071
122072
122089
122192
122193
122194
122195
122196
122198
122200
122211
(cont.). Summary of wolves taken by Department personnel the Susitna River
study area as part of an experimental wolf removal program from January
1976 through June 1980 in GMU 13 of southcentral Alaska.
Long-
Suspected pack bone Date of
affiliation Color Sex age harvest Location
Clearwater Black F p 3/20/77 Lower Clearwater Creek
Watana Gray M A 3/19/77 Delusion Creek
Keg Creek Gray F p 3/22/77 Keg Creek
Keg Creek Gray M p 3/22/77 Keg Creek
Keg Creek Gray M A 3/22/77 Keg Creek
Keg Creek Black F A 3/22/77 Keg Creek
Keg Creek Black F p 3/22/77 Keg Creek
Keg Creek Black F p 3/22/77 Keg Creek
Keg Creek Gray F A 3/22/77 Keg Creek
Clearwater Black F p 3/23/77 Valdez Creek
Clearwater Black M p 5/28/77 Valdez Creek
Jay Creek Gray M p 3/23/78 Clarence Lake
Jay Creek ·Gray M A 3/23/78 Clarence Lake
Jay Creek Gray M p 3/23/78 Clarence Lake
Clearwater Black F A 3/21/78 Middle Clearwater Creek
Watana Gray M A 3/20/78 Fog Creek
Watana Gray M p 3/22/78 Fog Creek
Watana Gray F A 3/22/78 Fog Creek
Clearwater Gray F Yrlg 7/08/78 Clearwater Creek
1 54
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L
L
L
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L
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Table 70. Summary of number of wolves harvested by public which possibly
could have been located in the Susitna River study area 1975-1980.
Possible
pack association Sex Age Color Date Comments
Brushkana pack M A Black 10/24/75
Valdez Creek M ? ? 10/75
Mouth Maclaren River M A Gray 2/25/77
Mouth Maclaren River F A Gray 2/25/77
Mouth Maclaren River M A Black 10/8/76
Susitna River F p Gray 3/7/77
Denali Highway Unk Unk Gray 9/18/77
Susitna River F A Black 2/3/78
Susitna-Watana M A Gray 2/12/78
Mouth Maclaren River M p Black 9/10/77
Mile 122 Denali Highway M p Gray 2/22/79
Denali Highway M p Gray 3/13/79
Denali Highway M p Gray 4/13/79
Denali Highway M A? Gray 9/29/79
Susitna River M ? ? 4/20/80
Susitna River F ? ? 4/20/80
Susitna River F A Gray 3/17/80
Coal Creek F ? Gray 2/13/80
Coal Creek M ? Gray 2/13/80
Coal Creek F ? Gray 2/13/80
Coal Creek F ? Gray 2/13/80
Butte Lake F A? Gray 2/25/80
Butte Lake F A? Gray 2/25/80
Butte Lake F A? Gray 2/25/80
1 5 s
Table 71. Numbers and density of wolves estimated to occupy the Susitna River study area from
(J1
Ol
June 1980.
Possible or Known
Pack Affiliation
Brushkana River-
Butte Creek
Coal Creek
Clearwater Creek
Jay Creek
Maclaren River
Middle Fork of
Susitna River
Watana-Deadman Creeks
Subtotal
Lone Wolves
(10% of subtotal)
Total
Sq. miles/wolf
Total area (2804 sq. mi.)
Habitat area (1858 sq. mi.)
' •'
Spring Fall
1975 1975
7 12
6? 10?
2 6
2 5
l7 33
2 3
19 36
148 78
98 52
F ,~~1 !"''' 1 r 1 lr'"''"" "'l r· :1 r· l r '! ., r·'" 1
Spring
1976
2
2
3
0
?
1
2
10
1
ll
255
169
Fall Spring Fall Spring Fall
1976 1977 1977 1978 1978
7 1 2 2 2
0 0 0 0 0
3 2 0 1 2
2 2 4 1 4
? l 0 0 0
2 0 2 2 0
4 2 6 1 3
18 8 14 7 11
2 1 1 l 1
20 9 15 8 12
140 312 187 351 234
93 206 124 232 155
spring 1975 through
Spring Fall Spring
1979 1979 1980
3-4 3-4 ?
0 0 0
0 ? 2
3-4 10 6
0 5 ?
0 0 2
2+ 7 5
10 26 15
1 3 2
11 29 17
255 97 165
169 64 1D9
1 u
J
J
J
J•i
D
D
0
0
D
J-
o-·-
D
D
;1 u
D
period (Table 60) suggests that wolf densities in the Susi tna
River study area were slightly lower than those elsewhere in the
Unit. This was probably attributable to the fact that
topography, vegetation, and snow cover in the Susi~~a study area
make wolves more vulnerable to hunting and trapping.
For purposes of evaluating the effects of wolf removal on
moose calf survival, spring wolf densities were used since they
would represent the number of adult wolves which could
potentially prey upon moose calves. Newborn pups would have low
food demands for a few weeks following birth, but their demands
would increase and probably approach those of an adult by late
November. Our estimates of wolf numbers in the study area
indicate that density was lower than the spring 1975 level by 42
percent, 53 percent, 58 percent, 42 percent, and 11 percent for
springs 1976, 1977, 1978, 1979, and 1980, respectively. If the
two large packs outside the study area were included for the
years that removal was maintained, reductions in wolf density
from 1976 through 1978 would have been 37,. 54, and 69 percent,
respectively. In either case, wolf numbers averaged less than
50 percent of the spring 1975 level from 1976 until removal was
terminated in spring 1979.
Wolf Repopulation of Removal Area
In spite of continuing public harvests, wolf populations in
the Susitna River study area had increased to within at least 89
percent of the spring 1975 population by spring 1980. In all
likelihood, the spring 1980 wolf population is even higher than
that presented in Table 71 because we lack data for two
previously occupied areas which we suspect are now occupied by
an unknown number of wolves.
The rapid repopulation of this study area can be attributed
to both immigration and to high reproductive rates for the few
remaining wolves which were not removedby either the Department
or the public. Immigration of wolves into the study area was
significant. Dispersal and movement of individual
radio-collared wolves such as number's 067, 204, 219, 002, 001,
199 and 116 (described under Pack Histories) demonstrate how
rapidly areas void of wolves can be repopulated.
During this study, annual rates of wolf population increase
were extremely high: 89.5 percent in 1975, 81.8 percent in
1976, 66.7 percent in 1977, 50.0 percent in 1978, and 163.6
percent in 1979. For the 5-year period, the average annual rate
of increase was 93 percent. We suspect that this rate will
decline as the habitat becomes saturated and ·Wolf harvests
increase.
Evaluation of Wolf Removal on Moose
Initially we planned to evaluate the effect of wolf removal
on moose calf survival by comparing fall calf:cow ratios as
1 57
determined by standard moose sex and age composition counts.
Pre and post wolf removal ratios within specific count units
were to be compared. Also, ratios in low wolf density count
units could be compared with those in medium to high wolf
density count units. This approach presented certain problems,
however. Calf:cow ratios in a count unit normally fluctuate
from year to year and factors such as moose density, climate and
habitat vary between count units. Therefore, results of direct
comparisons between count units or between years could be
obscured or exaggerated by these factors. Also, wolf densities
in. these comparison areas were never constant and fluctuated
widely from year to year in response to hunting and trapping
(Table 60).
In an effort to m~n~m~ze some of these Droblems, we
selected combinations of count units in which caif:cow ratios
had historically fluctuated in phase. We assumed that when
ca1f:cow ratios were positively correlated between groups of
count units over a period of years, there was a high likelihood
that factors influencing ratios in those units were similar. We
expected these units to continue to exhibit similar trends
unless some factor was altered in one group of units but not the
other. If wolf densities strongly influenced moose calf
survival, we expected a divergence in trends between a group of
count units in which wolf densities were substantially reduced
and another group of units in which densities remained "normal."
We selected combinations of composition count units in
which fluctuations in calf:cow ratios, yearling bull percentages
and number of moose seen per hour were similar prior to wolf
reduction (Figs. 12 through 14) . count units 3, 6 and 7 were
within the Susitna River study area (wolf removal area) while
count units 10, 13, 15, 16, and 17 were in areas of "normal"
wolf density (Fig. 15). Count areas 2, 5, and 14 were adjacent
to the Susitna Rivar study area and wolf densities in those
areas were occasionally influenced by removal activities. Not
all count units were surveyed each year. The strongest
correlation of calf:100 cow ratios prior to wolf control was
between comparison areas 1 and 4 {P<0.05, r=0.86).
Trends in calf:cow ratios between comparison areas 1 and 3
and 1 and 4 (Fig. 12) diverged in 1976 and 1977 but not in 1978
or 1979. Prior to wolf reductions, calf; cow ratios in the
Susi tna River study area fluctuated considerably and exceeded
30:100 in 4 of 7 years. Therefore, the changes observed in 1976
and 1977 could have resulted from either reduced wolf predation
or normal variation in ot.her factors. Because yearling cow
moose cannot be accurately identified from fixed-wing aircraft,
the number of cows older than 2 years of age was calculated by
subtracting the number of yearling bulls observed (representing
a minimum estimate of the number of yearling cows) from the
total number of cows observed {Ballard et al. 1980). This
method assumes an equal sex ratio at birth and does not include
yearling bulls taken by hu~ters prior to the survey. Therefore,
1 58
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40
35
30
25
20
15
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Correlation coefficients
prior to 1976
(p<0.05
Areas 1 & 3
Areas. 1 & 4
Areas 1 & 2
Areas 2 & 3
Areas 2 .& 4
.754)
r =
r =
r =
r =
r
.72
.86
.84
.54
.69
(initiated
winter 1975-76)
trol Area
3. 1111111111111111111
No Wolf Control
(Unit 13 (CA 10,13,16,17)
~-No Wolf Control
(Unit 13 (CA 10,13,15,16,17)
2.. ····················
Adjacent Count Units l/
L-------~---------.--------.---------.--------,lr--------r--------.---------r1--------,1 Influenced by Wolf Control-
1969 1970 1971 1972 1973 1974 1J75 19~6 1977 1Y78 {CA 2,5 ,14)
Figure 12. Calf
Game
I Survey Year 1
1979
per 100 cow moose ratit:"s from selected moose sex and age compositif? count areas within
Management Unit 13 of Southcentral Alaska from 1969 through 1979,-Only CA-5 surveyed in 1979.
8
7
6
5
4
3
2
1
Correlation coefficient
prior to 1976
(p<O.OS = .754)
Areas 1 & 3
Areas 1 & 2
Areas 2 & 3
4.
r =
r =
r =
.74
.80
.58
No Wolf Control
(Unit 13 (CA 10,13,15,16,17)
2. Adjacent Count Units
Influenced by Wolf Control
( CA 2, 5 ,14) !/
1969 1970 1971 1972 1973 1974 1975 1976 1977 1978
* Not plotted, nearly
identical to CA's 10,13,
15, 16, 17.
Figure 13.
Survey Year
Percent small bulls within selected moose sex and age compositjon count areas within Game
Management Unit 13 of Southcentral Alaska from 1969 to 1978. 1. Only CA-S survey in 1979.
;r···,., r· ·· ., r .... , r· ·1 ~· '' '] [' ':·'l
[''' "1 [ --i c.....J. c.:J
llO
100
90
80
70
60
50
40
30
20
'.
.... ····· ·····
,... ,.
Wolf Control Area (initiated w1nter .1975-76)
111111111111111111111
No Wolf Control
(Unit 13 (CA 10,13,16,17)
No Wolf Control
(Unit 13 (CA 10,13,15,16,17)
Adjacent Count Units ~------~--------~--------~------~~------~--·------~--------~------~~-------, Influenced by Wolf Control!/
1969 1970 1971 1972 1973 1974 1975 1976 1977 ,·1978 (Ca 2,5,14)
.survey Year
Figure 14. Moose per hour values from se]P~~Pn mnnse sex and a~e composition ce~nt areas within Game
Management Unit 13 of Southcentral Alaska from 1969 through 1978, lJ Only CA-5 survey in 1979.
[......]
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Figure 15. Boundaries of fall moose sex and age composition count areas within GMU-13 of Southcentral Alaska •
. ~ ~:-~ ' ~ S}; ....
at)
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the estimate of the number of cows older than yearlings is
exaggerated. Nevertheless, calf:cow ratios generated from these
estimates are more meaningful because they exclude some of the
sexually immature cows.
Using the above described method of computing calf:cow
ratios we analyzed statistical differences in ratios between
comparison areas 1 and 3 and 1 and 4 using an analysis of
residuals (Everith 1977:46) to determine which cells of a
chi-square table were significant in rejection of the null
hypothesis {Table 72). There were no significant (P>0.05)
deviations for the years Department wolf removal was in effect,
suggesting that reductions in wolf density did not increase
moose calf survival. Significant (P<O.OS) deviations did occur
in 1971 which might be explained by differences in winter
severity. This might also explain the significant (P<O.OS)
deviation of numbers of calves observed in the comparison count
area during 1975.
Percentages of yearling bulls {Fig. 13) observed in fall
composition counts were compared between the wolf removal area
and the comparison areas. We would have anticipated a
divergence in trends for percent yearling bulls beginning in
1977, had the 1976 and succeeding calf crops experienced greater
survival due to wolf control. This was not the case and,
although there was an increase in 1978, all of the comparison
areas continued to fluctuate in phase suggesting again that
Departmental wolf control had not resulted in increased calf or
short yearling survival.
We also would have anticipated an increase in moose
harvests in the wolf removal area if wolf reductions had
increased ~~e survival of bull calves, short yearlings and
adults. Numbers of bull moose harvested in the susi tna River
study 3.rea were compared with harvests in the remainder of
GMU 13 (Table 73). Significant deviations (P<0.10) in numbers
of moose harvested in the Susitna River study area occurred in
1976 and 1979 (Table 73B). In 1976, following the first year of
wolf removal, the bull harvest was lower than expected while in
1979, after both wolf removal and the transplanting of brown
bears (discussed below), the harvest was higher than expected.
This analysis again suggests that moose survival was not
enhanced by reductions in wolf density .
The only evidence which suggested that wolf reductions had
increased moose calf survival in the Susitna study area was a
p2sitive but nonsignificant correlation (P>O.OS, df = 3,
r = 0.70) between fall calf:cow ratios (corrected) and the
preceeding spring wolf zdensity. This relationship was not
evident (P>O.OS, 4db, r = 0.53) for comparative count areas
outside the Susitna River study area. We tend to discount the
former relationship because of only four data points and because
calf:cow ratios in the comparative areas had similar trends in
ratios.
1 6 3
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Table 72. Adjusted residuals (d) of number of calves and cows (~2 yr old) observed
in fall moose sex and age composition surveys conducted in the Nelchina
and Susitna River Basin of southcentral Alaska from 1969-1979.
Deviations from expected valuel/ Deviations from expected valuelf
Comparison Comparison
Year area Cow.s Calves area Cows Calves
1969 1 0.160 -0.240 1 0.276 -0.409
3 -0.095 0.163 4 -0.147 0.253
1970 1 0.331 -0.527 1 0.448 -0.708
3 -0.361 0.655 4 -0.487 0.892
1971 1 -1.237 2. 352~\-1 -1.039 1.943
3 1.083 -2.346* 4 0.855 -1.855
1972 1 0.540 -1.181 1 0.618 -1.342
3 -0.581 1.448 4 -0.605 1.521 F
1973 1 -0.198 0.457 1 -0.080 0.181
3 0.203 -0.533 4 0.077 -0.203
1974 1 0.286 -0.487 1 0.407 -0.688 r
3 -0.270 0.524 4 -0.383 0.750
1975 1 0.523 -1.185 1 0.287 -0.668 L
3 -0.988 2.552~\-4 -0.42 1.136
1976 1 0.151 -0.300 1 0.062 -0.123 r
3 -0.142 0.322 4 -0.055 0.128 L
1977 1 -0.293 0.535 1 -0.533 0.989
3 0.296 -0.616 4 0.508 -1.094
1978 1 0.511 -0.970 1 0.343 -0.659
3 -0.518 1.119 4 -0.329 0.733 L
1979 1 -0.620 1.174 1 -0.541 1.018
3 0.671 1.448 4 0.567 -1.237
y (/d/>1.96, P<0.05) * Significant deviations denoted by
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Table 73.
Area
Numbers of moose reportedly harvested in the
Susitna River study area in comparison to the
remainder of Game Management Unit 13 for harvest
years 1974 through 1979.
Number of moose harvested
1974 1975 1976 1977 1978 1979
Susitna River study area 145
562
103 83 104 148 163
Remainder of GMU 13 479 462 502 612 588
Table 73B. Adjusted residuals (d) of moose harvests from the Susitna
· River study area and the remainder of Game Management
Unit 13 for harvest years 1974-1979.
Deviations from expected value!!
Area 1974 1975 1976 1977 1978
Susitna River study area 0.996 -0.657 -1.962'\--0.974 0.376
Remainder of GMU 13 -0.481 0.317 0.947 0.470 -0.181
Significant deviation (/d/>1.67, P<0.10) denoted by*·
1979
1. 780*
-0.859
1 6 5
Analyses presented thus far in this report indicate that
wolf predation was not a major cause of moose calf mortality in
GMU 13 during this study. Winter food habits and rates of
predation on short yearling and adult moose, however, suggest
that wolf predation might constitute an important mortality
factor for these age classes. Experimental wolf removal also
provided an opportunity to test this thesis. If it were true,
the effects of wolf removal would be manifested by an increase
in adult moose survival. To test this possibility we compared
ratios of adult cow (~2 yr old) and calf (<6 mos.) moose
observed per hour of survey in comparison areas 1 and 3 to those
ratios observed in comparison areas 1 and 4 (Table 74). For all
comparisons no significant (P>0.05) deviations in ratios of cow
and calf moose observed per hour of survey were evident
following reductions in wolf density initiated in 1976. This
analysis suggests that adult and short yearling moose survival
was not significantly improved as a result of wolf control.
This evaluation of wolf removal was based on the premise
that one factor, wolf predation, was responsible for low calf
survival. If this assumption were correct, removal of this
factor should have produced substantial increases in calf: cow
ratios, particularly since moose pregnancy rates were high
(Ballard and Taylor 1980). Prior to the study, we did not
select a calf:cow ratio at which we would have concluded calf
survival had definitely improved as a result of wolf control.
However, a value of 50 calves to 100 cows appeared to be
reasonable if wolf predation were indeed primarily responsible
for low calf survival. Obviously the reported ratios were
considerably less. If, however, wolf predation were only one of
several factors a smaller increase would be expected.
Composition counts, as currently conducted, are probably not
precise enough to measure such small changes.
Calf:cow ratios for the stu?y area were compared with those
in other moose populations 1n the State (Fig. 16). Of
particular interest were the data from Mt. McKinley National
Park (Troyer 1976, 1977 and 1978). Wolf densities in ~e Park
have been estimated at approximately one wolf per 50 mi (Murie
1944 and Haber 1977). Wolf populations in the Park are for the
most part unexploi ted, and thus we would expect the moose
population to be characteristic of a population subjected to
high levels of wolf predaton. The study area, the remainder of
Unit 13 excluding the Susitna River study area, and McKinley
Park have all exhibited the same trends in calf:cow ratios since
1974 even though wolf densities differed greatly among these
areas. In contrast, however, Unit 20A calf:cow ratios have
increased tremendously following wolf removal in 1976
(Gasaway et al. 1977).
Another, more direct method of evaluating the effects of
wolf removal on moose calf survival was initiated in late spring
1977. Newborn moose calves were captured and fitted with
special transmitters which alerted biologists that mortality had
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Table 74. Adjusted residuals (d) of cow (~2 yr old) and calf (~6 mo) moose
observed per hour of survey during moose sex and age composition
counts conducted in the Susitna River study area in comparison
to comparative areas in the remainder of Game Management Unit 13
from 1969-1979.
Deviations from e~ected value!/ Deviations from exEected value!/
Comparison Comparison
Year area Cows Calves area Cows Calves
1969 1 -0.793 1.133 1 -3.025* -0.412
3 -0.403 1.174 4 2.065* 2.415*
1970 1 -1.889 0.054 1 0.266 1.272
3 0.693 1.579 4 -1.171 0.476
1971 1 -1.728 -0.200 1 -0.765 0.221
3 1.737 -0.573 4 0.856 -0.376
1972 1 1.614 0.149 1 0.848 -0.299
3 -0.806 -1.076 4 -0.234 -0.932
1973 1 0.447 -0.549 1 0.584 -0.581
3 . 0.507 -1.294 4 0.164 -0.901
1974 1 -0.304 0.440 1 -0.506 0.213
3 -0.107 0.350 4 0.246 0.314
1975 1 -0.417 -0.539 1 1.947 0.014
3 -0.068 -0.100 4 -1.514 -0.936
1976 1 -0.494 -0.324 1 -0.095 -0.228
3 0.649 -0.257 4 0.487 -0.537
1977 1 0. 719 1.092 1 0.138 0.633
3 -0.645 -0.721 4 -0.163 -0.599
1978 1 0.365 0.382 1 0.437 0.297
3 -0.223 -0.439 4 -0.261 -0.665
1979 1 0.651 1 0.534 0.902
3 -0.473 -0.960 4 -0.584 -0.839
y (/d/>1.96, P<O.OS) * Significant deviations denoted by
1 6 7
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00
50
40
35
30
25
20
15
10
5
1969 1970 1971
...-"'\
/ \
.... ---\
1972
\
\
\
\
\
\
\
\
'
1973 1974 1975
Survey Year
___ ,.,.. _,....-
Wolf control
'·~ .
1976 1977
/
/
/
/
/
/
Bears transplanted
from 1327 mi2 area
Wolf Control Area ----------
(initiated winter 1975-76)
Unit 20A
McKinley Park!;
Unit 13 excluding CA' s
3,6,7. 11111111111111
1978 1979
Figure 16. Comparison of calf p~r 100 cow moose ratios between Susitna River Study Area and other moose
populations within Alaska from 1969 through 1978.
!/Only small portion of Park survey in 1979, sample size of 103 (Troyer, pers. comm.) .
rc c 1 ' r 'll "" . c=J r,.,.,. en r .. ''1 r··· l t''''' ·~ r' "'TI [~ c.....J [=.:J c..:.__] I r:l·o'·l [ _ __]
l
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occurred (Ballard et al. 1979). During 1977 and 1978, 136
calves were radio-collared. Fifty-four percent of the calves
died of natural causes, with predation by brown bears accounting
for 79 percent of those deaths (Ballard et al. 1981). Causes of
calf moose mortality were compared between the wolf removal area
and two comparative areas; no significant differences (P>0.05)
in causes of mortality were detected, indicating that
differences in wolf density were not an important factor
influencing calf survival. Further, only 3 percent of the
radio-collared calf mortality was attributable to wolf
predation. ·
Results of moose calf mortality studies in 1977 prompted
initiation of a brown bear food habits study (Spraker et al.
1981; Ballard et al. 1981). This study confirmed the findings
of the moose calf mortality study in that adult bears ( ~3 yr.
old) were preying upon ungulates (predominately moose calves) at
the rate of 1 kill/6.1 days. Subsequently, in spring and summ~2
1979, 48 brown bears were transplanted from a 1,327 m~
(3,437 km 2 ) portion of the Susitna River study area, partially
in an attempt to determine if the moose population would respond
to a decrease· in bear density. Details of the transplant and
resulting bear population estimates were provided by Miller and
Ballard ( 1980) . Moose calf survival in the bear removal area
increased by 24 calves:lOO cows (~2 yr. old) over 1978 levels
(Ballard et al. 1980). Comparative moose count areas did not
demonstrate increases in calf survival during the same period.
High calf survival continued through winter 1979-80 as evidenced
by the high survival of radio-collared yearlings and a spring
calf:lOO cow (uncorrected) ratio of 56 (Ballard and Gardner,
unpub. data). This large increase in calf survival was the
result of a temporary reduction in bear density of approximately
58 percent (Ballard et al. 1980; Miller and Ballard 1980).
During this year of inc12eased ca~f survival, wolf densities
ranged from 1 wolf/169 mi (438 km ) in spring -J979 whe:JZ the
bear transplant was in progress to 1 wolf/64 mi (166 km ) by
fall 1979. ~y sprin~ 1980, wolf densities had increased to
1 wolf/109 mi (282 km ); within 89 percent of the spring 1975
pre-control wolf density. Thus, even though wolf densities
increased, calf and short yearling moose survival remained high
in the study area.
During 1976 and 1977, we grossly estimated the GMU 13 fall
moose population at 11,000 to 19,000 animals based upon
sightings of tagged moose (Ballard and Taylor 1980). Using this
estimate we attempted to grossly model the dynamics of this
moose population based upon our findings. Assuming a stable
moose population of approximately 15,000 moose, over 10,000
moose calves are produced annually in GMU 13 based upon a
pregnancy rate of 88 percent (Ballard and Taylor 1980) and a
parturition rate of 1.15 calves/cow (Ballard and Cornelius,
unpubl. data). According to predation rate data based on
sightings of radio-collared bears (Spraker et al. 1981;
Ballard et al. 1981) and wolf scat analyses from den and
1 c 9
rendezvous sites (this report), we calculated that brown bears
were annually killing over 4, 000 moose calves from mid-May to
mid•July, while wolf predation was responsible for 434 to 1,013
calves annually during the same time period. Based upon the
15, 000 moose population estimate and fall calf percentages
derived from fall composition counts (ADF&G files),
approximately 3, 000 calves could not be accounted for by our
simple model. Similar types of problems existed for adult moose
mortality.
Application of observed wolf and bear predation rate data
to the adult moose population estimate resulted in an estimate
that roughly 25 percent of the adults $Uccumbed to predation
alone. This crude modeling attempt indicates that our predation
rate and composition data are in error, and/or that adult moose
population estimates are too low, and/or that our understanding
of moose population dynamics are inadequate. Obviously more
sophisticated modeling techniques could solve some of these
problems. However, it is apparent that the lack of adequate
moose population estimates will hinder attempts to quantify the
effects of wolf . and bear predation on moose. Attempts to
quantify moose populations will be attempted in fall 1980 in
conjunction with studies on the impacts of susitna River
hydroelectric development on moose. Perhaps these estimates
will allow us to better quantify the effects of predation on
moose.
Ballard et al. (1981) reviewed the causes of newborn moose
calf mortality in three separate Alaskan moose populations:
Kenai Peninsula, Tanana Flats, and this study. In all three
studies predation was the leading cause of summer calf
mortality: by black bears on the Kenai (Franzmann et al. 1980);
wolves on the Tanana Flats (Gasaway et al. 1977); and brown
bears in the Nelchina Basin (Ballard et al. 1981). Although
predation by brown bears was especially significant in the
Nelchina Basin, the GMU 13 moose population is probably
receiving predation pressure from all three predator species in
addition to human harvest.
In summary, data collected in this study do not support the
hypothesis that wolves are responsible for the low calf moose
survival reflected in fall moose composition counts in GMU 13.
These data also appear to partially refute the hypothesis that
at the observed densities wolves are responsible for high rates
of adult moose mortality. tf wolf control increased moose
survival, the increases were not of sufficient magnitude to be
reflected in fall moose counts or calf mortality studies.
Predation rate studies, however, indicate that wolf predation
may be an important cause of yearling and adult moose mortality.
If moose survival increased as a result of lowered wolf
densities it was certainly overshadowed by the increases in calf
and probably adult moose survival resulting from the removal of
brown bears.
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RECOMMENDATIONS
To quantify the effects of both wolf and bear predation on
GMU' 13 moose; data from this and other related
predator-prey studies should be used to model the moose
population. This analysis would· aid game managers in
formulating strategies for managing predators and prey.
We should continue to annually monitor wolf numbers in
GMU's 11 and 13 by both aerial survey and radio telemetry
to determine changes in density, and aid in assessing the
potential impact of wolf predation on moose.
For GMU's 11 and 13, we propose that game managers consider
instituting a season bag limit of seven wolves in an effort
to stabilize wolf numbers at a post hunting density of
approximately 1 wolf/100 mi 2 of habitat. We speculate that
establishment of this objective would maximize sustained
wolf harvests and yet not allow predation on moose to
become excessive.
Hunters and trappers should be required by regulation to
return all tags, radio-collars, etc. from all species of
wildlife marked for scientific study. Although
implementation of this regulation will not guarantee
compliance it will at least· aid in informing the public
that the information is needed.
Further research is needed to determine desirable
predator-prey ratios for moose management. Bot.."l predator
and prey densities should continue to be manipulated in the
Susi tna study area and the effects moni tared to achieve
this goal.
Wolf repopulation of the Susitna study should be monitored
for at least 1 more year to gather additional data on
population dynamics and rates of increase.
ACKNOWLEDGEMENTS
A large number of individuals participated in various
portions of this project and it would be impossible to tJ.'lank
each separately. First, the senior author would like to thank
his wife, Artina, for her patience and assistance with this
project. Not only did she donate numerous hours to this
project, but she served as a springboard for ideas and made a
number of suggestions to improve and strengthen the project.
We were fortunate to have Karl Schneider as our supervisor
for this series of projects. During the studies he continually
made constructive suggestions for ways of improving the project.
His indepth editing of our reports and suggestions for
alternative ways of analyzing portions of the data are greatly
appreciated. We also appreciate his successful efforts in
assuring that these research projects were adequately funded to
accomplish project objectives.
1 7 1
Sterling Eide, Area Management Biologist, participated in
nearly all phases of this project from 1976-1980. He generously
shared ideas and made numerous suggestions for improving this
project to make it more applicable to management.
A number of temporary employees served as field assistants
during this program and deserve recognition. They included
R~ Ridgeway, T. -Balland,-L-~ Metz,-J. -Westlund, ··and-c~ Gardner~
Craig Gardner assisted with data collection, scat analyses, data
tabulation and analyses for this report.
A number of other Department biologists assisted with
various phases of this project. Albert Franzmann and
Robert Tobey advised us on various aspects of the project and
participated whenever their services were needed.
Charles Lucier and his laboratory staff provided excellent
laboratory support. Victor VanBallenberghe participated in the
first year of the project.
We wish to extend our thanks to bush pilots Mr. Alfred Lee,
Lee's Air Taxi, · and Mr. Kenneth Bunch, Sportsman's Flying
Service, for the many safe hours of flying they performed. Both
of these individuals donated their time on many occasions to
help make the project a success. Both Vern and Bud Lofstedt,
Kenai Air Service, piloted the helicopter for various tagging
operations. Both were always willing to participate in any
aspect of the project whenever they were needed.
SuzAnne Miller, biometrician, came to our rescue in the
latter stages of this project. She not only advised us on
various statistical procedures but performed many of t.."'lem on her
personal time.
Don Cornelius served as an assistant while this report was
being prepared and did an excellent job of keeping up with field
studies. He also reviewed and made constructive comments on
early drafts of this manuscript.
Both Raymond Kramer and SuzAnne Miller extensively read and
reviewed this report and made a number of recommendations for
its improvement.
We further wish to acknowledge the assistance provided by
other biologists in the Glennallen office, Messrs. Bird,
Potterville, Roberson, Webster and Williams, for helping out
when needed. We especially thank Kathy Adler for placing blood
and morphometric data on computer sheets and for developing an
excellent filing system in addition to handling budgets.
Lastly we would like to thank our supervisors James Faro,
Robert Rausch, Karl Schneider, Ron Somerville and John Vania for
providing support when it was needed and for allowing us the
freedom · to conduct this study in the manner we felt necessary.
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LITERATURE CITED
Adorjan, A. s., and G. B. Kolenosky .. 1969. A manual for the
identification of hairs of selected Ontario mammals. Ont.
Dep. Lands For. Res. Rep. Wildl. 90. 47 pp.
Alaska Department of Fish and Game. 1973. Alaska game
management policies. Alaska Dept. Fish and Game. Juneau.
65 pp.
Atwell, G., P. Garceau, and R. A. Rausch. 1963. Wolf
investigations. Alaska Fed. Aid Wildl. Rest. Proj. W6R3,
Work Plan K. Juneau. 28 pp.
Averbeck, G., J. c. Schlotthauer, and w. B. Ballard. In prep.
Abdominal parasites of wolves in Southcentral Alaska.
J. of Wildl. Diseases.
Baer, C. Harold, R. E. Severson, and S. B. Linhart. 1978.
Live capture of coyotes from a helicopter with ketamine
hydro~hloride. J. Wildl. Manage. 42{2):452-454.
Ballard, W. B., and K. P. Taylor. 1978. Upper Susitna
River moose population study. Alaska Dept. Fish and
Game. P-R Proj. Rep., W-17-10, Job 1.20R. 62 pp.
, A. W. Franzmann, K. P. Taylor, T. Spraker,
----~c~.~c~.~S~c~hwartz, and R. o. Peterson. 1979. Comparison
of techniques utilized to determine moose calf
mortality in Alaska. Proc. N. Am. Moose Conf.
workshop, Kenai, Alaska. 15:362-387.
, and T. Spraker. 1979. Unit 13 wolf studies.
------:::-A ... l-a-sk"a Dept. Fish and Game. P-R Proj. Rep., W-17-8,
Jobs 14.8R, 14.9R and 14.10R. 90 pp.
, and K. P. Taylor. 1980. Upper susitna Valley
____ m_o_o_s_e_p-opulation study. Alaska Dept. Fish and Game.
P-R Proj. Final Rep., W-17-9, W-17-10, and W-17-11.
102 pp.
, s. D. Miller,
----~c-a~l~f~-m-ortality study.
and T. H. Spraker. 1980. Moose
Alaska Dept. Fish and Game .
P-R Proj . Final Rep. ,
W-21-1, 123 pp.
W-17-9, W-17-10, W-17-11, and
-----~-----' T. H. Spraker, and K. P. Taylor. 1981. Causes
of neonatal moose calf mortality in sout.~central Alaska.
J. Wildl. Manage. In Press. ·
Bishop, R. H., and R. A. Rausch.
fluctuations in Alaska,
101:559-593.
1974. Moose population
1950-1972. Nat. Can.
1 7 3
Clark, K. R. F. 1971. Food habits and behaviour of the
tundra wolf on central Baffin Island. Univ. Toronto.
PhD. Thesis. 223 pp.
Cowan, I. MeT. 1947. The timber wolf in the Rocky Mountain
National Parks of Canada. Can. J. Res. 250(5):139-174.
Eide, s. 1979. Wolf survey-inventory report -1977-78,
Game Management Unit 13. pp. 95. In Hinman, R. A.
(Ed.) 1974. Ann. Rep. Survey-Inventory Activities,
Part II. Furbearers, Wolf, Wolverine, Small Game. Ak.
Fed. Aid in Wildl. Rest. Rep., Proj. W-17-10. 192 pp.
Etkin, W. 1964. Cooperation and competition in
behavior. pp 1-34. In Etkin, W. (Ed.). 1964.
behavior and organization among vertebrates.
Chicago Press, Chicago, Illinois.
social
Social
Univ.
Everith, B. s. 1977. The analysis of contingency tables.
John Wiley and Sons, Inc. New York. 128 pp.
Floyd, T. J., L. D. Mech, and P. D. Jordon. 1978. Relating
wolf scat content to prey consumed. J. Wildl. Manage.
42(3):528-532.
Franzmann, A. W., and P. D. Arneson. 1973. Moose Research
Center Studies. Alaska Dept. Fish and Game. Fed. Aid in
Wildl. Rest. Proj. Rept., W-17-5. 60pp. (multilith}.
, and 1976. Marrow fat in
----~~~-------Alaskan moose femurs ~n relat~on to mortality factors.
J. Wildl. Manage. 40(2):336-339.
----~~------~-' c. C. Schwartz, and R. 0. Peterson. 1980.
causes of summer moose calf mortality on the Kenai
Peninsula. J. Wildl. Manage. 44(3):764-768.
Fritts, s. H., and L. D. Mech. In Press. Dynamics,
movements, and feeding ecology of a newly-protected
wolf population in northwestern Minnesota. Wildl.
Monogr.
Fuller, T. K., and L. B. Keith. 1980. Wolf population
dynamics and prey relationships in nort..lJ.eastern
Alberta. J. Wildl. Manage. 44(3):583-602~
Gasaway, W. C., P. Haggstrom, and 0. E. Burris. 1977.
Preliminary observation on the timing and causes of
calf mortality in an Interior Alaskan moose population.
Proc. N. Am. Moose Conf. Workshop. 13:54-70.
Haber, G. c. 1977. Socio-ecological dynamics of wolves and
prey in a subarctic ecosystem. PhD. Thesis. Univ. of
British Columbia. 585 pp.
1 7 4
r
-':-=-
L.
r
L
L
F
L
F
i
L
l u
J
J
n
J
]'
u-·
D-~.
[ ..
!
0 .
J
D
Holleman, D. F. and R. 0. Stephenson. In press. Winter
prey selection and consumption by the Alaskan wolf.
J. of Wildl. Manage.
Markgren, G. 1969. Reproduction of moose in Sweden.
Viltrevy, 6(3):1-299.
Mcilroy, c. 1974. Moose survey-inventory progress report -
1972, Ga.Ille Management Unit 13. pp. 66-74. In
McKnight, D. E. (Ed.). 1974. Annual report of
survey-inventory activities, Part II. Moose, caribou,
marine mammals and goat. Ak. Fed. Aid in Wildl. Rest.
Rep., Proj. W-17-5. 269 pp.
-----=----· Moose survey-inventory progress report -1974,
Game Management Units 11 and 13. pp 49-55 and 61-79.
In McKnight, D. E. (Ed.). 1976. Ann. Rep. Survey-
Iiiventory activities, Part I I . Moose. Ak. Fed. Aid
in Wildl. Rest. Rep., Proj. W-17-7. 187 pp.
Mech, L. D. 1970. The wolf:the ecology and behaviour of an
endangered species. The Nat. Hist. Press. 384 pp.
~----~~· 1974. Current techniques in the elusive
w1.lderness carnivores. Proc. XI Internat. Congress of
Game Biol. 315-322 pp.
Miller, s. M., and W. B. Ballard. 1980. Estimates of the
density, structure and biomass of an interior Alaskan
brown bear population. In Ballard et al. 1980. Moose
calf mortality study. Alaska Dept. Fish and Game. P-R
Proj. Final Rep., W-17-9, W-17-10, W-17-11 and W-21-1.
Mohr, C. 0. 1947. Table of equivalent populations of North
American small mammals. Am. Midl. Nat. 37(1):223-249.
Murie, A. 1944. The wolves of Mount McKinley U.S. Natl.
Park Serv., Fauna Ser. 5. 238 pp.
Neiland, K. A. 1970. Weight of dried marrow as indicator
of fat in caribou femurs. J. Wildl. Manage.
34(4):904-907.
Nielson, C. A. 1977. Wolf necropsy report: preliminary
pathological observations. Alaska Fed. Aid Wildl.
Rest. Special Rept., Proj. W-17-8 and W-17-9.
Peterson, R. 0. 1977. Ecological studies of wolves on Isle
Royale. Ann. Rep. 1976-77. 12 pp.
1977. Wolf ecology and prey relationships
on Isle Royale. Natl. Park Serv. Sci. Monogr. Ser. 11.
21 pp.
1 7 s
. 1980. Wolf-moose investigation on the
---=K-e-n-a"'~---:P=-e-ninsula, Alaska. Quarterly Rep. #15. Kenai
Nat. Moose Range.
Rausch, R. A. 1967. Some aspects of the population ecology
of wolves, Alaska. Am. Zool. 7:253-265.
. 1969. A summary of wolf studies in
_____ s_o_u~th'L'·_.c-entral Alaska. 1957•1968. Trans. N. Am. Wildl.
and Nat. Resour. Conf., 34:117-131.
----~~-~·' R. J. Somerville, and R. H. Bishop. 1975.
Moose management in Alaska. Naturaliste Can.,
101:705-721.
Sergeant, D. E., and D. H. Pimlott. 1959. Age
determination in moose from sectioned incisor teeth.
J. Wildl. Manage. 23(3):315-321.
Skoog, R. 0. 1968. Ecology of caribou (Rangifer tarandus
granti) in Alaska. PhD. Thesis, Univ. of Cal.,
Berkeley, California. 699 pp.
Snedecor,G. w., and w. G. Cochran. 1973. Statistical methods.
Iowa State Univ. Press. 593 pp.
Sakal, R. F., and F. J. Rohlf. 1969. Biometry.
W. H. Freeman and Co., San Francisco. 776 pp.
Spraker, T. H., W. B. Ballard and S. D. Miller. 1981.
Game Management Unit 13 brown bear studies. Alaska
Dept. Fish and Game P-R Proj. Rept. W-17-10, W-17-11,
W-17-21. Job 4.13R.
Stenlund, M. H. 1955. A field study of the timber wolf
(Canis lupus) on the superior National Forest,
Minnesota. Minn. Dept. Conserv. Tech. Bull. 4. 55 pp.
Stephenson, R.
and Game.
18 pp.
0. 1975. Wolf report. Alaska Dept. Fish
P-R Proj. Rept., W-17-3 through W-17-7.
. 1978. Unit 13 wolf studies. Alaska
----=-~--~~ Dept. F~sh and Game. P-R Proj. Rept., W-17-8, Jobs
14.8R, 14.9R and 14.10R.
, and L. Johnson. 1972. Wolf renort.
---,A::-'!1.-a-s-;k-a-....,D=-e-p~t. Fish and Game. P-R Proj. Rept., W-17-3.
51 pp.
., and 1973. Wolf report.
---,A:-:1;-a-s...,k-a--,D=-e-p-.t. Fish and Game. P-R Proj. Rept., W-17-4.
52 pp.
1 7 6
1 ,J
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Tobey, R. 1980. Wolf survey inventory report -1978-79,
Game Management Unit 13. pp 81. In Hinman, R. A.
(Ed.). 1980. Annual report of survey-inventory
activities, Part IV. Furbearers, upland game,
wolf and wolverine. Ak. Fed. Aid in Wildl. Rest.
Rept., Porj. W-17-11. 112 pp.
Theberge, J. B., S. M. Dosenbrug, and D.
Site and seasonal variations in
Algonquin Park, Ontario. Canadian
92(1):91-94.
H. Pimlott. 1978.
food of wolves,
Field-Naturalist,
Van Camp, J., and R. Gluckie. 1979. A record long-distance
move by a wolf {Canis lupus). J. Mammal. 60:236.
Voigt, D. R., G. B. Kolenosky, and D. H. Pimlott. 1976.
Changes in summer foods of wolves in central Ontario.
J. Wildl. Manage. 40(4):663-668.
PREPARED BY:
Warren B. Ballard
Game B~olog1st
SUBMITTED BY:
Karl B. Schneider
Reg1onal Research Coord1nator
APPROVED BY:
Research Chlef, DlVlSlOn of GamE
Appendix I. Blood values of wolves sampled in Game Management Unit-13 of southcentra1 Alaska from July 1976 through May 1977.
Wolf Carbon Uric Total Packed
Accession dioxide BUN Creatinine Bilirubin Acid Protein Albumin Globulins Alpha l Alpha 2 Beta % Gama % A/G Cell· %
Number (meg/L) (mg/dl) (mg/dl) (mg/dl) (mg/dl) (gm %) (gm %) (gm %) (gm %) (gm %) (~m %) (gm %) Ratio Volume Hemoglobin
122001 4 47 1.3 0.0 3.6 7.90 3.72 4.18 0.25 0.21 1.86 1.86 0.89
122002 0 47 1.3 0.0 2.5 6.90 3.31 3.59 0.17 0.21 1.71 1.50 0.92
122003 ll 42 1.2 0.0 2.6 7.30 3.23 4.07 0.28 0.23 2.01 1.55 0.79
122004 0 50 1.2 B 0.3 7.30 2.86 4.44 0.30 0.18 2.45 1.51 0.64
122007 14 7l 0.7 0.5 6.95 3.36 3.59 0.47 0.47 1.90 0.60 0.95
122008 l3 45 0.7 0.1 6.10 3.12 2.98 0.45 0.53 1.55 0.45 1.05
122009 ll 59 1.0 0.2 6.20 3.40 2.80 ·o.24 0.57 1.37 0.62 1.21
122010 l3 21 0.2 0.0 5.70 3.19 2.51 0.43 0.23 1.52 0.32 1.27
122011 14 24 0.8 0.1 5.50 3.08 2.42 0.25 0.48 1.32 0.36 1.27
122012 3 70 1.7 1.5 8.3g/dl G
122017 ll 54 1.5 0.6 3.2 8.00 3.60 4.40 0.23 0.12 3.37 0.68 0.82
122018 12 27 1.2 0.4 2.2 6.50 3.41 2.09 0.29 0.44 2.07 0.29 l.ll
122019 10 16 l.l 0.1 1.3 2.80 1.85 0.95 0.12 0.12 0.64 0.06 1.94
122021 10 20 0.9 0.2 2.6 5.50 2.99 2.51 0.33 0.28 1.57 0.33 1.19
122022 10 29 0.8 0.2 2.6 6.70 3.23 3.47 0.39 0. 76 1.95 0.37 0.'93
122048 15 108 1.6 0.3 1.8 9.10 4.55 4.55 0.33 0.91 2.48 0.82 l.OO
122049 14 89 1.5 0.2 2.1 9.50 5.60 3.90 0.52 0.69 2.36 0.33 1.43
122056 7.00 4.16 2.84 0.27 0.75 1.04 0. 77 1.46 51
122057 12 36 1.1 0.0 0.9 6.70 3.30 3.40 0.25 l.OO 1.96 0.20 0.97
122058 10 18 1.3 0.1 1.4 7.00 3.35 3.65 0.34 0.26 2.12 0.92 0.92 56
122061 12 23 1.1 0.0 l.l 5.50 2.87 2.63 0. 39 0.52 0.85 0.87 l. 09
122062 14 26 1.1 0.0 l.O 6.30 3.50 2.80 0.28 0. 72 1.25 0.56 1.25 55 19.5
122063 15 52 0.8 0.0 0.5 4.70 2.44 2.26 0.33 0.31 0.99 0.63 1.08 49 20
122064 14 68 l.l 0.0 0.8 5.40 3.22 2.18 0.28 0.67 0.59 0.64 1.47 49 19.8
122067 13 25 1.3 0.0 1.3 5.50 2.40 3.10 0.27 0.95 1.64 0.24 0. 77 43
122069 8 27 1.3 0.0 1.6 5.60 2.94 2.66 0.40 0.28 1.24 0.74 l.ll 45
122070 18 48 0.9 0.0 2.1 5.30 2.92 2.38 0.33 0.42 1.27 0.36 l. 23 37
122072 9 36 1.3 0.1 5.1 5.60 3.49 2.11 0.19 0.61 0.38 0.93 1.66 50
122083 12 60 1.3 0.1 1.4 7.60 4.18 3.42 0.40 0.54 1.31 1.17 1.23 58
122084 12 67 0.9 0.0 1.6 6.70 2.85 3.85 0.49 0.41 2.49 0.45 0. 74 39
122085 12 79 0.8 0.0 1.9 6.20 3.21 2.99 0.43 0.37 1.80 0.40 1.07
122086 17 28 1.2 0.0 1.1 6.30 3.23 3.07 0.43 0.35 1.97 0.32 1.05 46
...., 122087 12 28 1.2 0.1 0.4 6.80 3.45 3.35 0.46 0.26 2.08 0.55 1.03 40
00 122089 8 39 l.l 0.3 5.5 7.20 3.01 4.19 0.28 0.45 3.14 0.31 0. 72
. . .. 'II r··"'l r 1~ r· 'I l r 'i'"1 r··1···l r., .... , .. l r··· ., r·"· 1 r· ' .. , I'T '1 r··"' l f''"'' l [ ' _] [L_J [____]
Appendix I (cont.).
Wolf Alkaline
Accession Glucose Cholesterol Triglycerides LDH SGOT SGPT Phosphatase Phosphorus Calcium Iron Sodium Potassium Chloride
Number (mg/dl) (mg/dl/) (mg/dl) (U/L) (U/L) (U/L) (U/L) (mg/dl) (mg/dl) . (ug/dl) (meg/L) (meg/L) (meg/L)
122001 9 207 18 G 89 D 247 12.2 198 157 G 125
122002 266 227 86 G D D 67 G 13.3 295. 153 G 118
122003 79 217 7 G 63 57 119 G 11.7 259 G G 130
122004 0 194 87 G T T 38 G 12.9 359 158 G 114
122007 70 113 73 323 300 T 28 4.3 8.8 188 G 6.4 124
122008 104 117 22 94 41 75 109 7.5 9.9 165 145 6.1 114
122009 79 163 23 232 238 229 64 2.7 8.1 189 G 6.1 130
122010 102 180 6 137 108 103 176 8.7 12.3 98 159 6.4 122
122011 106 161 25 127 96 86 182 6.8 9.9 112 155 5.7 122
122012 241 145 G T T 69 G 8.0 231 148 G 94
122017 11 262 110 G 20 T 130 G 12.8 250 151 G 104
122018 154 241 30 G D D 358 G 13.8 195 148 G lOS
122019 479 110 20 165 138 64 113 4.0 6.6 73 110 6.5 82
122021 73 216 29 G 307 161 10.0 11.3 173 132 G 95
122022 56 221 35 G 308 162 G 12.5 148 143 G 102
122048 51 232 152 262 220 101 69 5.7 13.4 317 G 6.3 G
122049 50 268 78 389 157 121 76 5.8 G 353 G 6.6 G
122056
122057 137 222 15 177 336 384 163 3.6 10.4 186 146 4.4 123
122058 133 180 18 1070 1070 830 30 3.6 9.6 196 149 7.2 U8
122061 274 145 31 95 50 38 78 2.9 10.2 U2 147 4.4 121
122062 95 140 27 163 93 65 35 1.5 10.3 154 148 4.8 119
122063 176 125 54 128 134 111 64 2.8 8.7 121 147 4.5 U9
122064 132 116 57 109 157 127 24 2.1 9.3 150 148 4.2 121
122067 106 128 37 3500 2780 892 114 6.9 8.9 184 142 9.9 110
122069 58 135 82 1222 1250 400 131 13.0 13.6 150 142 14 107
122070 59 148 229 510 486 208 134 9.6 11.6 110 147 8.6 105
122072 334 140 52 213 126 63 32 7.4 10.6 88 148 9.6 115
122083 32 177 18 444 182 112 38 4.1 9.8 269 153 5.4 123
122084 UO 255 62 234 121 71 78 3.9 9.3 148 145 5.9 119
122085 83 152 126 214 264 us 33 2.8 8.0 139 148 5.2 us
~ 122086 92 191 ll 206 92 76 86 3.3 10.6 128 150 5.1 117
--.:I 122087 53 185 30 G 161 88 38 2.3 U.O 79 148 5.0 120
w 122089 239 240 T T 252 G ll.l 178 148 G 111 102 G
Appendix II. Hair mineral element values (ppm) from wolves harvested and radio-collared in GMU's 11 and 13 of
southcentral Alaska from June 1976 through May 1977.
Accession
Number Zn Cu Ca Mg K Na Cd Co Fe Pb Mn Cr
122001 237 28 1540 185 1460 6030 0.9 0.10 112 6 0.8 0.1
122002 178 15 690 95 540 5770 1.2 0.16 45 2 0.3 0.1
122003 164 19 1210 120 1030 5960 l.l 0.14 144 8 0.8 0.2
122004 160 11 870 95 580 5710 l.l 0.14 74 6 0.6 0.5
122008 171 14 400 60 610 5700 1:0 0.13 . 59 <l 0.3 0.3
122009 222 14 680 120 790 5720 1.3 0.16 58 6 0.4 0.8
122010 172 10 380 60 820 5790 l.l 0.15 59 <l 0.3 0.3
122011 223 17 520 95 1120 5820 1.3 0.14 63 4 0.4 0.1
122012 169 10 480 40 390 5670 l.l 0.27 61 <l 0.4 <0.1
122013 175 17 sao 70 490 5300 0.8 0.17 80 4 0.2 <0.1
122014 202 10 710 110 1470 6320 l.l 0.19 76 <l 0.4 0.6
122015 85 10 250 45 4690 6120 1.0 0.15 74 <l 0.7 <0.1
122016 272 35 850 100 1920 5930 1.2 0.14 87 <l 0.8 0.3
122017 165 13 420 65 410 5520 1.3 0.14 64 :<1 0.6 0.3
122018 175 10 390 30 1200 5720 1.3 0.21 65 3 0.2 0.2
122019 150 16 740 85 1960 5790 1.3 0.18 71 <1 0.6 <0.1
122020 142 18 620 95 790 5870 0.9 0.13 76 14 0.3 0.1
122021 177 32 1140 125 2800 5910 1.4 0.20 105 •8 0.6 0.4
122022 151 14 650 55 2030 5860 1.0 0.18 74 16 0.4 <0.1
122023 193 12 335 145 1585 2470 1.2 0.21 50 l 1.3 0.4
122024 149 20 700 95 1270 5220 IS IS IS . IS IS IS
122025 169 6 630 125 965 1350 1.4 0.19 7 <1.0 1.1 0.2
122027 195 9 465 95 445 730 1.5 0.25 20 ,2 0.9 0.4
122028 196 13 1285 235 470 1715 1.4 0.24 53 6 1.3 0.2
122029 180 6 275 80 525 1390 1.5 0.16 17 8 1.0 0.3
122030 168 8 365 125 915 2090 1.7 0.25 35 <1.0 1.2 0.2
122031 188 16 395 200 1940 2970 1.4 0.19 121 12 1.0 0.4
122032 213 13 215 110 585 880 1.5 0.14 30 7 0.9 <0.1
122033 194 8 200 120 1090 860 1.3 0.19 15 <1.0 0.9 <0.1
00
0
Appendix II (cont.). Hair mineral element values (ppm) from wolves harvested and radio-collared in GMU's 11 and 13
of southcentral Alaska from June 1976 through May 1977.
Accession
Number Zn Cu Ca Mg K Na Cd Co Fe Pb Mn Cr
122075 168 9 220 55 430 815 1.4 0.21 20 9 1.2 0.7
122076 172 8 220 90 855 3235 1.4 0.27 29 9 1.4 1.0
122077 174 8 255 95 1245 3175 1.6 0.25 31 8 1.1 0.8
122078 133 7 230 70 470 3275 1.6 0.26 25 13 l.l 0.9
122081 155 <l.O 465 115 600 910 1.1 0.19 23 <1.0 0 .. 6 <0.1
122083 241 1 1170 285 700 1600 l.S 0.30 98 9 1.1 0.4
122084 197 <l.O 665 125 300 780 1.1 0.10 179 8 0.8 0.1
122085 183 4 450 115 345 555 1.5 0.19 52 9 0.5 0.3
122086 206 5 1185 165 720 1610 1.1 0.21 69 14 0.7 <0.1
122087 161 <1.0 890 200 715 490 1.1 0.17 49 <1.0 1.0 <0.1
122088 194 3 760 215 845 550 1.0 0.20 so 9 0.6 <0.1
122089 171 <1.0 895 185 495 1385 1.3 <0.1 170 l3 0.5 <0.1
co
Appendix I I I. Copy of paper presented at Portland Wolf
Symposium, held in Portland, Oregon in August 1979
(J. 0. Sullivan and P. C. Paquet, co. eds.
GRAY WOLF -BROWN BEAR RELATIONSHIPS IN THE
NELCHINA BAS IN OF SOUTHCENTRAL ALASKA
Warren B. Ballard
Alaska Department of Fish and Game
P.O. Box 47
Glennallen, Alaska 99588
ABSTRACT: From June 1976 through June 1978, 16 wolf (Canis
lupus) packs were observed at 130 kills in the Nelchina
Basin. seventeen of these kills were contested by brown
bears ( Ursus arctos) • Nine of the 17 kills contested by
both bears and wolves were observed in conjunction with
studies of the Mendeltna wolf pack. Comparisons of
predation rates, predator densities and prey densities were
made between two packs which were intensively studied during
late May and June 1977 and 1978. It was suggested that the
disproportionate number of bear-wolf encounters at kill
sites for the Mendeltna pack was primarily the result of a
lower moose (Alces alces gigas) density in the Mendeltna
area. Descriptions of bear-wolf encounters are given.
Mortality of each predator species as a result of encounters
is also described. Possible significance of these
observations to predator-prey relations is discussed.
INTRODUCTION
The wolves of southcentral Alaska have been the focus
of interest and study for over 30 years. From 1948 to 1953
poisoning by the Federal Government reduced populations of
wolves to low levels. In 1953 only 12 wolves were estimated
to survive in the Nelchina study area described by Rausch
(1969). Bears, wolverines (Gulo gulo), other carnivores and
some omnivores were probably reduced by ~~ese Federal
poisoning efforts. The wolf population gradually increased
and reached a peak of 400 to 450 animals in 1965. Rausch
(1969) summarized the status of wolves in this region from
1957 through 1968.
Rausch (1969), Bishop and Rausch (1975), and Mcilroy
( 1974) described the history of the Nelchina Basin moose
population. The moose population began declining after the
severe winter of 1961-62. The decline continued with severe
winters occurring in 1965-66 and 1971-72. Although wolf
predation was not suggested as the main reason for the
population decline, it was thought to have at least
accentuated the decline, and perhaps more importantly,
prevented recovery during mild winters (Bishop and Rausch
1 8 2
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1975). This trend of thought, coupled with the findings of
Stephenson and Johnson ( 1972 # 1973), which revealed high
percentages of calf moose in wolf scats, suggested that wolf
predation on moose calves was preventing the moose
population from recovering. Consequently, in 1975 a series
of studies on wolf-moose relationships were initiated.
These studies were later expanded to include brown
bear-moose relationships. Information pertaining to these
studies was reported by Stephenson (1978), Ballard and
Taylor (1978a,b), Ballard and Spraker (1979), Spraker and
Ballard (1979) and Ballard et al. (in press). Considerable
attention was focused on gathering wolf food habits
information during the late spring and summer when most
moose calf mortality occurred. The purpose of this paper is
to report on observed encounters between bears and wolves
during these studies, and to discuss the significance of
such interactions.
MATERIALS AND METHODS
Radio-collared wolves were tracked and visually
observed, when possible, from fixed-wing aircraft. These
methods were similar to those described by Mech (1974).
Monitoring intensity varied seasonally and from pack to pack
but consisted of at least bimonthly efforts during winter
months. Two wolf packs were intensively studied during the
summers of 1977 and 1978 and were located either once or
twice daily from late May to mid-July in 1977 and to late
June in 1978.
Ages of captured wolves were determined on the basis of
tooth eruption and wear. During radio-tracking, ages of
unmarked wolves were occasionally estimated on the basis of
relative sizes and also by the criteria described by
Jordan et al. (cited in Mech 1970). The age-sex structure
of certain packs was not ascertained until the animals had
been killed by hunters and trappers. Hunters and trappers
were encouraged to provide us with wolf carcasses taken in
the study area by offering $10.00 per carcass. Ages of
harvested wolves were determined by examining epiphyseal
cartilage of the longbone according to methods described by
Rausch (1967).
Moose kills were classified as calf, yearling or adult
from fixed-wing aircraft based on combinations of size,
pelage, and antler growth. When practical, wolf kills were
examined on the ground. Cause of death was determined
according to ·methods described by Stephenson and Johnson
(1973) and Ballard et al. (in press).
Observations of circumstances surrounding bear-wolf
encounters were recorded, as they occurred, in a notebook
and then elaborated upon after the flight ended. When it
became apparent that more than two or three observations
1 8 3
would be collected, notes were recorded on a Sony tape
recorder (Model TC-55) and transcribed after the flight had
terminated.
Wolf summer home ranges were determined by plotting all
radio locations for individual packs and then connecting the
outermost observations. Sizes of home ranges were determined
with a compensating polar planimeter.
Study Areas and Wolf Pack History
The study was conducted in Game Management Unit 13 of
southcentral Alaska. The area, commonly referred to a~ the
Nelchina Basin 2 consists of approximately 61,595 km of
which 18,798 km is over 1,200 m elevation.
The study was conducted primarily in those portions of
the unit lying north of the Chugach Mountain Range and east
of the Talkeetna Mountains. Year-round studies involved up
to 16 wolf packs. Only data pertaining to wolf-bear
relationships ?-nd their implication to wolf summer food
habits will be presented here.
Mendeltna Wolf Pack
The Mendel tna wolf pack occupied the Lake Louise and
Oshetna River Range Units described by Skoog (1968).
Boundaries of its year-round territory are depicted in
Figure 1.
Generally, the area consists of a level plateau of wet
muskeg interspersed with numerous ponds and lakes. Drier,
rolling hills on the western portion of the area comprise
the foothills of the Talkeetna Mountains. Elevations range
from about 600 m to 1, 170 m on the western edge in the
Talkeetna Mountains. Much of the lowland areas is
vegetated with sparse to dense stands of black spruce (Picea
mariana) and white spruce (P. glauca) interspersed with wet
muskegs containing several species of sedges ( Carex sp. ) ,
grasses, willow (Salix sp.), and birches (Betula sp.).
Drier, better drained sites contain a mixture of white
spruce, willow and shrub birch (Betula glandulosa). In tl::le
higher, western portion of the area spruce densities decline
and the area is a transition between spruce-muskeg and
subalpine-tundra. All stream courses are vegetated with
willows and birch. Sparse stands of spruce occur on
southerly exposures. Well-drained, sandy sites on the
lowlands often contain homogenous stands of aspen (Populus
tremuloides). Understory vegetation is comprised of varying
densities of low bush cranberry (Vaccinium vi tis-idaea);
high bush cranberry (Viburnum edule) and two species of
blueberry (Vaccinium ovalifolium and V. uliginosum).
Lichens are found in varying density throughout the area.
All study areas contain old burns which are more than 30
years old.
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Fig. 1. Map of Game Management Unit 13 and year-round
territories of two wolf packs intensively studied during
summers 1977 and 1978 in the Nelchina River Basin of
southcentral Alaska.
1 8 5
During the 1977 season, the Mendel tna pack numbered
seven adults of which two to four were yearlings. Two
adults and one yearling were radio-collared. This pa~k
occupied a summer home range of approximately 829 km .
During the 1977 season the pack had two den sites at which
two litters, totaling at least eight pups, were raised.
Hogan.Hill Wolf Pack
This pack was located in the Alphabet Hills (Fig. 1) .
The area comprised portions of Skoog's ( 1968) Lake Louise
and Alphabet Hills Range Units. The northern two-thirds of
the area is comprised of "low rounded hills" with elevations
reaching about 1600 m. Higher elevations are characteristic
of subalpine tundra. Lower elevations are thickly vegetated
with white and black spruce. Several creeks bisect the
area, draining primarily into the Gulkana River. The
southern slopes of the Alphabet Hills 1 which were
predominantly utilized by the Hogan Hill pack during summer
1978 1 are thickly vegetated with spruce and willow along
stream bottoms and adjacent to ponds. As the area levels
out, vegetation becomes similar to that of the Lake Louise
Flats although spruce 1 willow and birch densities appear
greater.
During 1978 the Hogan Hill pack was comprised of eight
adults of which at least two were yearlings. One adult and
two yearlings were radio-collared. In 1978 the pack
maintained one den site at which at least five pups wer2
raised. They ranged over an area of approximately 570 km
during early summer 1978. Boundaries of the year-round
territory are shown in Fig. 1.
RESULTS AND DISCUSSION
From June 1976 through June 1978, 16 study wolf packs
were observed on 130 kills, of which approximately 75
percent were moose (Ballard and Spraker 1979). Of that
total, 17 ( 13 .1%} were contested by brown bears . In most
instances I was unable to determine which predator species
had made the kill. During the summers of 1977 and 1978 I
intensively studied one different wolf pack each year and
was able to document some of the circumstances surrounding
bear-wolf encounters at, and away from, kill sites. Because
such observations are rarely witnessed, my notes and
interpretations of bear-wolf encounters for the Mendeltna
Pack during summer 1977 are summarized chronologic~lly
below.
On 11 June, at 2500 hr., adult gray-black female wolf
07 was observed. being chased by a cow moose which was
exhibiting aggressive behavior (mane ruffed-up and ears
down). Wolf would veer off a straight line in what appeared
to be an attempt to lose the pursuing cow. When wolf appeared
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to lose cow by crouching in brush, the cow would search for
the wolf and in three instances was able to find it. The
cow gained ground on the wolf which appeared to tire. On
one occasion the wolf stopped and crouched in the brush.
The cow ran over to the area and appeared to trample the
wolf. Wolf then continued running but at a much slower pace
and with a limp. The chase lasted approximately 15 minutes,
at which time the wolf appeared to head for den site, while
the cow began traveling back the direction from which-it had - -·
come. Cow continued to exhibit aggressive behavior. Cow
began swimming a pond on the other side of which I observed
a brown bear sow with three yearling cubs. The cubs were
huddling over and dragging around a calf moose carcass. Cow
ran around the bears within a 40-50 m radius. Wolf 08, a
yearling male, was present in dense spruce some 150-175 m
away from the bears, but was not observable. I returned to
the calf kill at 2230 hr. via helicopter and frightened the
bears, which were huddled over the calf carcass, away from
the site. Examination of kill revealed puncture marks on
the neck and either puncture or claw marks on the anus.
Only the head had been fed on. The skull was cleaned out so
that all that remained was skin casing. Tongue, eyes and
ears had been eaten which was characteristic of bear-killed
calves (Ballard et al., in press). Imprints were noted in
area and bear hair was noticeably evident on surrounding
brush. Interpretation: Bears made the kill and wolves were
attempting to scavenge, but were chased away by cow or bears
or both.
On 12 June, at 0910 hr., wolf 83, an adult gray male,
and a smaller, drabber gray adult of unknown sex were
observed chas.ing and harassing same bears observed the
previous even~ng. The wolves stayed fairly close to each
other while chasing the fleeing bears. When the bears
stopped running one wolf would crouch and approach the sow.
The sow would charge the approaching wolf at which time the
other wolf would would charge and chase the yearling cubs
causing the sow to charge the second intruding wolf. On one
occasion the wolves treed all three cubs. The wolves
appeared to press their charge when the bearsf direction of
movement was towards den *2 which was less than 2 km away to
the east. On one occasion the radio-collared wolf was
observed sneaking around and crouching down in front of the
bears' direction of movement. Apparently the sow detected
this action because when she was approximately 10 m away,
she charged the crouched wolf and almost caught it by the
hind quarters. It appeared that when the bears finally
established a trend of movement away from the den, the
wolves no longer pursued and began heading back towards the
den. These activities lasted 15 minutes and covered 0. 6 km
from where we first observed the bears. Interpretation:
Wolves discouraged bear movement towards wolf den.
1 e 7
On 14 June, at 1720 hr., wolf 08 (yearling gray male)
was observed alone resting on sand bar. Approximately 60 m
away a single adult brown bear was feeding on an adult moose
kill estimated to be 80 percent consumed. Wolf appeared to
have swollen abdomen, indicating it also had fed on the
kill. Interpretation: Kill made by wolves and wolves
displaced by bear.
On 15 June, at 0850 hr. , -wolves 83 (adult gray male) ,
08 (yearling gray male), and one black yearling were observed
approaching moose calf kill which had one sow and one
yearling brown bear feeding on it. Kill was estimated to be
80 percent consumed with guts and hide remaining. Approach
of airplane and perhaps wolves frightened bears causing them
to run from kill. Wolves went directly to kill and began
feeding. Interpretation: Kill made by bears, observer
approach and/or wolves caused bears to leave kill which was
taken over by wolves.
On 16 June, at 1945 hr., wolves 83 (adult gray male),
08 (yearling gray male), a small gray adult of unknown sex,
and one black yearling of unknown sex were observed
attacking an adult brown bear which possessed an adult moose
kill. Initially three wolves were observed equally spaced
surrounding the bear. One of the wolves was observed
attempting to nip the bear in rump. Bear made several short
charges at wolves which were approaching to within 3-5 m.
Wolves easily out manuevered the bear and three of the
wolves appeared to keep the bear away from the kill as a
fourth wolf fed on it. The bear's direction of movement was
toward the kill and after 15 minutes of encountering the
wolves, the kill was reached. When the bear reached the
kill the wolves stopped harassing the bear and began
traveling in the direction of the main den. Kill was
estimated to be 50 percent consumed. Interpretation:
Either bear or wolves made kill and wolves were attempting
to displace bear. ·
On 22 June, at 082 9 hr. , wolves 83 (adult gray male)
and one adult gray were observed feeding on what I
identified as a moose calf. Ground inspection of the kill
site at 1200 hr. revealed the kill had been misidentified.
Instead of a calf moose, the wolves had been feeding on a
yearling brown bear. A portion of the carcass had been
buried, but most had been consumed. The kill site contained
tracks of a small bear and wolf. Interpretation; Yearling
bear was killed by wolves.
On 24 June, at 1655 hr., wolf 83 {adult gray male) was
observed alone resting approximately 10 m from an adult
moose kill with one adult brown bear on it. Head, rear
quarters, guts and skin were all that remained.
Interpretation: Kill was made by either wolves or bear.
Wolf may have been attempting to scavenge and/or displace
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bear. On 29 June three wolves were observed feeding at kill
site.
On 27 June, at 2200 hr., wolves 83 (adult gray male),
one gray adult and one black yearling were observed resting
close to one adult brown bear which was feeding on calf
moose kill. Estimated kill to be 50 percent consumed with
head and front quarters missing. Bear seemed unconcerned by
presence of wolves. Bear was still present on 28 June at
1000 hr. and had carcass almost consumed. Interpretation:
Kill made by either bears or wolves. Wolves attempting to
scavenge and displace bear.
On 8 August, at 0730 hr., wolves 83 (adult gray male),
one gray adult, one gray yearling and one black yearling
observed scattered around an adult moose kill with one brown
bear on it. Two grays and one black were observed huddled
together touching noses and wagging tails before separating
and charging bear, running it away from kill. A third gray
hidden by a large spruce ran to the kill and tore off a
large ch~nk of flesh as the returning bear charged. Another
gray followed carrying the meat into the dense spruce.
Several bear charges were observed. Bear remained in
possession of kill. Interpretation: Either bear or wolves
made kill. Wolves attempted to scavenge and/or displace
bear.
Aggressive behavior between the two predator species
occasionally results in mortality to the participants.
Joslin (1966) reported that an adult female wolf was killed
close to a den by a black ·bear (Ursus americanus). In
September 1976, a member of the Mendeltna pack was killed by
a brown bear probably as a result of competition over an
adult moose kill. Details of this particular observation
were presented in Ballard (in press). Mech (1970) thought
that occasionally wolves killed bears, but that the victims
were probably cubs, young bears, or older weakened bears.
Murie (1944) suggested that bear-wolf encounters were more
intense on the part of wolves when they occurred close to
wolf dens. The Mendeltna wolves exhibited agonistic
behavior towards bears both at kills and in areas close to
den sites. Observations recorded during this study
substantiate that wolves do occasionally kill bears. The
result of brown bear-gray wolf encounters, therefore, may at
times be an additional source of natural mortality not
previously documented for either predator species. Whether
it is a significant source of mortality for either species
is unknown.
Reasons for Contested Kills
During summer 1977 the Mendeltna pack had six contested
kills in addition to several bear encounters away from kill
sites. In contrast, during a similar study period in 1978
1 B 9
when the Hogan Hill pack was studied, none of six kills were
contested and no bear encounters away from kills were
observed. Reasons for the larger number of contested kills
for the Mendeltna pack may be related to a number of factors
including: (1) observability, (2) predator density and
(3) prey density.
If there was a difference in observability between the
areas, it was not detectable. During summer 1977, the three
radio-collared members of the Mendeltna pack were observed
on 188 of 224 ( 83. 9%) occasions they were located. In
comparison, the three radio-collared members of the Hogan
Hill pack were observed on 97 of 114 ( 85 .1%) occasions.
Differences in bear density are unlikely to have caused
the disproportionate number of contested kills in the
Mendeltna area. Although no accurate estimates of bear
density exist, tagging data and sightings of bears (Ballard
and Taylor l978a, Spraker and Ballard 1979) suggest the
study2 areas had similar densities, approaching one bear per
39 km • There were, however, differences in wolf densities
(Table 1). Based upon areas occupie~ during summer, wolf
densities ranged from o2e wolf/ 73 km for the Hogan Hill
pack to one wolf/119 km for the Mendeltna pack. Thus the
area with the lowest wolf density had the largest number of
kills contested by bears. Differences in wolf density may
have been partially related to the maintenance of two den
sites, 8 km apart, by the Mendel tna pack, but was more
likely related to differences in prey density.
Numbers of moose counted in fall sex and age composition
surveys from 1976 throu9h 1978 were utilized to calculate a
crude approximatio~of moose density (Table 1). The number
of moose per 2.6 km counted in the Alphabet Hills count unit
containing the Hogan Hill2 wolf pack terri tory was 1.10,
while .88 moose per 2.6 km was counted in two count units
containing the Mendel tna wolf pack terri tory. Therefore,
the area with the highest wolf density also had the highest
moose density.
I speculate that the bear-wolf encounters observed
while studying the Mendeltna wolf pack were due primarily to
lo~er moose densities in that area. This speculation ·was
supported by predation data collected by monitoring
radio-collared bears. These data indicate that bears took
substantial numbers of moose in all of the areas studied
(Ballard and Spraker, in prep.). Thus, for the Hogan Hill
area, where moose were more abundant, no kills were
contested, because, I suspect, sufficient moose were
available for each predator during the study period.
Speculation that low prey densities were responsible
for the disproportionate number of contested kills for the
Mendeltna wolf pack was also supported by data on the
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Name
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Mendeltna
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Summary of predator-prey statistics for two wolf Pffk areas intensively studied during early summer 1977 and
1978 in the Nelchina Basin in southcentral Alaska.-
Wolf Density Mo,ose fl No. and % of Available Kg. Prey
Within Summer Density of Kills Bear-Wolf Known Feeding Prey Biomass
Study Range (Moose~ Wolves Contested Kill Rate Rate Biomass Per Adult
Period (Wolf/km 2) 2.6 km ) Present Kills (Days/Kill) (Days/Feed) (kg) Wolf/Day
28 May -l/73 1.10 6 0 = 0% 4.0 4.0 841 4.4
21 June 1978
27 May -l/119 .88 ll 6 = 55% 10.0 4.6 2,173 6.2/4.7
15 July 1977
!/ Biomass of available food based upon following assumptions: Weight of adult moose= 427.5 kg (from Franzmann and
Bailey 1977) and yearling moose= 197.5 kg. (from Franzmann and Arneson 1973, 1975) of which approximately 75%
(from Peterson 1977) available as food yielding 321 kg. and 148 kg., respectively.
Newborn calf moose weights 13.3 kg. (from Ballard and Taylor 1978) and gains weight at rate of 1.3 kg/day (from
Franzmann and Arneson 1973). Therefore, 15 day old calf weighs 32.8 kg., of which 90% is consumable yielding
29 .s kg.
Yearling brown bear weighs 45 kg. (from Spraker and Ballard 1979) of which 75% is consumable yielding 34 kg.
Snowshoe hare weighs 1.4 kg. (from Burt and Grossenheider 1964) of which all is consumable yielding 1.4 kg.
chronology of calf moose mortalities. During 1977 and 1978,
79 percent of radio-collared calf mortality was attributed
to predation by brown bears (Ballard et al., in review).
During 1977, when the Mendeltna pack was being intensively
monitored, 53 percent of all calf mortalities had occurred
by 11 June. These data correspond with the date of the
first observed contested kill between the Mendel tna wolf
pack and brown bears. This suggests that ample moose were
available for both predator species until mid-June but not
afterwards. If correct, then bear-wolf encounters at kills
for the Hogan Hill wolf pack would be expected to occur at a
later date had a declining prey base influenced its
occurrence. Although daily contact with the Hogan Hill pack
terminated on 21 June, two and possibly three of four kills
observed between 1 July and mid-November were contested by
bears with the first contested kill occurring on 4 August.
Therefore, I suspect that had I continued intensively
monitoring the Hogan Hill Pack beyond 21 June, I would have
observed contested kills before 4 August.
Significance of Contested Kills to Predator Ecology
During this study I was unable to quantify how much was
eaten by each predator species at a particular kill site
because the observation periods were too short and in some
cases, my presence may have interfered. In many cases,
however, it it was apparent that both species were able to
feed at many of the kills for varying lengths of time. The
amount consumed by wolves at a particular kill site could
alter kill rates and influence how kill data are
interpreted.
From 27 May through 15 July 1977, during intensive
monitoring of the Mendel tna wolf pack's activities, all or
some, Mendeltna wolves were observed on 11 kills. The kills
included 6 adult moose, 3 calf moose, 1 yearling
moose and 1 yearling brown bear. In comparison, from 28
May through 21 June 1978 members of the Hogan Hill Pack were
observed on 6 kills comprised of 2 adult moose, 2 calf
moose, 1 yearling moose and 1 of unknown species. Based
upon these data, wolf kill rates were calculated for kills
which were known to have been made by wolves and for kills
when bears were involved (Table 1). The latter rates are
referred to as feeding rates. For known wolf kills there
was a large difference in kill rates: one kill every 4. 0
days for the Hogan Hill Pack which numbered eight adults
versus 10.0 days for the Mendeltna Pack which numbered seven
adults. However, when bear contested kills were added the
Mendel tna Pack rate increased to one kill every 4. 6 days
while the Hogan Hill Pack rate remained unchanged.
The amount of prey biomass available per adult wolf was
calculated for both study packs (Table 1). Two values were
calculated for the Mendeltna Pack: the first value assumed
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that all of the prey biomass was available to wolves even
though bears were present on some kills while for the second
value I arbitrarily assumed that only SO percent of the
biomass on bear contested kills was available to wolves.
Although I could not determine how much was eaten by either
predator at a kill site, I did observe that both usually fed
on some quantity. Regardless, the range of 4.7 to
6.2 kg/wolf/day of available food for the Mendeltna Pack was
greater than the rate of 4.4 kg/wolf/day calcul-ated-for -the
Hogan Hill Pack.
Both the kill and consumption rates during summer for
the Mendel tna and Hogan Hill wolf packs fall within the
range of values reported in the literature for the winter
season. Mech (1970) reported that a pack of 15 to 16 wolves
had a kill rate of one moose per 3.0 days to one moose per
3. 7 days on Isle Royale. Fuller and Keith (in press)
reported a kill rate of one moose per 4.7 days for a pack of
nine wolves in northern Alberta. Peterson { 1977) reported
food availability of 4.4 to 10.0 kg/wolf/day for Isle Royale
wolves from 1971 through 1974 which was considerably less
than the average consumption rate of 22 kg/wolf/day derived
by Mech ( 197 0) . Mech (cited in Peterson 19 77) determined
that one Minnesota wolf pack declined after a winter when
only 3.0 to 3.4 kg/ wolf/day of food was available,
increased at 5.8 kg/wolf/day, and remained stable at
3.6 kg/wolf/day. Mech (1970) reported that a higher kill
rate occurred when calf moose comprised a larger percentage
of the prey taken. The same appeared to be true during this
study.
Stephenson (1978) speculated that competition from
bears at wolf kills could result in an increase in the wolf
predation rate. Data presented from this study indicate
that, if true, the increase may not be detectable with the
study methods used. Competition at kill sites could
increase bear predation rates, however, no data were
collected on this aspect.
Within recent years, scat analyses have been used to
determine wolf food habits. Although most such studies have
acknowledged that the derived data represent what was eaten
rather than what was actually killed, the observations of
wolf-bear encounters further emphasize the need for caution
when analyzing both wolf and bear scat data and interpreting
their significance to predator-prey relationships. If both
species were feeding on the same kill, the resulting food
data could only be viewed as that obtained by scavenging.
Only within recent years have both black and brown bears
been identified as significant predators of cervids
(Schlegel 1976, Franzmann and Schwartz 1978, Ballard and
Taylor 1978, and Ballard et al. In Press). The fact that
both predator species have potential to not only prey upon
1 9 3
ungulate species, but also to scavenge and interact with one
another could greatly complicate our attempts to understand
predator·prey relationships.
ACKNOWLEDGEMENTS
The study was funded in part by Alaska Federal Aid in
Wildlife Restoration Project W-17·R.
Sterling Miller, Karl Schneider, Donald McKnight, and
Karen Wiley, all of the Alaska Department of Fish and Game,
reviewed earlier drafts of the manuscript and made many
helpful suggestions. Appreciation is also expressed to
Rolf Peterson, Michigan Technological University, for
reviewing the manuscript.
REFERENCES
Ballard, W.B. and K.P. Taylor. 1978a. Moose calf mortality
study, Game Management Unit 13. Alaska Dept. Fish and
Game. P·R Proj. Rep., W-17-9 (2nd half) and W-17-10
{1st half), Job 1.23R. 43pp.
and 1978b. Upper Susitna River moose
populat~on Study. Alaska Dept. Fish and Game. P-R
Proj. Rep., W-17-10, Job 1.20R. 62pp.
_____ , A.W. Franzmann, K.P. Taylor, T. Spraker, c.c.
Schwartz, and R.O. Peterson. In press. Comparison of
techniques utilized to determine moose calf mortality
in Alaska. 15th N. Am. Moose Conf. Workshop. Kenai,
Alaska.
and T. Spraker. 1979.
Dept. Fish and Game.
14.8R, l4.9R and 14.10R.
Unit 13 Wolf studies. Alaska
P-R Proj. Rep., W-17-8, Jobs
Juneau. 90pp.
. In press. Brown bear
-----canadian Field-Naturalist.
kills gray wolf.
---, T. Spraker and K.P. Taylor. In review. Causes of
neonatal moose calf mortality in southcentral Alaska.
J. Wildl. Manage.
Bishop, R.H. and R.A. Rausch. 1975. Moose population
fluctuations in Alaska, 1950-1972. Naturaliste Canadien
101:559-593.
Burt, W.H. and R.P. Grossenheider. 1964. A field guide to
the mammals. 2nd Ed. Houghton Mifflin Co. , Boston.
284pp.
Franzmann, A.W. and P.D. Arneson. 1973. Moose Research
Center studies. Alaska Dept. Fish and Game. P-R Proj.
Rep., W-17-5. 60pp.
1 9 4
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and 1975. Moose Research Center Report.
Alaska Dept. Fish and Game.
129pp.
P-R Proj. Rep., W-17-7.
and T.N. Bailey. 1977. Moose Research Center Report.
Alaska Dept. Fish and Game. P-R Proj. Rep., W-17-9.
76pp.
and C.C. Schwartz. 1978. Moose calf mortality study,
---.Kenai Peninsula. Alaska Dept. Fish and Game. P-R Proj.
Rept. W-17-10, Jobs l.24R and 17 .3R. 20pp.
Fuller, T.K.
dynamics
Alberta.
and L.L. Keith. In review.
and prey relationships
J. Wildl. Manage.
Wolf population
in Northeastern
Joslin, P.W.B. 1966. Summer activities of two timber wolf
(Canis lupus) packs in Algonquin Park. Unpubl. · M.S.
thesis. University of Toronto. 99pp.
Mcilroy, C. 1974. Moose survey-inventory progress report -
1972,-Game Management Unit 13. 66-74pp. In Mc~~ight,
D.E. (Ed.). 1974. Annual report of survey-inventory
activities, Part II. Moose, caribou, marine mammals
and goat. Ak. Fed. Aid in Wildl. Rest. Rep., Proj.
W-17-5. 269pp.
Mech, L.D. 1970. The wolf: the ecology and behavior of an
endangered species. The Nat. Hist. Press. 384pp.
___ . 1974. Current techniques in the study of elusive
wilderness carnivores. Proc. XI. Internat. Congress
of Game Bio., 315-322pp.
Murie, A. 1944. The wolves of Mount McKinley. U.S. Natl.
Park Serv., Fauna Ser. 5. 238pp.
Peterson, R. 0. 1977. Wolf ecology and prey relationships
on Isle Royale. Natl. Park Serv. Sci. Monogr. Ser. 11.
210pp.
Rausch, R.A. 1967. Some aspects of the population ecology
of wolves, Alaska. Am. Zool. 7:253-265.
1969. A summary of wolf studies in southcentral
Alaska, 1957-1968. Trans. N. Am. Wildl. and Nat.
Resour. Conf., 34:117-131.
Schlegel, M. 1976. Factors affecting calf elk survival in
northcentral Idaho. A progress report. Proc. 56th
Ann. Conf. W .. Assoc. State Game Fish Comm. 342-355pp.
Skoog, R.O. 1968. Ecology of caribou (Rangifer tara.ndus
granti) in Alaska. PhD. Thesis, Univ. of California,
Berkeley, California. 699pp.
1 9 5
Spraker, T. and W.B. Ballard. 1979. Unit 13 Brown Bear
Studies. Alaska Dept. Fish and Game. P-R Proj. Rep.,
W-17-R. Juneau.
Stephenson,
Aalska
Juenau.
R.O. and L. Johnson.
Fed. Aid Wildl. Rest.
Slpp.
1972.
Rept.
Wolf
Proj.
report.
w-17-3.
and L. Johnson. 1973. Wolf report. Alaska Fed. Aid
Wildl. Rest. Proj. W-17-4. Juenau. 52pp.
. 1978. Unit 13 wolf studies.
----~Rest. Prog. Rept. Proj. W-17-8.
Alaska Fed. Aid Wildl.
Juneau. 75pp.
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Appendix IV. Copy of manuscript submitted to Canadian Field
Naturalist
Surplus Killing of Caribou by Wolves
STERLING H. EIDE AND WARREN B. BALLARD Alaska Department of
Fish and Game, P.O. Box 47, Glennallen, Alaska 99588
Eide, S. H. and W. B. Ballard.
Caribou by Gray Wolves.
1981. Surplus Killing of
Canadian Field-Naturalist.
Seven apparently healthy adult caribou were killed and only
partially consumed by a pack of two to four Gray Wolves in
late March 1979 along the Copper River in southcentral
Alaska. Deep snow contributed to the vulnerability of the
Caribou.
Key Words: Gray Wolf, Canis lupus, Caribou, Rangifer
tarandus, surplus killing.
Several investigators have documented instances in
which Gray Wolves (Canis lupus) have killed more prey than
they consumed. Most of these incidents of surplus killing
( Kruuk 1972) occurred during winter months. Mech et al.
(1971) reported surplus killing of White-tailed Deer
(Odocoileus virginianus) in Minnesota; in sweden, Bjarvall
and Nilsson (1976) reported surplus killing of eight
domestic Reindeer. Miller and Broughton ( 1974) observed
surplus killing of Caribou (Rangifer tarandus) calves during
summer 1970. All of these observations, although of
interest to students of predator/prey relationships,
represent the exception to generally accepted predation
principles. To better interpret their significance to both
predator and prey, further explanation of circumstances
surrounding such observations is needed. We describe what
appeared to be surplus killing of Caribou by Gray Wolves
during winter 1979 in the Nelchina Basin of southcentral
Alaska and provide an interpretation of the events leading
to this phenomenon.
On 31 March 1979, seven Caribou carcasses were observed
within a 1 km radius along the Copper River near its
confluence with the Indian River. On the basis of tracks,
puncture marks and subcutaneous hemorrhaging, we inferred
that these Caribou had been killed by a pack of two to four
Gray Wolves. The amount of flesh taken from these carcasses
was subjectively estimated as follows: 1 -90%, 5 -SO%,
1 -20%. Because all carcasses had been scavenged by Bald
Eagles (Haliaeetus leucocephalus), Ravens (Corvus corax) and
Red Foxes (Vulpes fulva), it was difficult to estimate the
amount consumed by Wolves.
Ages of Caribou were estimated on the basis of mandible
wear (Skoog 1968). Percent of fat of the femur was used as
1 9 7
an indicator of physical condition using methods described
by Neiland ( 1970). Ages ranged from 2 to 5 years and the
percentage of fat in femurs ranged from 52 to 88 percent,
and averaged 74 percent (Table 1). All would have been
placed in Cheatum's (1949) visual index classification 3 or
4, indicating that based on femur marrow the animals were in
relatively fair physical condition. Neiland (pers. cornm.)
considered Caribou with less than 25 percent fat to be in
relatively poor physical condition.
Mech (1970) discussed the likelihood of deep snow
contributing to excessive killing by Wolves. On 29 March
1979 (U.S. Geological survey 1979) snow depth at a snow
course near the Sanford River was 92 em. This was 44% above
the 12-year average of 64 em. Only during the severe winter
of 1971-72 have snow depths exceeded those recorded on
29 March 1979. Therefore, deep snow possibly made these
Caribou especially vulnerable to Wolves. Four of the
Caribou were killed on the Copper River where snowpack was
hard and footing good, but tracks indicated they were chased
through deep snow before reaching the river and likely were
exhausted. The other carcasses were located in deep snow
which clearly had impeded their movements.
During relatively mild winters, Wolves completely
consume most Caribou killed (Ballard and Spraker 1979).
However, during relatively severe winters, trappers, guides
and others have reported the occurrence of multiple,
simultaneous killings of Caribou and subsequent failure to
consume large percentages of the flesh. Perhaps during
severe winters surplus killing by Wolves may be more common
than previously suspected. If this were true, predation by
Wolves could be considered an important factor contributing
to the decline of Caribou herds.during severe winters.
We thank L. D. Mech, U.s. Fish and Wildlife Service
and K. B. Schneider and D. E. McKnight, both Alaska Department
of Fish and Game, for providing helpful comments. The study
was funded in part by Alaska Federal Aid in Wildlife
Restoration Project W-17-R.
LITERATURE CITED
Ballard, W. B., and T. H. Spraker. 1979. Unit 13 wolf
studies. Alaska Dept. Fish and Game.· P-R Proj. Rep.,
W-17-8, Jobs 14.8R, 14.9R and 14.10R. 90 pp.
Bjarvall, A. and E. Nillson. 1976. Surplus killing of
reindeer by wolves. J. Mamm., Vol. 57, No. 3.
Cheatum, E. L 1949. Bone marrow as an index of
malnutrition in deer. New York State Conservationist
3(5):19-22.
1 9 8
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Kruuk, H. 1972. Surplus killing by carnivores. J. zool.
166:233-244.
Mech, L. D. 1970. The wolf: the ecology and behavior of
an endangered species. P. 246.
Mech, L. D., L. D. Frenxe1 and P. D. Karns. 1971. The
effects of snow conditions on the vulnerability of
white-tailed deer of wolf predation. Pp. SS-56 in
Ecological Studies of the Timber Wolf in Northeastern
Minnesota. L. D. Mech and L. D. Frenzel, ed., 1971.
Miller, F. L. and E. Broughton. 1974. Calf mortality on
the calving ground of the Kaminuriak caribou during
1970. Canadian Wild. Ser. Rep. series No. 26.
Neiland, K. A. 1970. Weight of dried marrow as indicator
of fat in caribou femur. J. Wild. Mgmt., 34(4):904-907.
Skoog, R. o. 1968. Ecology of caribou (Rangifer tarandus
granti) in Alaska Ph.D. Thesis. Univ. California,
Berkeley. 699.
u.s. Soil Conservation Service. 1979.
water supply outlook for Alaska.
Anchorage, Alaska.
Snow surveys and
U.S. Dept, Aqr. ,
1 9 9
Appendix V.
Naturalist.
Copy note published in Canadian Field
Brown Bear Kills Gray Wolf
WARREN B. BALLARD
Alaska Department of Fish and Game, P.O. Box 47, Glennallen,
Alaska 99588
Ballard, W. B. 1980. Brown Bear Kills Gray Wolf. Canadian
Field-Naturalist 94(1):000-000.
Ground examination indicated a Brown Bear (Ursus arctos) had
killed a Gray Wolf (Canis lupus) at an adult Moose (Alces
alces gigas) carcass in southcentral Alaska. The
observation represents first published evidence of a Gray
Wolf mortality inflicted by a Brown Bear.
Key Words: Gray Wolf, Canis lupus, Brown Bear, Ursus
arctos, mortality, Moose kill, Alces alces gigas,
interspecific relationships.
Several known and potential natural mortality factors have
. been described for Gray Wolves, Canis lupus (Mech 1970) .
This report describes a cause of mortality not previously
documented.
On 25 September 1976, while conducting wolf ecology studies
in the Nelchina Basin of southcentral Alaska, I aerially
tracked a wolf pack ( 6 gray and 3 black wolves) t·o an
adult Moose (Alces alces gigas) kill. The Moose kill
appeared to be less than 2 days old, on the basis of both
color of exposed flesh and degree of consumption (50%).
The next day Alfred Lee, Lee's Air Taxi Service, found a
dead Gray Wolf close to the kill and observed a Brown Bear
(Ursus arctos) running from the site. The site was examined
from the ground the following day. The dead adult male Gray
Wolf was at the base of a White Spruce tree (Picea glauca}
approximately 10 m from the Moose, and the Moose had been
buried. Tracks, plus extensively disturbed vegetation and
soil indicated a fight had occurred.
The wolf had numerous punctures on its throat and around
its anus . At least three cervical vertebrae and the rear
portion of the occipital condyle were crushed. The left
rear femur was fractured.
Evidence indicated that the Brown Bear had killed the Gray
Wolf. An alternate explanation is that otJ.,.er wolves had
killed the animal. Because the victim had been a regular
pack member, and the bone damage was not typical of other
wolf-killed wolves examined, it seems unlikely that the wolf
was killed by pack members.
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Murie (1944) mentioned that it was not infrequent for
bears to discover kills made by wolves and disposess the
wolf to assume ownership. This observation, however, is the
first published evidence of Gray Wolf mortality inflicted by
a Brown Bear. The importance of this type of mortality to
Gray Wolf populations is unknown.
I am grateful to Thomas Balland for assisting with field
observations.
Literature Cited
Mech, L. D. 1970. T~e wolf: the ecology and behavior of an
endangered spec1es. Natural History Press, Garden
City, New York City, New York 384 pp.
Murie, A. 1944. The wolves of Mount McKinley.
States National Park Service, Fauna Series 5.
United
238 pp.
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