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SUSITNA HYDROELECTRIC PROJECT
FINAL PHASE I REPORT
BIG GAME STUDIES
VOLUME V WOLF
Warren B.Ballard,
Craig L.Gardner,
John H.Westlund
and
James R.Dau
ALASKA DEPARTMENT OF FISH AND GAME
Submi tted to the
Alaska Power Authori ty
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ARLIS
JUaskaResources
Library &Information Service
Anchorage,Alaska s
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PREFACE
In early 1980,the Alaska Department of Fish and Game contracted with
the Alaska Power Authority to collect information.useful in assessing
the impacts of the proposed Susitna Hydroelectric Project on moose,
caribou,wolf,wolverine,black bear,brown bear and Dall sheep.This
information,along with information on furbearers,small mammals,birds,
and plant ecology collected by the University·of Alaska,is to be used by
Terrestrial Environmental Specialists,Inc.of Phoenix,New York,in
preparation of exhibits for the Alaska Power Authority's application for
a Federal Energy Regulatory Commission license to construct the project.
The studies were broken into phases which conformed to the anticipated
licensing schedule.Phase I studies,January 1,1980 to June 30,1982,
were intended to provide information needed to support a FERC license
application.If the decision is made to submit the application,studies
will continue into Phase II to provide additional information during the
anticipated 2 to 3 year period between application and final FERC approval
of the license.
Wildlife studies did not fit well into this schedule.Data collection could
not start until early spring 1980,and had to be terminated during fall 1981
to allow for analysis and report writing.(Data continued to be collected
during winter 1981-82,but could not be included in the Phase I report.)
The design of the hydroelectric project had not been determined.Little
data was available on wildlife use of the immediate project area,although
some species had been intensively studied nearby.Consequently,it was
necessary to start with fairly general studies of wildlife populations
to determine how each species used the area and identify potential impact
mechanisms.This was the thrust of the Phase I Big Game Studies.During
Phase II,we expect to narrow the focus of our studies to evaluate specific
impact mechanisms,quantify impacts and evaluate mitigation measures.
Therefore,the Final Phase I Report is not intended as a complete assessment
of the impacts of the Susitna Hydroelectric Project on big game.
The reports are organized into the following eight volumes:
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Volume I.
Volume II.
Volume III.
Volume IV.
Volume V.
Volume VI.
Volume VII.
Volume VIII.
Big Game Summary Report
Moose -Downstream
Moose -Upstream
Caribou
Wolf
Black Bear and Brown Bear
Wolverine
Dall Sheep
SUMMARY
In 1980 and 1981,wolf packs which potentia~ly could be impacted
by the proposed Susitna Hydroelectric Project were studies.
Thirty-six wolves from six separate packs were radio-collared and
relocated a total of 1,175 times from fixed-wing aircraft.Sev-
eral packs occupying areas in or adjacent to the impoundments
were not studied because poor snow conditions precluded their
capture.
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From radio-relocations,ground observations,and previous studies
we were able to describe the histories of individual wolves and
their associated packs.Interactions between pack members and
between separate packs were observed throughout the study.
Wolves in the Watana pack were most frequently observed in shrub
(50%)habitats with ecotones being of particular importance.No
slope or aspect selection was observed.Approximately 1/3 of all
radio-locations of the Watana pack were located in areas to be
inundated by the proposed impoundment.
Of 83 wolf kills observed for six packs,57%were moose,33%were
caribou and the remainder were small mammals.Fifty-one percent
of the moose killed and 7%of the caribou killed by wolves were
calves.Food habits and predation rates data suggested that
wolves were annually preying upon from 11-13%of the moose popu-
lation and from 2-10%of the Nelchina caribou herd.These data
suggested that the percentage of caribou in wolves'diet may be a
function of their availability.As caribou become more abundant,
they make up a greater proportion of the wolves'diet.
The minimum estimated wolf population in the study area ranged
-from a fall high of 80 to a spring low of 40.These wolves were
divided into 13 packs.Pack territory sizes ranged from 346-981
mi 2 and averaged 545 mi 2.An estimated 6-7 packs will be di-
rectly affected by the proposed impoundment and an additional
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14-16 packs could be indirectly affected due to changes in either
moose or caribou movements and numbers.
Wolf harvests in GMU 13 from 1971-1981 ranged from a high of 128
in 1977-78 to a low of 45 in 1980-81.The low harvest in 1980-81
was attributed to poor weather and relatively low wolf densities.
Shooting was the most common harvest method throughout the
period.
Twenty-three wolf den and rendezvous sites were examined.Most
dens were located on slightly elevated,well-drained sites with a
south or east exposure usually near the center of the territory.
Average distance between contiguous natal dens was 28 miles.
Although the number of actual and potential den sites which would
be inundated by the impoundment was not determined it was sug-
gested that loss of the sites would be of minor significance.
Intensive ground observations of active wolf den sites in May and
June of 1980 and 1981 allowed us to detail summer activity pat-
terns and food habits.These observations revealed that wolves
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were present at den sites throughout the day.Helicopters flying
dens always annoyed wolvesi however,they became more toler--near
able over time.Judging from observed behavior patterns and a
review of the literature,it was recommended that all human
ground activities be restricted from a 1.5 mile radius of active
dens.This is particularly important in spring to avoid den site
abandonment.If human activity must occur near dens,these ac-
tivi ties should be limited to early morning and late evening
hours.
The most important potential impact of the Susitna Hydroelectric
Project on wolves would occur indirectly due to reductions in
prey density,part'icularly moose.Di sruption of movements or
reductions in numbers of migratory moose and caribou may reduce
wolf densities long distances from the impoundments.Temporary
increases in wolf density may occur in the project area due to
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their displacement and the displacement of moose and caribou from
impoundment areas.Direct inundation of wolf habitat in the long
term may long lower wolf densities.Addi t~onal .wolf mortality
will probably occur due to increased hunting and trapping activi-
ties resulting from publicity concerning the area's wildlife and
as access becomes developed.
We propose a continuation of all Phase I studies into Phase II to
increase our data base,particularly on packs which occupy im-
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poundment areas which have not been studied and to gather infor-
mation on winter predation rates.Phase I I studies should al so
include a comprehensive investigation of the relationships of all
large predators to their ungulate prey.Of particular importance
are the effects of wolves and bears on moose and caribou calf
_mortali ty.Finally,we suggest that wolf studies be extended to
downstream areas .
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TABLE OF CONTENTS
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SUMMARY
LIST OF TABLES
.LIST OF FIGURES
INTRODUCTION
METHODOLOGY
RESULTS AND DISCUSSION
Pack Histories
Susitna Pack
Susitna-Sinona Pack
Tolsona Pack
Tyone Creek Pack
Fish Lake Pack
Watana Pack
Elevational and Seasonal Usage of Habitat Types
Food Habits
Fish Lake Pack
Susitna Pack
Susitna-Sinona Pack
Tolsona Pack
Tyone Creek Pack
Watana Creek
Summary of Food Habits
Predation
Winter Rates
Susitna Pack
Tyone Creek Pack
Tolsona Pack
Summer Rates
Watana Pack
Wolf Territories l Population Numbers l
and Numbers to be Impacted by the Project
Impacts of Wolves on Moose and Caribou
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1
3
11
15
15
23
25
30
33
34
36
40
42
42
49
49
49
56
56
64
64
66
67
68
68
68
73
79
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TABLE OF CONTENTS (cont'd)
Wolf Hunting-Trapping Mortality
Den Sites
Summer Activity Patterns
Susitna Pack
Watana Pack
POTENTIAL IMPACTS OF SUSITNA HYDROELECTRIC
PROJECT ON WOLVES
PROPOSED PHASE II STUDIES
ACKNOWLEDGEMENTS
REFERENCES
APPENDIX A
APPENDIX B
APPENDIX C
APPENDIX D
APPENDIX E
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83
104
104
114
119
120
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132
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146
163
190
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LIST OF TABLES
Table 1.Summary of statistics associated
with wolf radio-collaring activities
for Susitna Hydroelectric Studies in
GMU 13 of southcentra1 Alaska during
1980 and 1981.
Table 2.Summary of numbers of location obser-
vations of radio-collared wolves by
individual and pack affiliation during
1980 and 1981 in the Susitna Hydro-
electric project area.
Table 3.Summary of individual movements of
wolves 122330 and 122331 from the
Susitna wolf pack f~om 10 May
through mid-November 1981.
Table 4.Habitat type use by month of occur-
rence for the Watana wolf pack in
southcentra1 Alaska from April 1980
through mid-October 1981.
Table 5.Elevation and aspect usage by members
of the Watana wolf pack from April
1980 to mid-October 1981 in the
Susitna Riv.er Basin of southcentra1
Alaska.
Table 6.Chronological summary of kills at
which the Fish Lake wolf pack was
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14
20
37
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LIST OF TABLES (cont'd)
Table 6.cont'd
observed from 19 March 1981 through
26 October 1981 in GMU-13 of south-
central Alaska.
Table 7.Summary of incidence of food remains
in wolf scats collected at a Fish Lake
pack rest area during summer 1981 in
southcentral Alaska.
Table 8.Chronological summary of kills at
which the Susitna wolf pack was ob-
served from January 1980 through
December 1981 in Game Management
Unit 13 of southcentral Alaska.
Table 9.Summary of incidence of food items
in wolf scats collected at the Susitna
wolf den occupied in late spring and
summer 1980 in GMU-13 of southcentral
Alaska.
Table 10.Summary of incidence of food remains
in wolf scats collected at the Susitna
wolf pack rendezvous site occupied
during summer 1980 in GMU-13 of south-
central Alaska.
Table 11.Incidence of food remains in wolf
scats collected at the Susitna den
utilized in late spring and early
summer 1981 in GMU-13 of southcentral
Alaska.
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LIST OF TABLES (cont'd)
Table 12.Chronological summary of kills at
which the Susitna-Sinona wolf pack
was observed from mid-April 1980
through August 1981.
Table 13.Chronological summary of kills at
which the Tolsona wolf pack was ob-
served from 6 January through 17
February 1980 and July 1980 through
8 September 1981 in GMU-13 of south-
central Alaska.
Table 14.Incidence of food remains in wolf
scats collected at the Tolsona wolf
rendezvous site (old Mendeltna 2nd
den)utilized by pack members during
summer 1980 in GMU-13 of southcentral
Alaska.
Table 15.Incidence of food"remains in wolf
scats collected at the Tolsona wolf
den (old Mendeltna rendezvous site)
which was utilized by pack members
in late spring and early summer 1981
in southcentral Alaska.
Table 16.Incidence of food remains in wolf
scats collected at the Tolsona wolf
rendezvous site (old Mendeltna 2nd
den)occupied during summer 1981.
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51
52
53
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LIST OF TABLES (cont'd)
Table 17.Chronological summary of kills at
which the Tyone Creek wolf pack (2
adults,6 pups)was observed from
January through December 1980
in GMU-13 of southcentral Alaska.
Table 18.Chronological summary of kills at
which the Watana wolf pack was
observed from mid-April 1980
through 30 October 1981 in GMU-13
of southcentral Alaska.
Table 19.Incidence of food remains in wolf
scats from the Watana pack den
occupied during late spring and
early summer 1980 in GMU-13 of
southcentral Alaska.
Table 20.Incidence of food remains in wolf
scats collected at theWatana pack
rendezvous site which was used by
pack members during summer 1980 in
GMU-13 of southcentral Alaska.
Table 21.Incidence of food remains in wolf
scats collected at the Watana pack
den occupied during late spring and
summer 1981 in GMU-13 of southcentral
Alaska.
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57
58
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60
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LIST OF TABLES (cont'd)
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Table 22.Comparison between years of food
remains in wolf scats collected at
den and rendezvous sites in 1980 and
1981 from GMU-13 of southcentral Alaska.61
Table 23.Comparison by wolf pack of the annual
percentages of observed prey from
1975 through 1981 in the Susitna and
F Nelchina River Basin of southcentral
Alaska.
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Table 24.-Age,sex,condition (as determined
by percent fat),and cause of morta-
lity of moose and caribou kills exam-
ined in GMU 13 of southcentral Alaska
during 1980.
Table 25.Daily movements,observed prey and
location of the Tolsona wolf pack
from 3 through 26 January 1981 in
GMU-13 of southcentral Alaska.
Table 26.Summary of territory sizes for wolf
packs studied as part of the Susitna
Hydroelectric Project studies during
1980 and 1981 in southcentral Alaska.
Table 27.Estimate of numbers of wolves by
individual pack inhabiting the
Susitna Hydroelectric study area in
spring and fall 1980 and 1981.
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69
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LIST OF TABLES (cont'd)
Table 28.Summary of reported wolf harvests
for Game Management Unit 13 of south-
central Alaska by method of take from
1971 through 1980-81 season.
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84
Table 29.External characteristics of wolf den a
nd rendezvous sites examined in GMU-13
of southcentral Alaska from 1976 through
summer 1981.93
Table 30.Dimensions of hole entrances and
chambers for wolf den and rendezvous
sites examined in GMU-13 of south-
central Alaska from 1976 through 1981.
Table 31.Characteristics of vegetation at
wolf den and rendezvous sites examined
inGMU-13 of southcentral Alaska from
1976 through 1981.
Table 32.Summary of wolf den and rendezvous
sites discovered from 1975 through
1981 occurring within a 50 mile radius
of the proposed Susitna Hydroelectric
Project in southcentral Alaska.
Table 33.Average distance between contiguous
natal wolf dens from 1975 through
1981 in GMU-13 of southcentral Alaska.
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99
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105
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LIST OF TABLES (cont'd)
Page
Table 34.Chronological summary of Susitna
radio-collared wolf observations
conducted from fixed-wing aircraft
in late May and June 1980 in conjunction
with den site studies in GMU 13 of
southcentral Alaska.110
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LIST OF FIGURES
Page
Fig.1.
Fig.2.
Fig.3.
Fig.4.
Fig.5.
Map of Susitna Hydroelectric wolf
study area.
Location of proposed gravel borrow
areas to be used for construction
of the Susitna Hydroelectric Project.
Location of proposed access routes
to the Watana and Devil Canyon hydro-
electric facilities.
Suspected locations and territorial
boundaries of wolf packs inhabiting
the Susitna Hydroelectric Project
area during 1980 and 1981.
Distribution of main Nelchina radio-
collared caribou 1 14 April 1980 through
29 September l 1981 (from Pitcher 1981)
in relation to known and suspected wolf
packs and concentration areas within the
Susitna and Nelchina River Basins of
southcentral Alaska (from this study
and Ballard et al.1981).
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9
10
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Fig.6.Migration routes and movement patterns
of radio-collared moose in the Susitna
and Nelchina River Basins (from Ballard
and Taylor 1980i Ballard and Gardner 1981 1
1981ai Ballard et al.1981)in relation
to known and suspected wolf territories
(from this study and Ballard et al.1981a).80
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Fig.7.
Fig.8.
Fig.9.
LIST OF FIGURES (cont'd)
Relative densities of moose as deter-
mined from stratification and census
flights made on 2 through 4,8 and 29
November 1980 in the Susitna River
Hydroelectric Project study area of
southcentral Alaska (from Ballard et
ale 1981 -upstream moose report).
Diagram of hunting-trapping location
of wolf harvests in GMU-11 and 13 of
southcentral Alaska for 1976-77
regulatory year.
Diagram of hunting-trapping location
of wolf harvests in GMU-11 and 13 of
southcentral Alaska for 1977-78
regulatory year.
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81
86
87
Fig.10.Diagram of hunting-trapping location
of wolf harvests in GMU-11 "and 13 of
southcentral Alaska for 1978-79
regulatory year.
Fig.11.Diagram of hunting-trapping of
location of wolfh~rvests in GMU-11
and 13 of southcentral Alaska for
1979-80 regulatory year.
Fig.12.Diagram of hunting-trapping location
of wolf harvests in GMU-11 and 13 of
southcentral Alaska for 1980-81
regulatory year.
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89
90
LIST OF FIGURES (cont'd)
Page
Fig.13.General location and year of use of obser-
ved wolf den and rendezvous sites dis-
covered in the Susitna Hydro-electric
Project area from 1975 through 1981.91
Fig.14.Occurrence of three radio-collared wolves
at the Susitna wolf pack den studied
from 1 May through 6 June 1980 in GMU 13
of southcentral Alaska.107
Fig.15.Occurrence of lone adult wolves at the
Susitna wolf den from 1 May through 6
June 1980 in GMU 13 of southcentral Alaska.108
Fig.16.Frequency of occurrence of adult wolf
associations at the Susitna wolf den
from 1 May through 6 June 1980 in GMU 13
of southcentral Alaska.108
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Fig.17.Presence or absence of the two adult
members of the Susitna wolf pack at a
den site located in GMU 13 of south-
central Alaska from 19 May through
10 June 1980.
Fig.18.Presence of wolf 308 (Adult alpha
female)at the Watana den site from
7 May through 22 June 1981 in the
Susitna River Basin of southcentral
Alaska (solid line indicate presence
at den).
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INTRODUCTION
Development of hydroelectric power facilities along the Susitna
Ri ver is expected to have detrimental affects on a number of
wildlife species occupying habitats in and adjacent to proposed
reservoirs (Taylor and Ballard 1979).Of particular importance
are the potential effects these developments could have on both
moose (Alces alces)and caribou (Rangifer tarandus)populations.
Equally important are the effects of these impacts on predators
and scavenging species which depend on ungulates for food.Three
large predator.species besides man occur in the Susi tna Basin.
They include gray wolves (Canis lupus),brown bear (Ursus arctos)
and black bear (Ursus americanus).This report concerns studies
conducted to date for determining the potential impacts of
Susi tna hydroelectric development on wolves.
Wolves in Game Management Unit (GMU)13,commonly referred to as
the Nelchina Basin,have been the focus of.interest and study for
over 30 years (Ballard 1981).History of GMU 13 wolves from 1957
through 1968 was summarized by Rausch (1969).From 1948 to 1953
poisoning and aerial shooting by the Federal Government reduced
populations of predators to low levels.By 1953 only 12 wolves
were estimated to remain in the basin.This small population
quickly expanded and by 1965 was thought to have peaked at 400-
450 (Rausch 1969).Although no systematic studies were conducted
from 1969 through 1974,McIlroy (1976)suggested that a second
population peak occurred in 1970.
During the period of wolf population growth,moose populations in
GMU 13 declined suggesting a cause-effect relationship.Subse-
quently in 1975 a series of predator-prey relationships studies
involving wolves were initiated.Results of these studies were
provided by Stephenson (1978),Ballard and Spraker (1979),
Ballard and Taylor (1980),Ballard et al.(1980)and Ballard et
al (1981a and b).Portions of the aforementioned studies in-
volved experimentally manipulating wolf densities in part of the
area which could be impacted by Susitna hydroelectric development
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(Ballard et al.1980).Wolf control activities were conducted
from 1976 through July 1978.By 1980 wolf densities in the re-
duction area had returned to pre-control levels (Ballard 1980)
and thus studies to determine the potential impact of hydro-
electric development on wolves would not be influenced by the
earlier wolf control activi ties.Description of the proposed
Devil Canyon and Watana Dam projects have been briefly described
elsewhere (Taylor and Ballard 1979).
Obj ectives of Susi tna hydroelectric wolf studies during Phase I
were as follows:
(1)To identify wolf packs occupying areas that will be impacted
by the Susi tna Hydroelectric Proj ect.
(2)To delineate the territories of each pack and identify den
si tes,rendezvous sites and maj or feeding areas.
(3)To determine the numbers of wolves and rates of turnover for
each pack.
(4)To determine the food habi ts for each pack.
Data collected from earlier and ongoing GMU 13 wolf studies were,
in some cases,combined with those collected during this study in
an effort to provide a better understanding of wolf ecology in
the Susi tna Basin.
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METHODOLOGY
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).
Wolves were immobilized with either 2 to 2.5 mg of etorphine
(M-99,D-M Pharmaceuticals,Inc.,Rockville,MD)or a combination
of Icc phencyclidine hydrochloride (100 mg/cc,sernylan,Parke-
Davis Co.)and Icc of promazine,hydrochloride (100 mg/cc,
Sparine,Wyeth Laboratories).After being processed and radio-
collared,each wolf which had been immobilized with etorphine was
given an equivalent cc dosage (2 mg/ml)of the antagonist
diprenorphine (M 50-50,D-M Pharmaceuticals,Inc.,Rockville,MD)
which was injected into the radial vein.No antagonist is avail-
able for sernylan.A paper comparing M-99 with sernylan for im-
mobilizing wolves was prepared and accepted by the Journal of
Wildlife diseases (Appendix A).
Captured wolves were equipped with an adjustable radio collar
made of fiberglass and urethane manufactured by Telonics (Mesa,
AZ).Blood samples were taken from each wolf using methods simi-
lar to those described for calf and adult moose (Ballard et al.
1979).Blood samples were shipped frozen to Pathologist Central
Laboratory in Seattle,Washington for SMAC analysis and protein
electrophoresis.When practical,the following body measurements
were recorded:Weight,total length,hearth girth,chest height,
neck circumference,shoulder height,tail length,and length of
canines.
Radio-collared wolves were tracked and,when possible,visually
observed from fixed-wing aircraft using the methods described by
Mech (1974).Radio signals were received with a programmable
scanning receiver (Telonics).Monitoring intensity was variablerbutanattemptwasmadetolocateeachpackonceortwice/week.
Approximate ages of captured wolves were determined on the basis
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of tooth eruption and wear.Estimates of the ages of wild wolves
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were based upon their relative size and by criteria described by
Jordan et al.(cited by Mech 1970).In some cases,age and sex
structures of certain packs were not ascertained until the ani-
mals had been killed by hunters and trappers.Hunters and trap-
pers were encouraged to provide the Department with wolf car-
casses taken in Unit 13 by offering $10.00 per carcass.Ages of
harvested wolves were determined by both tooth eruption and wear,
and by examining epiphyseal cartilage of the longbone according
to methods described by Rausch (1967).
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-col-
lared wolves which had dispersed were not included.Sizes of
wolf territories and study areas were determined with a compen-
sating polar planimeter.All study areas and wolf territories
were planimetered at least three times and then averaged to com-
pute mi 2 (km2 ).This method was selected to provide estimates
comparable to those reported in other published studies.
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 autoclaved 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 re-
mains.Comparisons of hair scale impressions were made with know
samples by imprinting them on a slide containing clear fingernail
polish.
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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.(1979).A femur or meta-
tarsal and the mandible were collected from each kill to aid in
establishing the animals physical condition on the basis of per-
cent 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 January through April 1980 an attempt was made to locate
and examine all kills made by selected radio-collared wolf packs
during a 2-3 month period.An attempt was made to radio-locate
these packs every other day and to backtrack them to their pre-
vious location by following tracks.
In early May 1980 two members of the Susi tna wolf pack and in
1981 seven members of the Watana wolf pack were equipped with
activi ty transmitters (Telonics)in an effort to determine the
daily activity patterns of a denning wolf pack.Each activity
transmitter was equipped with a tip switch which altered the
pulse rate of the transmitter which was dependent on the position
of the animal's head.When the animal's head was down,the pulse
of the radio decreased and conversely when the animal was stand-
ing,the pulse rate increased.A semi-permanent monopole an-
tennae was erected ~mi Ie away from the pack's den si te .Both
amp Ii tude and period of each radio transmitter was monitored
during the denning season with a portable digital data processor
(Telonics TDP-1)which was connected to a portable programmable
scanning receiver and a rustrak recorder (Gulton Inc.,Man-
chester,N.H.).All three instruments were powered by a 12volt
battery,all of which were housed in a large plastic container
for weather protection.While activity patterns were moni tored
electronically,the Susitna den area was observed from a blind
for a 31 day period in 1980.The Watana den was observed for a
47-day period.During ground observations the presence or ab-
5
sence of radio-collared animals was determined by manually scan-
ning the den site area with a hand-held antenna (Ballard et al.
1977).Both ground and aerial observations were used to verify
activity data which were plotted on the rustrak recorder.
The recorder continuously plotted radio signal information on a
paper spool at the rate of 8 inchesjhour.A separate bench mark
transmi tter was used as a control to calibrate the receiver so
that a continuous comparison could be made between the known lo-
cation and activi ty of the bench mark transmitter to data col-
lected from the wolf transmi tters.
Descriptions of the proposed Devil Canyon and Watana Dam projects
have been described elsewhere (Taylor and Ballard 1979)Because
moose are the principal wolf prey,the boundaries of the wolf
study area were the same as those described for upstream moose
studies except that some packs were studied outside the boundary
area because of wolf dispersal from the primary study area.
Boundaries of the primary study area were as follows:
The Denali Highway on the north to its confluence with the
Maclaren River on the east,the r-iIaclaren River to its confluence
with the first unnamed creek in R4E,T13N (Gulkana Quad)upstream
to Monsoon Lake,then a straight line to Tyone Village continuing
up Lake Louise to the Lake Louise Road to its intersection with
the Glenn Highway,on the south the Glenn Highway to the Little
Nelchina,then upstream to the peak of the Talkeetna Mountains,
on the west the upper elevations of the Talkeetna Mountains to
the confluence of the upper north and south forks of the
Talkeetna River,then northwest to the mouth of Portage Creek,
then upstream of Portage Creek to its headwaters to the head-
waters of Brushkana Creek to its confluence with the Denali
Highway (Fig.1).
Vegetation,topography and general climate of the area has been
described by Skoog (1968),Bishop and Rausch (1974),and Ballard
(1981)and thus no further descriptions are needed until vegeta-
tion studies under Subtask 7.12 are completed.
6
-
-
_.
-
.......
Fig.1.Map of susitna Hydroelectric wolf study
For purposes of this analysis we assumed the proposed proj ect
-.._-----------._---
would consist of a 2-dam system;a dam on the Susitna River be-
tween Deadman and Tsusena Creek;and a dam on the Susitna River
at Devil's Canyon.The upper Watana impoundment would inundate
an area up to 2,200 foot elevation,while the Devil's Canyon im-
poundment would inundate an area up to the 1,450 feet elevation.
Characteristics and timing of drawdown and fill were not avail-
able for this report.
At the time this report was written the plan of construction was
poorly understood.Final decisions on locations of access
routes,barrow pits,construction camps,etc.,had not been made.
Therefore,impacts of a range of al ternatives were considered.
Location of proposed borrow areas was provided by Acres American,
Inc.,in October 1981 (Figure 2),however,details concerning
access,volume of gravel to be extracted,method of extraction,
etc.,were not provided.Proposed alternative access routes are
depicted in Figure 3.Apparently any of the routes could be used
for ei ther a road or in some cases a railroad bed.
Preliminary plans call for 5,000 construction workers at height
of construction,requiring at least two or more camps.Operation
of the project will include an unknown number of bUildings and
permanent housing for approximately 120 permanent personnel.
8
-.
Figure 2.Location of proposed gravel borrow areas to be used for
construction of the Susitna Hydroelectric Project.
9
.~
!U8-
u.S.DEPARTMENT OF ..TEHIOR
FISH All)Wl.Dl.IfE SERVICE
UPPER SUSITNA RIVER BASIN
TWO DAM.PLAN
\
I
\-."~<~1/"
)
\\
l
\-.r?J-.,.,
/
r-./
.-'_-1
.Jc:,
\
"
J
~,p-C1
(-""1
\
)---~
r-_\-"--..J "\
.,,
I
(
'""'l
rr-------
I
I
"/
o
/0
J ........
'"'t
t'-\.."
($)"1 '"\
\S}..""'.....~
6\'l:-.'~'t"11»;.\
J
I
""r
\
),
I
20M,...
....-......-..................-...
UPPER SUSITNA RIVER PROFILE
11M.llLEI 1Z0-ZtO
$CALE
Q
,---.-··10--15
.......o
I~
I
OCTOBER 1111
Figure 3.location of-proposed access routes to the Watana and Devil Canyon hydroelectric facilities.C(IIPS QI'IHGMlRS IlK MAP
J J t J J 1 J I 1 J
......
-
RESULTS AND DISCUSSION
From 20 February 1980 through May 1981,36 gray wolves from six
individual packs were captured and radio-collared for this study.
Seven wolves were recaptured on one or more occasions for recol-
1aring.Twenty-one (57%)of the captured wolves were males (6
pups and 15 adults)and 15 (43%)were females (7 pups and 8
adults (Table 1).Six of the 23,were recaptured from earlier
studies.Induction and recovery times of wolves immobilized with
00-99 are compared to those of wolves immobilized with Sernylan in
Appendix A.
Morphometric measurements and blood analysis of captured wolves
are in the process of being entered in to a computer and there-
fore,except for weights,packed cell volume,and percent hemo-
globin,will not be presented at this time.Data from this and
earlier wolf studies (Appendix B)will be combined and analyzed
jointly to describe morphometric growth and development in this
wolf population.
One thousand one hundred and seventy-five individual radio loca-
tions were obtained for the 36 radio-collared wolves from January
1980 through October 1981 (Table 2),yielding an average of ap-
proximately 33 locations per animal.A total of 2,255 wolf
sightings were made while locating the radio-collared packs which
represented 437 pack days (pack day is defined as any day on
which one or more members of a pack was located on'e or more
times).
Radio contact with at least four and perhaps as many as six wolf
packs occupying habitats along the Susi tna River near the pro-
posed impoundments was not established during this study.Lack
of adequate snow cover and clear sunny weather seriously hampered
capture efforts between spring 1980 and fall 1981.With the ex-
ception of the Watana wolf pack,radio contact with the packs
studied was possible only because a few members o'f each pack had
functioning radio collars from earlier studies facilitating loca-
tion of the packs for additional collaring.
11
Table 1.Summary of statistics associated with wolf radio-collaring activities for Susitna Hydroelectric studies in
GMU 13 of southcentral Alaska during 1980 and 1981.
Drug Induction Antagonist Reverse Packed
qrig.Pack Accession Date Est.Weight Dosage Location of Time Dosage Time Cell %
Affiliation Number Captured Age Color Sex (lbs)(mg)Injection (min)(cc)(min)volume Hb
--
Fish Lake 122326 04/14/81 Adult Gray F 92 M-99 2.5 Top back 5.0 2.5 2.0 49.3 16.7
122327 04/14/81 Pup Gray M 94 M-99 .2.5 Rump 8.0 2.5 10.0 54.3 18.8
122329 04/17/81 Pup Gry/Blk F 57 M-99 2.5 '1 5.0 2.5 2.0
Susitna 122229*04/16/80 Yrl.Gray M 105 M-99 2.5 Left rump 5.5 2.5 1.7 57 20+
122295*03/20/80 4 yr.Gray F -Sernylan:1:1 7 -N/A N/A
Sparine
04/15/80 100est M-99 2.0 Left leg 7.0 2.0 1.0 49.5 17.9
122296*02/20/80 6-7 yr.White M 110est Sernylan:1:1 '1 -N/A N/A
Sparine
I-'122302 02/20/80 Pup Gray F -Sernylan:1:1 --N/A N/ANSparine
10/16/80 Yrl.75 M-99 2.0 Tail 5.0 2.5 1.5 63 20+
122303 02/20/80 Yrl.Gray M 80 Sernylan:2.0 Left rump -N/A N/A
Sparine
122305 04/13/80 3 yr.Gray M 100 M-99:2.0-.25 7 8.0 3.0 1.0 55 20+
Rompun
10/16/80 106 M-99 2.5 Right rump 5.5 2.5 2.0 68 20+
122306 04/13/80 Pup Gray M 85est M-99:2.0:4 7 8.0 3.0 1.0 53 18.9
Rompun
122330 4/17/80 Yrl Gray M 97 M-99 2.5 Back 8.0 2.5 2.0
On ground 1.0
122331 4/17/80 Yrl Gray M 105 M-99 2.5 Rump 3.0 2.5 2.0
Susitna-
Sinona 122312 10/14/80 Pup Gray H 46 M-99 2.5 Top back 5.0 2.5 .8
122313 10/14/80 2-3yr Gray M 106 M-99 2.25 Left leg -2.5 1.0 63 20
Tolsona 122220*07/06/80 2.5yr Black F 68 M-99 2.25 Front leg 20.0 2.5 1.0
2nd dart 1.5 7 3.0
122315 10/16/80 3-4yr Black F 78 M-99 2.5 Top back 5.0 5.0 5.3 68 20+
2nd dart 2.5 '1
122316 10/16/80 Yrl Gray H 82 M-99 2.5 7 6.0 2.5 1.5 62 20+
122210*04/14/81 Adult Gray M 97 M-99 2.5 Leg 11.0 2.5 12.0 57.9 19.2
122328 04/14/81 1-2yr Gray F 70 M-99 2.5 Back 7.0 2.5 6.0 58.9 19.3
~J J J .~
.II I 1 )J ]!I J )..__1
--Cl J J 1 :"""]1 .~J -]
Table 1.(cont'd)
Drug Induction Antagonist Reverse Packed
Orig.Pack Accession Date Est.Weight Dosage Location of Time Dosage Time Cell %
Affiliation Number Captured Age Color Sex (lbs)(mg)Injection (min)(cc)(min)volume Hb
Tyone 122215*02/20/80 3-4yr Gray M 111 Sernylan:1.0:1.0 1 1 N/A N/A
Sparine
122216*02/20/80 3-4yr Gray F 95 Sernylan:1.0:1.0 1 1 N/A N/A 51 20
Sparine
122298 02/20/80 Pup Gray F 84 Sernylan:1.0:1.0 1 1 N/A N/A 44 20
Sparine
10/16/80 1.5yr 82 M-99 2.5 1 12.0 2.5 .8 56 19.5
2.5 1 12.0
122299 02/20/80 Pup Gray F 82 Sernylan:1.0:1.0 1 1 N/A N/A
Sparine
I--'122300 02/20/80 Pup Gray M 93 Sernylan:1.0:1.0 1 1 N/A N/A 48 20wSparine
122301 02/20/80 Pup Gray M 100 Sernylan:1.0:1.0 1 ?N/A N/A
Sparine
122317 10/16/80 2-3yr Black M 1 M-99 2.5 Left shoulder 40 2.5 1.0 60 20+
Watana 122308 04/24/80 3-4yr Gray F 91 M-99 2.5 Top back 17.0 8.0 5.0 47 25+
122309 04/19/80 Pup Gray F 79 M-99 2.0 Chest cavity 4.0 4.0 .8 52 20+
122310 04/23/80 2yr Gray M 101 H-99 2.0 Left side 26.0 5.0 1.5 52 20+.
2.0 Back
03/06/8'1 107 M-99 2.5 Rump 4.0 2.5 1.0 62 20+
05/10.81 102 M-99 2.5 Shoulder 10.0 2.5 -62.8 19.4
122311 04/23/80 2yr Gray M 112 M-99 2.0 1 1 5.0 2.0 50 20
122323 03/07/81 Pup Gray F 93 M-99 2.5 Shoulder 9.0 2.5 1.0
122324 03/07/81 Pup Gray F 90 M-99 2.5 1 3.25 2.5 1 53 20+
05/10/81 Yrl Gray 78 M-99 2.5 Back 3.0 2.5 1.0 54.4 16.8
122325 03/07/81 Pup Gray M III M=99 2.5 Shoulder 2.0 2.5 1.5 50 20+
05/10/81 Yrl 99 M-99 2.5 Side 8.0 2.5 2.0 55.7 17.7
122344 05/10/81 Yrl Gray M 64 M-99 2.5 Back 9.0 2.5 1.0 52.2 17.8
122345 05/10/81 Yrl Gray M -M-99 2.5 Side 10.0 2.5 1
122346 05/10/81 Yrl Gray F 77 M-99 2.5 Side 5.0 2.5 1 56.0 18.4
*Recapture
Table 2.Summary of numbers of location observations of radio-collared wolves by
individual and pack affiliation during 1980 and 1981 in the Susitna
Hydroelectric Project area.
Pack Accession No.Radio No.Wolf No.
Name Number Locations Sightings Pack Days
Fish Lake 122326 10 8 14
122327 7
122329 8
122300 10
Subtotal 35
Susitna 122295 110 536 126
122296 9
122302 81
122303 9
122305 33
122306 39
122330 14
122331 21
Subtotal 316
Susitna-Sinona
122229 27 94 33
122312 3
122313 11
Subtotal 41
Tolsona 122220 78 798 83
122315 41
122316 25
122210 21
122328 8
Subtotal 173
Tyone 122215 31 323 77
122216 11
122298 42
122299 37
122300 41
122301 13
122317 5
Subtotal 180
Watana 122308 88 436 104
122309 14
122310 61
122311 18
122323 50 ~
122324 43
122325 48
122344 25
122345 42
122346 41
Subtotal 430
Totals 37 1,175 2,255 437 ~
14 -
....
r
:1
r
Pack Histories
Histories of individual radio-collared wolves and their respec-
tive packs which either occupy areas to be inundated or areas
where prey species would be affected are described in the fol-
lowing section.
Individual wolves are identified in the text by the last three
digi ts of their assigned accession number.Data collected by
\
Ballard et ale (1981)before the initiation of this project are
included.An abstract of this latter report is provided in
Appendix B.
Susitna Pack
Radio contact with this pack was established in February 1979
within the Deep Lake pack territory (Ballard et al.1981).At
that time the Susitna wolves were observed fleeing from the car-
cass of the Deep Lake female (#009)which they had just killed.
Reasons for thi s conflict and the fate of an uncollared gray
which had been accompanying wolf 009 are not known,particularly
since thi s Susi tna pack never returned to the Deep Lake area.
When collared,the pack was comprised of at least two adults and
seven pups.On the basis of size and later capture records,a
tenth wolf,an adult male was suspected to have been in the pack.
Following initial capture,the pack moved to the area south of
the big bend in the Susitna River.Whether these wolves had al-
ways occupied the area west of Lake Susitna and Tyone is unknown,
but seems likely based on the identified gaps between territories
of other packs for the period 1975 through 1978 (Ballard et ale
1981).By late spring 1979,the pack numbered six or seven.
Pack losses between fall and spring were probably the result of
one or two wolves being shot and at least one dispersal.
The pack was first observed at the 1979 den site on 13 April.At
least six pups were raised but were not observed until 3 August.
15
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 towards the Wrangell Mountains.In·late
January 1980,wolf 229 1 a yearling male l and at least two gray
associates appeared to follow the caribou migration and dispersed
to the east.This group was subsequently referred to as the
Susi tna-Sinona:pack and its history from January 1980 through
mid-October 1981 is provided in the Susi tna-Sinona portion of
this report.By early February 1980,the original Susitna pack
numbered seven.
In early March 1980 the pack was reduced by two members during a
conflict wi th the Tyone pack.Details of this conflict follow:
On 8 March while during a study of predation rates l wolf 295 (the
adult gray female of the Susitna pack)was tracked to a location
2 miles (3.2 km)south of Vermillion Lake.She was alone.By
backtracking her in the snow for several miles to the west to the
confluence of Sanona and Tyone Creeks,seven additional wolves
were located in one area.This was more than had been observed
in the Susi tna pack during the previous 2 weeks.A check of
other wolf radio frequencies revealed that radio-collared members
of the Tyone pack l which was compri sed of two adults and six
PUPSI were also present.A search was made for other radio-col-
lared members of the Susitna pack and wolf 296 1 the light adult
gray male which on the basis of tail posture and leadership in
the pack was assumed to be the alpha male l was found dead 0.5
miles (0.8 km)north of Tyone Creek.A SUbsequent examination of
wolf 296 revealed puncture marks on the neck and shoulders.In
addition l at least seven distinct wolf trails radiated from the
areal leaving little doubt that wolf 296 had been killed by other
wolves.Following blood in the snow wolf 296 was backtracked to
the location where the struggle had begun.At this site a fresh
adult moose kill was discovered.
16
-
-
-
-
-
-
--
.....
I
At the moose kill there were at least two wolf beds in the snow
approximately 20 feet away.A Inoose fetus,a dead ptarmigan
(Lagopus sp.)and two wolf beds were also found 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.Approximately 100 yards separated the carcass of the moose
and the carcass of wolf 296.
One of the wolf trails radiating from the moose kill site was
\
spotted with blood.This trail was followed for approximately
0.25 miles upstream where the trails of four wolves carne to-
gether,suggesting that an apparent pur sui t continued.Approx-
imately 0.25 miles (0.4 krn)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 according to signs in the snow.
Wolf 302 of the Susi tna pack was radio-located at 1130 hr.3.5
miles east of the carcass of wolf 296.When originally located
at 1030 hr.she had been within 0.25 miles (0.4 krn)of the Tyone
wolves.At this time the location of four of seven Susi tna
wolves and all eight Tyone wolves was known.
As we left the site an additional fresh calf moose kill was ob-
served 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 wolves 295 and 302 of the Susi tna pack were lo-
cated.Wolf 295 had moved to the east side of the Tyone River.
Wolf 302 was within 5 miles (8 krn)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,apparently with punctures in the neck and around the
ears.
17
Based upon ground and aerial observations,it was concluded that
-----_.----_.-._-
the Susitna pack had come upon a moose kill made by the Tyone
wolves.The moose kill was located on the territory boundary of
each pack .Therefore,in addition to competition for possession
of the kills,the conflict may have involved a territorial dis-
pute.The Susitna area appeared to have a relatively poor avail-
abili ty of prey in comparison to the Tyone pack terri tory during
this winter.Comparison of prey abundance between the two areas
will be discussed in the predation rates section of this report.
Although the literature indicates that conflict with and between
pack members occasionally results in wolf mortality,all reported
incidences have involved only one mortali ty.
Following the dispersal of wolf 229 and its associates,and the
deaths of wolves 296 and 303,the Susi tna 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,two of four remaining pack mem-
bers (#'s 295 and 305)were fitted with activity radio trans-
mi tters.In addi tion,ground observations at the den site were
made from 1 May through 6 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.Preliminary results of this 1 month study are
presented in the summer activities section of this report.
The pack moved the pups to the first rendezvous site,located
approximately 0.75 miles west of the den site between 4 and
6 June.A second rendezvous site,located approximately 5 miles
to the northwest,was briefly occupied between 1 and 7 August.
No pup morta1i ty was observed in 1980.In addition no trapping
or hunting mortality occurred in 1980 due to poor snow condi-
tions,and,thus,the pack was still comprised of 10 wolves by
the end of February 1981.
18
....,
r,,
During March and April 1981 wolf 295 was frequently observed
alone away from the remainder of other members.By mid-Apri 1
pack size had declined to 6 wolves,probably due to either dis-
persal,unreported harvest,or both.Radio contact with wolves
305 and 306 was lost between February and late March 1981.In
mid-April wolves 330 and 331 (both yearlings)were captured and
radio-collared while accompanying wolf 302 and one uncollared
gray.
On 10 May wolf 331 was observed on the northern edge of the
Susi tna pack terri tory accompanied by two gray wolves from the
Watana pack (=If 's 310 and 323)and four unradio-collared grays
which could have come from either pack.Both Watana wolves (see
Watana section,this report)were well outside of their normal
home range.During this brief observation,members of the two
packs interacted with one another apparently with no mortality or
injury to any individuals.
After members of the two packs had been observed together,wolves
330 and 331 were not subsequently observed with the remaining
Susi tna Pack members.Instead,they slowly dispersed to the
west,generally inhabiting the upper reaches of the Black and
Oshetna Rivers through mid-summer.Table 3 summarizes the indi-
vidual movements of wolves ~30 and 331 from 10 May through mid-
November 1981.From 10 to 26 May both wolves traveled indepen-
dently.The number of associates accompanying both wolves sug-
gested that 1-2 other grays from the Susi tna Pack were also in
the process of dispersing.During 1981 the Nelchina caribou herd
calved in the Oshetna and Black River.area.Both wolves were
frequently observed in close proximity to caribou and we suspect
that they preyed heavily upon them.
By mid-October wolves 330 and 331 had separated.Wolf 331 was
observed near the mouths of Coal Creek and on Clearwater Creek
during October,accompanied by one uncollared gray.By mid-
November wolf 331 and we assume the uncollared gray (determined
to be a 2 yr.old yearling male)were found near Stephan Lake;
19
Table 3.Summary of individual movements of wolves 122330 and 122331 from the Susitna Wolf Pack from 10 May thru mid-November 1981.
WOLF 122330 WOLF 122331
Dates
&Tillie
Distance be-
tween (obs.&
appx.direc-Final
tion traveled Location
iI Wolves Misc.
observed Note
Dates
&Time
Distance be-
tween (obs.&
appx.)direct-Final
tion traveled Location
fI Wolves
observed
Misc.
Note
5/10-5/26
1730-1730 32 W Gilbert Ck
5/10-5/11
gray alone 1730-1235 ~_SW Lone Butte 3 gray
killed calf
moose
10/17-11/uncollared
gray alone 1515 WSR StephaI!nl.~k~__2 g~_gray ID as "
5/30-6/4 Wolf 330
gray Wolf 331 present 1500-2J05 2_7_N Gilbert Ck 3~ay __present
5/29-5/30
1400-1500 0 -Landslide Ck No ""
8/20-10717 Mouth L.Clear
gray alone 1455-1515 18 NW Water Ck._2 gray
"
near beaver
house
chasing
caribou
caribou present
2.5 mi from 330
3 gray
No
2 gray
2 gray
up Tyone Ck 2 gray
Lake Oshetna R
TyoneCk 3 gray
Landslide Ck
"
"
"
"
"
"
"
"
"
"
SW
E
WSW
SW
"
"
"
"
"
"
8
"
"
"
"
16
5/11-5/12
1235-1750
5/12-5/13
1750-1800
5/13-5/16
1800-0915
675-6/6
1900-1100
6/4-6/5
2105-1900 same as wolf 330
6/15-6/16
1408-0958
5714
0740
6/13-6/15
1118-1408
6/12=6/13
1913-1118
6119-8/20 Mouth of
1127-1455 40 NW Coal Ck.
6/11-6/12
0852-1913
6/16-6/19
0958-1127 " " " "
"
"
"
"
"
"
"
"
""
"
"
alone on calf caribou 5/16~5/29
kill 2.5 mi from 331 0915-1400 t4
Wolf 331 present &3
caribou 1 moose
present
gray Wolf 331 present
gray
Wolf 331 present &
gray 4 caribo~pre6ent
gray
gray
gray
gray
Wolf 331 present &
gray 100 caribou_present
Roaring Ck.1 grayS
N Gilbert Ck 3
Clarence Lk
NE Ck 3
SW ---2
ESE Susitna Lake 3
W upper Bl.R.2
-
NE BI.R.3
SW Gold Ck.3
NE Oshetna R.3
NW Bl.Lk.3
NW Brushkana Ck.
Brushkana Ck •
5/26-5729
1730-1400 25
~5/29-6/4
1400-2105 25
6/4-6/5
2105-1900 8
6/5-6/6
1900-1100 3
6/6-6/11 36
1100-0852
6-11-6/12
0852-1913 35
6/12-6/13
1913-1118 6
6/13-6/15
1118-1408 12
6/15-6/16
1408-0958 6
6/16-6/19
0958-1127 10
6/19-10/17
1127-1540
10/17-
1540
.,.",
/
l ,J ;~_J _J I .1 J )a ••
),.J •J(
....
I
.-
-I
.....
.-
I
approximately 76 miles to the west from the Susi tna Pack terri-
tory.During the same time interval wolf 330 was observed alone
at Brushkana Creek where he remained through November.Whether
both of these wolves have completed their dispersal is not yet
known.
Wolf 295 was first observed at the wolf den,which had been uti-
lized in 1979 and 1980,on 23 May 1981.Since she had not been
radio-located earlier in the month we do not know the exact date
of when she regularly began visiting the site;however,she was
observed within 3 miles of the den in late March.Between 23 May
and 29 May wolf 295 was observed at the den site accompanied by
from 1 to 2 grays (including female wolf 302).From these obser-
vations we assumed she had whelped pups or was in the process of
doing so.At 10:30 a.m.,on 30 May wolf 310 from the Watana Pack
was located at the Susi tna den site.No other radio-collared
wolves were at the site.Interestingly,on 28 May wolves 310,
344,345 and one uncollared gray had been observed within 2.5
miles of the Susitna den (see W~tana Pack Section).After 28 May
the three Watana wolves were no longer observed together and
wolves 344 and 345 appeared to disperse from the area.Wolf 295
and one uncollared gray were observed resting approximately 8
miles northwest of the den site,on the same day,while 302 was
observed alone,S miles north of the den.By 4:30 p.m.,wolf 295
had returned to the den site while 302 remained about 2 miles NE
from the den.Because wolf 310 had been observed at the Susitna
den,we speculated that either he had been accepted into the
Susi tna Pack or that some sort of conflict was occurring.
On 31 May at 1515 wolf 310 was again located at the Susitna den,
while wolves 295 and 302 were located 5 miles north of the den.
By 0750 hr.,on 1 June wolf 302 had returned to the den site
while wolf 295 was observed bedded approximately 9 miles north-
west of the den site.At 1738 the same day,however,wolves 295
and 310 were both present at the den site and wolf 302 could not
be located.On both 2 and 3 June wolf 302 was again at the den
site while wolf 295 was observed approximately 6.5 miles bedded
21
to the northwest within about 0.5 mile of wolf 3l0's location.
Between 3 and 6 June both wolves 295 and 302 were periodically
observed at the den site while the location of 310 was unknown.
On 6 June wolf 310 was observed alone at the Susitna 1980 rendez-
vous site while wolves 295 and 302 were located approximately 3
miles north of the den,separated by about 2 miles.Wolf 344
also from the Watana Pack was also present in the area but he was
not accurately located.The next day wolf 302's location had not
changed and upon intensive searching from fixed-wing aircraft,we
observed that she was dead.Wolf 302 was examined on 8 June.
Her carcass was founG lying on its right side at the base of a
clump of small black spruce.Several puncture wounds were evi-
dent along the sternum of the chest cavity.At least one and
perhaps two of the punctures had pierced a lung.There were also
puncture holes on the inside of both rear legs.Except for the
punctures which pierced the lung,the wounds did not appear
fresh.Based upon the size of the puncture marks and the recent
interactions with wolf 310,it appeared wolf 302 had been killed
by other wolves,probably 310.
Following the death of 302,wolf 295 did not return to the den
site and was observed either alone or accompanied by 2 uncollared
grays through the remainder of June.Wolf 310,however,was
again observed at the Susitna den on 9 and 10 June accompanied by
one uncollared gray.
During July and early August no radio-locations were obtained on
ei ther wolf 295 or 310 due to poor weather conditions.By 20
August 1981 both wolves were observed together.Between late
August and mid-October they were continually observed together
and we suspected that perhaps a bond had been formed.The fate
of the two remaining uncollared members of the Susitna Pack (ex-
cluding wolves 330 and 331)is unknown.
Unfortunately,because of poor weather and work schedules,we
were unable to examine the Susi tna den until late August.At
that time we were unable to find·any significant evidence con-
22
-
.....
.!fPI!l
firming the presence of pupsJ~h<:?w~~~r!__o~e pup scat was found at
the entrance to the rendezvous site,located several hundred
yards to the northwest.Because of wolf 295's frequent presence
at the den during late May we suspect that pups were present and
that they and wolf 302 had been killed by wolf 310.Since no
pups had been observed from aircraft,they probably died before
they were old enough to spend much time outside of the den and
few,if any,scats would have been present.
Reasons for the conflict between wolf 310 and the Susi tna Pack
are not known andean only be speculated upon.However,the
Susitna pack appears to have disintegrated in part due to abnor-
mal family structure.To our knowledge,when wolf 310's presence
was first confirmed,the Susi tna pack was without an alpha male.
and was comprised of yearlings and two-year-olds (in 1980 the
alpha male was killed by the Goose Creek Pack and in 1981 the
alpha male was shot by aerial trappers).Thus many of the pack
members were of an age when dispersal normally occurs (Ballardet
al.1981).Perhaps these relatively young wolves were unable to
defend the den site from an older and perhaps more aggressive
.wolf,such as 310.If wolf 310 did in fact kill the Susitna
pups,it is uncertain why wolf 295 and 310 would then start ac-
companying one another.
At the time this report was prepared wolves 295 and 310 appeared
to be the only wolves occupying the Susi tnaterri tory .
From February 1979 through mid-October 1981 the Susitna pack oc-
cupied an area of 561 mi 2.Their territory ranged from Sinona
Creek on the west to the headwaters of the West Fork of the
Gulkana River on the north to Minnesota Lake on the east and to
Moose Lake on the south.
Susi tna-Sinona Pack
As mentioned in the Susitna Pack history section,wolf 229,which
had originally been capturectand radio-collared as a pup in April
23
1979,was a regular member of the Susitna Pack until late
December 1979.In December 1979 and January 1980,wolf 229 and
at least two other unradio-collared grays from the Susitna pack
appeared to follow migratory caribou eastward to the Gakona River
and did not return to the Susitna Pack.From 6 to 9 January 1981
they moved 24 miles northeast to the West Fork of the Gulkana
River and by 11 January had moved an additional 17 miles east to
an area just north of Hogan Hill.After that date they remained
in that area and by 25 January were accompanied by a fourth un-
collared gray.In mid-April 1980 wolf 229 was recaptured and its
transmitter was replaced.We also attempted to capture a second
wolf which we suspected was an adult pregnant female,but the
.wolf escaped by running into a hole at the old Sinona den site
which had been used by the original Sinona Pack in 1978 (Ballard
et al.1981).
During May and June 1980 we frequently radio-located the wolves
in the vicinity of the old Sinona den site,but subsequent ground
examination in September 1980 suggested it had not been used and
thus we were unable to locate a den site.We did conclude,how-
ever,that the pack denned because in mid-October the pack was
comprised of 4-5 wolves,2 of which were judged to be pups-of-
the-year.At that time two additional wolves were captured and
radio-collared;a female pup ·(#312)and a 2-3 year-old adult male
(#313).We suspected that a third wolf had been struck by a dart
but it escaped.
Following the capture of wolves 312 and 313 only four wolves were
observed in the pack.On 19 November 1980 wolf 312 was killed by
a trapper,leaving the pack with 3 wolves,for the remainder of
the winter.
On 31 March 1981 wolf 313 was observed alone at the 6,000 foot
elevation of Nathlie Mountain,approximately 36 miles from the
pack territQry.By 13 April,however,he was observed alone back
in the pack area,but apparently soon left again since we could
not locate its radio signal.Sometime between 13 April and 23
24
-
~,
-
-
July 1981 wolf 313 traveled to~h~_~~l~o~~Pack area,a straight
line distance of approximately 52 miles,where he became a regu-
lar member of that pack.As of mid-October 1981,wolf 313 was
still associated with the Tolsona Pack,but its radio signal was
deteriorating quickly.While wolf 313 was dispersing,wolf 229
apparently also began to disperse.
Wolf 229 was last located in the Susitna-Sinona pack area on 23
April 1981.Subsequently,we lost radio contact with him until 4
August 1981,when he was observed alone,traveling west approxi-
mately 3 miles west of the big bend in the Maclaren River,a
straight line movement of approximately 56 miles.His subsequent
whereabouts were unknown as of mid-October 1981.
From February 1980 to April 1981 the Susitna-Sinona Pack occupied
an area of 466 mi 2 •The area was bounded by the main fork of the
Gulkana River on the west,the Copper River on the south,the
Gakona River on the east,and a line from Roundtop Mountain to
the Chistochina Hills on the north.This area corresponds very
closely to the Sinona II Pack terri tory which existed from 1977
through mid-winter 1979 (Ballard et al.1981).During winter
1979 most members of the Sinona II Pack were ground shot with-
possibly one or two wolves surviving.However,one of the re-
maining wolves was radio-collared and it began making long dis-
tancemovements from its normal area,a characteristic of a wolf
in the process of dispersing.Therefore,we suspected the area
to be vacant,except perhaps for one lone wolf.We concluded
that the temporary establishment of this pack represented the
colonization of a vacant terri tory by a group of dispersing
wolves.The status of this pack as of mid-October 1981 was un-
known but a trapper reported observing a single wolf in the area
which could be the third unaccounted for pack member.
Tolsona Pack
Prior to mid-June 1978 contact with this pack consisted of public
sightings,track counts and harvest records.These data indi-
25
cated that in early fall 1977 the pack had numbered at least 11.
By the end of winter,trappers had reduced the pack to three in-
dividuals.
Radio contact with this pack was established in early June 1978.
In search of a potential den site,aspen-covered knolls were
examined from fixed-wing aircraft unti 1 the den site was found
and a yearling gray male (*210)was radio-collared.At that time
the pack was compri sed of wolf 210,an adult gray female I and a
small black wolf which may have been a yearling.At least eight
pups were rai sed at the site.Pups were moved to a rendezvous
site 3.5 miles (5.6 kIn)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 forages wolf 210 was always observed
alone.In mid-September 1978 radio contact was lost.At that
time the pack numbered 10.
There was no radio contact with this pack from September 1978
until late January 1979 at which time a live black yearling pup
was purchased from a local trapper.The pup had been caught by
the toes and was in good enough condition to be radio-collared.
At the time her pack affiliation was uncertain.Within 2 weeks
of capture,however,she had rejoined the Tolsona pack which then
numbered seven (three blacks and four 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.
An accurate count of the number of pups produced was never ob-
tained.In mid-October,however,the pack numbered 16 (11 grays
and 5 blacks).On the basis of size and the scruffy appearance
usually exhibited by pups at that time of the year,it is be-
lieved at least six and perhaps nine pups were raised (3 blacks
and 6 grays).
26
~-
~.
-
-
During summer 1979 members of this pack appeared to continuously
---~--- --~
expand their range westerly into the old Mende1tna territory.In
mid-October,when the largest count of the pack (16)was ob-
tained,they were located close to Moore Lake which had been the
northern territory boundary of the Mende1tna pack.Although wolf
210 had not been radio-located since late August of 1978,it was
present wi th the pack at that time.
During winter 1979-80 the pack suffered attrition due to trapping
and perhaps from dispersal even though portions of their terri-
tory were included in an area closed to hunting and trapping.
Radio contact was temporarily lost when wolf 220 I S radio trans-
mi tter failed prematurely.Contact with this pack was not re-
established 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 compri sed of a minimum of two black adults I
seven gray adults and six pups.In late June the pack moved to
the Nicholson Lake rendezvous site which had been used as a den
site by the Mendeltna pack in 1977.This movement was the result
of an attempt to ground capture and radio collar one wolf.On
6 July wolf 220 was recaptured by helicopter darting.
By late fall 1980,the pack numbered 16 (4 blacks and 12 grays)
suggesting that seven pups may have been raised.
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's territory into the Mendeltna area,it is suspected the
black wolf may have been a survivor of the Mendel tna pack
(Ballard et ale 1981).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,fol-
lowing 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.
27
Through early winter 1980-81 our counts suggested that 16 or 17
-_.--~-.-_.----
wolves were present in the Tolsona Pack.Number and coloration
varied through winter ranging from 4 blacks and 12 gray wolves to
5 blacks and 10 gray wolves.In mid-October 1980 two additional
wolves were captured and radio-collared;wolf 315 ,a black adult
female believed to be the alpha female in 1980 and wolf 316 ,a
yearling male.
During winter 1980-81 the Tolsona pack frequented the west and
northern portions of its terri tory more frequently than it had
during the winters of 1978-79 and 1979-80.Reasons for the shift
in di stribution appear to be related to the winter di stribution
of caribou during those years.During winters 1978-79 and 1979-
80,large numbers of caribou wintered on the Lake Loui se flats
east of Old Man Lake.However,during winter 1980-81 caribou
appeared to be more abundant west and north of Old Man Lake.
Wolf radio-location data suggest that the Tolsona wolves rarely
utilized the southeastern portion of their territory during this
relative absence of caribou.
During early and mid-winter 1980-81 no mortalities due to either
hunting or trapping were observed primarily because of poor snow
and weather conditions,which were not favorable for tracking
wolves from fixed-wing aircraft.By March 1981,however,our
counts of this pack varied from 10 to 13 wolves,suggesting that
some had either dispersed or had been trapped.Radio contact
with wolf 316 was lost during this time period.Since wolf 316
was a yearling in October 1980 and because yearlings frequently
disperse at this time of year (Ballard et al.1981),we suspect
di spersal accounted for most of the losses.
On 14 April 1981 two additional members of the Tolsona Pack were
captured and radio-collared;wolf 210 an adult male whi.ch origi-
nally had been captured as a yearling in June 1978 and was last
observed in September 1978,.and wolf 328 a young adult female.
Wolf 328 apparently dispersed immediately after capture because
radio contact was not established until 2 June 1981,when she was
28
1lIQ1:
-
'"""'
-.
.....
,~
.....
-
observed alone on an old moose kill several miles north of the
--","-...-.- -------_.-
Susi tna River gauging station;a straight line movement of ap-
proximately 44 miles from her capture location.She remained in
that area for 3-4 days,killing at least one calf moose.Radio
contact was again terminated until 4 August 1981 when she was
observed on the East Fork of the Susitna River where contact was
again lost.We suspect that she may have continued traveling
northward into GMU-20 north of the Alaska Range.The straight
line distance from her capture location to the last observation
in August was approximately 72 miles.We suspected that several
other Tolsona Pack members also dispersed because by 5 June the
largest number of adult wolves counted was 8 (5 grays and 3
blacks).
Tolsona Pack members were first observed at the Nickolson Lake
den site,also utilized in 1980,on 14 May 1981.On 4 June at
approximately 1400 hrs.,the den site was approached by a photo-
grapher and the site was immediately abandoned by the wolves.
Between 1400.hrs.,on 4 June and approximately 1800 on 5 June the
pack had moved the pups from the den site to the Nickolson Lake
rendezvous site,approximately 1 mile to the west.At that time
3 pups (2 blacks and 1 gray)were observed.Five pups (3 blacks
and 2 grays)were observed on 6 June while the largest number,6
(3 blacks - 3 grays),was observed on 11 June.Pups were first
observed with adults away from the rendezvous site on 20 August
1981 at an adult caribou kill.
During mid-July 1981 wolf 313 from the Susitna-Sinona Pack was
detected with the Tolsona Pack.Wolf 313 apparently was in the
process of dispersing from the Sinona Pack because in late March
1981 he was observed alone in the Wrangell Mountains (see
Susitna-Sinona Pack history).Thus between 31 March and at least
23 July 1981 the wolf had moved a minimum straight line distance
of 40 miles to join the Tolsona Pack.He remained with the pack
through summer and fall 1981 when his radio prematurely failed.
Counts of the pack in mid-October suggest that the wolf is still
present and therefore it appears the wolf has been successfully
accepted into the pack.
29
Apparently there was no pu~~o_r-t:a~_i~¥~~n 1981 because by late
fall the pack numbered between 14-15 wolves (8 original adults,6
pups and wolf 313).
From June 1978 through October 1981 the Tolsona pack occupied an
area of 981 mi 2 (2,541 km 2 ).Their range extended from Tazlina
Lake to Lake Loui se,west to Tyone Creek and then east several
miles past Tolsona Creek.At this time no other pack territories
were believed to overlap the Tolsona terri tory.
Tyone Creek Pack
Prior to establishment of radio contact with this pack in
November 1977,data consisted of track counts and public sight-
ings.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 re-
trieved.The remaining wolf (#116)dispersed from the area.By
27 February he was observed accompanied by a black 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 during ex-
perimental wolf control studies.Wolf 116 continued to reside in
the Susitna study area and by 20 June was observed with a year-
ling 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 kIn)from his original capture location.Fol-
lowing this latter observation,radio contact with wolf 116 was
lost and it is assumed he dispersed farther to the north or west.
During fall 1977 and early winter 1978 the TyoneCreek pack oc-
cupied an area of 253 mi 2 (655 km 2 ).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.
30
-
-
....
-
.-
-
In November 1978 contact with wolves in this area was re-estab-
lished when two adults (#215 and 216)were radio-collared.
Whether these wolves were descendants of the original Tyone Creek
pack,which was thought to have been eliminated by ground
shooting in 1978,or represented wolves colonizing a new area is
not known.They did not,however,occupy the area previously
occupi ed by the Tyone Creek pack.
During winter 1978-79 no other wolves were ever observed with
this pair.On 23 April they were first observed at the 1979 den
si te.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.
Wolf hunting and trapping within this pack's territory was closed
in both 1978-79 and 1979-80 so that wolf numbers would remain
stable for the predation rate studies.However,five wolves,
three of which were radio-collared (#215,216 and 301),were re-
moved from the pack.One of these was legally ground shot when
the pack ventured out of the closed area in January 1980.In
March 1980,four others were killed illegally by aerial hunters.
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 were apparently unable to kill either moose or
caribou.From mid-March through December 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 the pack
began 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 Susi tna River ap-
31
proximately 50 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.
By late June 1980 the pack of four was joined by an uncollared
black wolf.By mid-July the three radio-collared wolves began
exhibi ting a propensity to travel independently of one another.
Radio contact with wolf 299 was "lost after 24 October 1980 when
it was located at Boulder Creek above the Denali Highway.On 17
January 1981 wolf 299 was killed by a trapper 10 miles southwest
of Fairbanks.
Wolf 298 continued to frequent the old Tyone Creek territory ex-
cept on 27 August she was observed approximately 40 miles (64 km)
to the south at the head of Chitina Creek accompanied by one gray
and one black.Following this observation wolf 298 was always
observed with one black wolf in the old territory.This black
was captured and radio-collared (#317)on 16 October and was de-
termined to be a young adult male,suggesting that a bond had
been formed.In late December 1980,the carcasses of three
wolves were found on a lake bed in the vicinity of the area oc-
cupied by wolves 298,317 and the uncollared gray.We were un-
able to find the radio-collared wolves following this observation
and,therefore,we assumed they had been killed.Neither the
kills or the radio-collars were reported to the Department.
Between 26 August and 18 September 1980 wolf 300 dispersed ap-
proximately 65 miles (104 km)northeast to the vicinity of Dickey
Lake.Thi s young male remained"in the vicinity of Dickey Lake
through early fall and by 23 October was accompanied by an uncol-
lared black wolf.We originally suspected this uncollared wolf
was a female and that a pair bond had been formed because the
wolves were occupying an area which was believed to be vacant
(Ballard et al.1981).Between 24 November 1980 and 19 March
1981 we lost radio contact with wolf 300.On 31 March 1981,he
32
'"""
-
--
~,
-
-
was located approximately 3 miles east of Crosswind Lake and was
"---.-------------
accompanied by 3 black and 5 gray wolves having moved a straight
line distance of 38 miles to the south.Subsequent observations
suggested wolf 300 had joined an existing pack located near Fish
Lake.Additional history of wolf 300 from 31 March through mid-
October 1981 is provided in the Fish Lake Pack section of this
report.
As mentioned earlier following the removal of the adult pair·in
March 1980,the remaining four pups were never observed on a
fresh ungulate kill.They did revisit old kills which had been
made prior to march 1980.Even following the separation of the
four yearlings in late summer,none were subsequently observed on
a fresh ungulate kill until 31 March 1981 after wolf 300 had
joined the Fish Lake Pack.These observations suggest that young
wolves are unable to successfully hunt and kill ungulates without
the experience and leadership provided by adults.Young wolves
(10 months)were able to survive without adults,however,they
subsisted on old kills and small game.
Prior to the removal of the adul t wolves from the Tyone Creek
pack in March 1980,the pack had occupied a relatively small ter-
ritory of 364 mi 2 (943 km 2 )from November 1978 through March
1980.Following the apparent deaths of wolves 298 and 317 and
the dispersal of wolves 299 and 300,the Tyone Creek Pack Area
was believed to be.vacant.This speculation was at least par-
tially confirmed by the dispersal of Susitna wolves 330 and 331
during summer 1981.
Fish Lake Pack
Radio contact with the Fish Lake Pack was established in mid-
March 1981 when wolf 300,originally from the Tyone Pack,as de-
scribed in the previous section,dispersed into the area.With
the addition of wolf 300 and we assume the uncollared black,the
Fish Lake Pack numbered 9 wolves (3 blacks and 6 grays).Wolf
300 was fitted with a new transmitter in mid-April 1981 at which
33
time three additional pack members were radio-collared (#326-an
adul t gray female,#327-a pup gray male,and #329-a pup black
female).
On 23 May 1981 wolves 300 and 326 were observed near Fish Lake at
what from the air appeared to be a den site.No pups were obser-
ved at the area and ground examination in September suggested
that although the area had been heavily frequented by wolves,it
was not a den si te as suspected.
Evidence that the Fish Lake pack successfully produced pups in
1981 was not obtained until 23 September when 2 gray and 1 black
pup were observed with 4 adults.Throughout fall 1981 the ra-
dioed adult members of the pack were rarely observed together.
Wolves 300,326 and 327 all exhibited a tendency to periodically
travel independently from the main body of the pack.
From March through fall 1981 the Fish Lake Pack occupied a rather
small area of 346 mi 2 which must be considered representative
only of the pack's summer home range.Boundaries of the area are
depicted in Fig.4.
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.Contact with this wolf was lost in
Apri 1 for unknown reasons.
From April 1978 to April 1980 data for this pack consisted only
of track counts and observations by Department personnel.By
fall 1978 the pack numbered three and may have remained at that
level through spring 1979,although the presence of only two
wolves could be ascertained.The pack apparently denned in 1979
because seven wolves were present by fall.
34
-
....
..."
-
.....
-
-
-
-
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.Prior to parturition the
pack was known to be comprised of at least five and perhaps as
many as eight wolves.Between 8 and 14 July pups were moved to a
rendezvous site approximately 1.0 mile WNW of the den site.No
other rendezvous sites were observed and the pups began traveling
wi th the adults regularly by late September.During September
wolf 311 was shot by a hunter on Watana Creek.·Radio contact
with wolf 309 was lost after 12 August either due to dispersal or
radio failure.By late December,13 wolves still remained in the
pack suggesting that pup survival was high and no further adult
mortali ty had occurred.From April through December the Watana
pack occupied an area of 400 mi 2 (1036 km 2 ) •
On 7 March 1981 an additional three wolves were captured and ra-
dio-collared (#323-pup gray female,#324-pup gray female,and
#325-pup gray male).
Wolf 310 was observed dispersing from the Watana Pack on 6 March
1981 when he was observed alone on the Susi tna River above the
big bend,18 miles from the eastern boundary of the Watana Pack.
He returned to the pack at Fog Creek by 24 March and remained
wi th them at least through 17 April.
On 18 April wolf 308,which was pregnant,was recaptured and fit-
ted with an acti vi ty transmitter for den site studies to be con-
ducted in early summer.Wolf 308 was first observed at the den
si te (same site used in 1980)on 6 May 1981.On 10 May both
wolves 324 and 325 were recaptured and also fitted with activi ty
transmitters.However,only three (#'s 308,324 and 325)wolves
from the Watana Pack could be located near the den area.
Late on 10 May 1981 wolves 310 and 323 apparently wi th several
r'"
uncollared members of the Watana Pack were observed with wolf 331
and perhaps other uncollared gray wolves from the Susi tna Pack
35
(see Susi tna Pack history this report).The Watana wolves were
approximately 26 miles east of the Watana Pack's eastern bound-
ary.At that time,four wolves,including 310 which was recap-
tured,were fitted with activity transmitters for the Watana den
study (#344-yearling gray male,#345-yearling gray male,and
#346-yearling gray female).
Wolf 308,the alpha female,was first observed at the Watana den
site,(same site used in 1980)on 6 May 1981.A blind for ground
observation was established at the site on 7 May.The daily ac-
tivi ties an,d food habits of the pack from 8 May through 1 July
will be discussed in food habits and activity patterns section of
this report.No attempt was made to observe pups from fixed-wing
aircraft in an effort to minimize disturbance.Consequently pups
were not observed from fixed-wing aircraft until 1 July 1981 when
3 pups were observed at a rendezvous site,located 0.8 miles
north-northwest of the den site.The move occurred between 21
June and 1 July.Six pups were rai sed during 1981.Pups re-
mained at the rendezvous si teat least through 24 August 1981.
By fall 1981 the pack numbered 15.
Elevation and Seasonal Usage of Habitat Types
Habitat use was examined by plotting radio location data for the
Watana Pack on 1:63,000 scale vegetation maps provided by the
Agricultural Experiment Station in Palmer,Alaska.Data points
which did not specifically fall within one habitat were tallied
as in an ecotone between the two types.Results of this analysis
are provided in Table 4.According to this analysis 8 of 24 hab-
itat types occurring in the area were not used,either singly or
in combination as an ecotone by the Watana wolf pack during the
study period (4/80-10/81).The unused types included snow and
ice,wet sedge grass,closed balsam poplar,open balsam poplar,
willow shrub,grassland,disturbed,and lakes.Because most of
the sampling period occurred during warm months the non-use of
lakes is misleading.Wolf packs frequently make kills on or at
the edge of lakes and streams during winter.
36
1 J J J J J J 1 1 )J .~
B
Table 4.Habitat type use by month of occurrence for the Watana wolf pack in Southcentral Alaska from
April 1980 through mid-October 1981.
Habitat Type Months Total %
(Mono type)Jan.Feb.Mar.Apr.May June July Aug.Sept.Oct.Nov.Dec.
low shrub 3 3 3 9 10.5
Woodland black spruce 1 6 4 3 1 1 16 18.6
Closed tall shrub 2 1 3 3.5
Open tall shrub 2 2 2.3
Open black spruce 1 3 1 1 6 7.0
Birch shrub 4 1 3 1 9 10.5
Open mixed forest 1 1 2 2.3
w
-...,J
Closed birch forest 1 1 1.2
Closed mixed forest 1 1 1.2
Mat-cushion tundra 1 1 1.2
Subtotal 0 0 1 3 20 10 3 3 2 8 0 50 583
Ecotones Jan.Feb.Mar.Apr.May June July Aug.Sept.Oct.Nov.Dec.Total
woodland bl.spruce-open
bI.spruce 3 3 6 7.0
open w.spruce -
open black spruce 1 1 1.2
closed m.for.-woodland
bI.spruce 1 1 1.2
sedge-grass t -open tall shrub 1 1 1.2
woodland bl.sp.-low shrub 3 1 1 5 5.8
Table 4.(cont'd)
Ecotones Jan.Feb.Mar.Apr.May June July Aug.Sept.Oct.Nov.Dec.Total
open white sp.-
open bl.sp.I I 1.2
closed mix f.-
woodland bl.spruce I I 1.2
birch shrub -
open bi.spruce I I 1.2
birch shrub -low shrub 3 2 5 5.8
low shrub -open t.shrub I I I 3 3.5
woodland bi.sp.-
sedge sh.tundra I I 1.2
w open birch forest -00
open white spruce I I 1.2
sedge grass to -
mat-cushion tundra I I 1.2
birch shrub -sedge
grass tundra I I 1.2
closed birch f -
woodland white sp.I I 1.2
birch shrub -
,open tall shrub I I 1.2
sedge grass t.-
low shrub I I 1.2
closed birch f.-
low shrub I I 1.2
J J J .J I J ~J i J }J t J l ]j J
I )J }}J l 1 ]CJ --1 •...]
W
\0
Tabl.e 4.(cont'd)
Ecotones Jan.Feb.Mar.Apr.May June July Aug.Sept.Oct.Nov.Dec.Total
open tall sh.-
woodland b 1.sp.1 1 1.2
closed mixed for.-
open mixed for.1 1 1.2
rock -sedge grass tundra 1 1 '1.2
Subtotal 0 2 2 2 15 7 2 0 2 4 0 36 42.5
Total 0 2 3 5 35 17 5 3 4 12 0 86 100.8
Of the 18 habitat or ecotone types used by the Watana wolf pack,
10 were monotypes.Wolves were located in the monotypes on 58%
of the occasions they were located,assuming the radio-location
data and the boundaries of the habitat types were accurate.Of
the 10 monotypes,low shrub,woodland black spruce,closed tall
shrub,open black spruce,and birch shrub accounted for 86 per-
cent of the use.Thirty-six (43%)of 86 locations occurred in
ecotone areas.As the vegetation maps do not show many ecotones
present,ecotone use might have been even higher.Of the 21
classified ecotone areas 17 (81%)had only one use occurrence.
However,20 (56%)of 36 used involved one of the shrub habitat
types.Overall,shrub habitat types accounted for 50 percent of
all use.
Elevation and slope for each radio location of Watana wolf pack
members is provided in Table 5.The average monthly elevation
occupied by pack members ranged from,2,207 feet in April to
3,350 feet in November.Of the 96 locations,34 (35%)were
located wi thin the area to be inundated by the Watana impound-
ment.Sixty-three percent of all locations occurred at or below
the 2,500 foot level.Sample sizes were too low to compare sea-
sonal changes in elevationaluse.
Slopes at each radio-location were classified as follows:flat
(F)- 0
0 to 10 0 ,gentle (G)-110 to 30°,moderate (M)-31 0 and
above,and riverine (R),occurring along stream or river.Based
upon the latter classification,the 96 locations for the Watana
Pack were as follows based on increasing order of occurrences:
F -15 (16%),R -19 (20%),M -35 (37%),and G -27 (28%).On a
year-round basis there did not appear to any selection for either
slope (x z =7.54,P>O.l)or aspect (x z =12.2,P>O.l)by the Watana
wolf pack.
Food Habits
During 1980 and 1981 six radio-collared wolf packs were observed
on 83 kills.Moose comprised 57 percent of the kills,while
40
~,
-
-
"...
'I )1 )1 })J -1 )-J J 1
Table 5.Elevation and aspect usage by members of the Watana Wolf Pack from April 1975 to mid-October 1981
in the Sus{tna River Basin of south-central Alaska.
Elevation (N)II o f o c cur r e n c e s
Month x Range Exposure F N S E W NE SE SW NW impoundment
Jan 2475 2150-2800 2 1 1
Feb 2575 2550-2600 2 1 1 0
March 2850 2150-3300 3 1 1 1 1
Apr 2207 1750-2600 7 2 2 3 4
May 2295 1600-3900 38 8 5 7
2 2 2 4 5 3 18
Jun~2484 1600-3300 22 1 6 3 1 3 4 4 8
+'July 2516 2450-2600 3 1 1 1 0
I-'
Aug 2367 2000-2800 3 1 1 1 1
Sept 2983 2500-3400 6 1 2 1 1 1 0
Oct 2614 1900-3500 7 2 1 2 1 1 1 1
Nov 3350 2700-4000 2 1 1 '0
Dec 3200 1 1 0
Totals 96 10 16 16 8 6 5 14 12 8 34
caribou comprised 33 percent.Other prey such as snowshoe hare,
._-"---------------
beaver,muskrat,and other small mammals comprised the remaining
11 percent.Moose calves comprised 51 percent of the moose kills
while for caribou,calves comprised only 7 percent of the kills.
Detailed descriptions of individual pack food habi ts follow.
Fish Lake Pack From 19 March through 26 October 1981 members
of the Fish Lake Pack were observed on five kills and on two
other occasions the wolves were in the process of making a kill
when observed (Table 6).The confirmed kills were as follows:2
caribou,1 adult moose,and 2 small mammals.Because of the lo-
cation of this pack on the Lake Louise flats,we would suspect
that caribou would comprise a large portion of this pack's diet
during 1980 and 1981.Al though we were unable to locate a den
site,we were able to locate a rest area and collected 19 scats
which contained mostly caribou (40%)remains (Table 7).The im-
portance of caribou prey to this pack undoubtedly fluctuates de-
pending on their availability.
Susitna Pack During 1980 and 1981 this pack was observed on
23 kills which were comprised of 10 caribou (43.5%),11 moose
(47.8%)and 2 small mammals (8.7%)(Table 8).In 1979,however,
only 9 percent of the kills (n=22)were comprised of caribou.
During 1979 caribou were essentially unavailable to this pack
while in 1980 and 1981 fair numbers of caribou wintered wi thin
this pack's territory.
Scats were collected at the Susi tna den and rendezvous site in
1980 and at the den site in 1981 (Tables 9,10 and 11).Moose
was the most important single summer food item in both 1980 and
1981,comprising 33.3 and 28.6 percent,respectively,of the
diet.However,as a group small mammals were equally important,
35%in 1980 and 41%in 1981,with beaver constituting the most
important small mammal food item.Moose were also the most im-
portant food item at the 1980 rendezvous site.There did not
appear to be any maj or changes in summer food habits from 1980
and 1981.
42
-
-
......
-
-Table 6.Chronological summary of kills at which the Fish Lake wolf pack was
observed from 19 March 1981 through 26 October 1981 in GMU-13 of
southcentral Alaska •...
Date of Kill Species and Age Percent Estimated
Observation Time Made Consumed Date of Kill
or Comments
3/31/81 9:25am Yes Moose-adult 90%9 wolves
4/20/81 4:00pm Yes Beaver or muskrat 1 wolf
5/22/81 11 :45am ?Injured bull moose No sign of kill
at bay on 5/23
5/23/81 pm Probable Injured bull caribou
at bay 1 wolf
9/23/81 10:22am Yes Caribou-calf 20%9/23 1 wolf
9/23/81 10:02am Yes Caribou-adult 10%9/23 2 wolves
10/26/81 2:39pm Yes Small mammal F 10/26 7 wolves
43
~I
~.
Table 7.Summary of incidence of food remains in wolf scats collected at a Fish
Lake pack rest area during summer 1981 in south central Alaska.-
Item in 19 Items in 0 Items in 19
Food Item Adult Scats Pup Scats Combined Scats """II %{I %II %
Moose,adult 0 0 0 0 0
Moose,calf 3 13.04 0 0 3 13.04 -Caribou,adult 4 17.39 0 0 4 17.39
Caribou,calf 1 43.5 0 0 1 4.35
Caribou,age unknown 4 17.39 0 0 4 17.39
Moose or caribou 1 43.5 0 0 1 4.35
Microtine 3 13.04 0 0 3 13.04
Bird 2 8.70 0 0 2 8.70
Wolf 1 4.35 0 0 1 4.35 -Unknown 4 17.39 0 0 4 17.39
Total 23 100.00 0 0 23 100.00
~
Grouped Data for 19 Scats
Food Item {I %
Ungulate 13 56.52
Small Mammal 3 13.04
Other 3 13.04
Unknown 4 17.39
Total 23 100.00
44
-
Table 8.Chronological summary of kills at which the Susitna wolf pack
was observed from January 1980 through December 1981 in Game
Management Unit 13 of southcentral Alaska l/.
Date of
Observation Time
Kill
Made Species and Age
Percent Estimated Date
Consumed of Kill or Comments
9:35am No
12:20pm Yes Caribou -adult M 90%
12:30pm No 95%
11:30am Yes Moose -calf
1:00pm Yes Moose -adult
10:45am No
10:45am Yes Caribou -adult
3:40pm Yes Moose-assumed calf
1 wolf
3 wolves
7 wolves
4 wolves
7 wolves
7 wolves
1 wolf
4 wolves
1 wolf 2/
2 wolves
8 wolves
7 wolves
3 wolves
Had cow w/nb
calf at bay.
2 wolves
1 wolf
Also had cow
w/nb calf at bay
1 wolf
4 wolves
4 wolves
7 wolves
7 wolves
2/11 or
2/12
4 wolves
3/13 4 wolves
Still on kill
of 3/14 4 wolves
1/22 or
1/23
1/26
1/28
2/2
3/18
3/20 or
3/21
3/23 or
3/24
4/13-14
5/17-18
5/25
6/2
5/22-23
old
8/26
11/18
5/23
75%
90%
90%
50%
75%
0%
90%
100%
100%
100%
100%
100%
50%
25%
old kill
5%
0%
50-75%
100%
0%
95%
Caribou -adult
Caribou -assumed
adult
Caribou -adult M
Moose -adult
Moose -nb calf
Caribou -calf
Unknown
Caribou -adult
Caribou -?
Moose -adult or
L.yearling
Caribou -adult
Beaver
Moose -adult?
Moose -?
Moose -calf
Caribou -?
Moose -calf
Moose -nb calf
11:08am No
12:23pm Yes
9,:23am No
11:05am Yes
2:32pm No
12:03pm Yes
10:30am No
2:48pm No
11 :50pm No
2:00pm Yes
4:46pm Yes
8:45am No
12:30pm Yes
7:45 pm Yes
1:00pm Yes
11:30am Yes
7:lOpm Yes
12:15pm -
9:30am Yes
12:40pm Yes
12:35pm Yes
Yes
6:46pm Yes
12:00pm Yes
6:05pm ?
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
4/15/80
5/18/80
5/25/80
6/2/80
6/23/80
8/7 /80
8/26/80
11/19/80
5/11/81
5/23/81
5/28/81
5/29/81
6/5/81
6/9/81
-.
.-
I
1/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.
~/Possible bear kill.
45
Table 9.Summary of incidence of food items in wolf scats collected at the Susitna
wolf den occupied in late spring and summer 1980 in GMU-13 of southcentral
Alaska.
Item in 45 Items in 34 Items in 79
Food Item Adult Scats Pup Scats Combined Scats
If %If %IF %
Moose,adult 3 5.36 2 5.00 5 5.21
Moose,calf 8 14.29 17 42.50 25 26.04
Moose,age unknown 2 3.57 0 0 2 2.08
Caribou,adult 5 8.93 2 5.00 7 7.29
Caribou,calf 2 3.57 1 2.50 3 3.13
Caribou,age unknown 1 1.79 0 0 1 1.04
Moose or caribou '7 12.50 3 7.50 10 10.42
Beaver 11 19.64 3 7.50 14 14.58
Muskrat 6 10.71 5 12.50 11 11.46
Snowshoe hare 4 7.14 1 2.50 5 5.21
Microtine 1 1.79 3 7.50 4 4.17
Vegetation 1 1.79 1 2.50 2 2.08
Unknown 5 8.93 2 5.00 7 7.29
Totals 56 100.01 40 100.00 96 100.00
Grouped Data for 79 Scats
Food Item If %
Ungulate 53 55.21
Small MaIIJIIlal 34 35.42
Other 2 2.08
Unknown 7 7.29
Total %100.00
46
-
-
-
-
~,
.......
..-
Table 10.Summary of incidence of food remains in wolf scats collected at the
Susitna wolf pack rendezvous site occupied during summer 1980 in
GMU-13 southcentral Alaska.
Items in 0 Items in 13 Items in 13
Food Item Adult Scats Pup Scats Combined Scats
1/%1/%1/%
Moose,adult 0 0 1 6.67 1 6.67
Moose,calf 0 0 10 66.67 10 66.67
Moose,age unknown 0 0 1 6.67 1 6.67
Bird 0 \0 1 6.67 1 6.67
Vegetation 0 0 1 6.67 1 6.67
Unknown 0 0 1 6.671 1 6.67
Totals 0 0 15 100.00 15 100.00
Grouped Data for 13 Scats
Food Item if %
Ungulate 12 80.00
Small Manunal 0 0
Other 2 13.33
Unknown 1 6.67
Total 15 100.00
47
Table 11.Incidence of food remains in wolf scats collected at the Susitna den
utilized in late spring and early summer 1981 in GMU-13 of southcentral
Alaska.
Items in 41 Items in a Items in 41
Food Item Adult Scats Pup Scats Combined Scats
il %/I %/1 %
Moose,adult 1 1.59 -0-a 1 1.59
Moose,calf 17 26.98 0 0 17 26.98
Caribou,adult 6 9.52 0 0 .6 9.52
Caribou,calf 3 4.76 a a 3 4.76
Beaver 10 15.87 0 a 10 15.87
Muskrat 2 3.17 0 a 2 3.17
Snowshoe hare 2 3.17 0 a 2 3.17
Hicrotine 10 15.87 0 a 10 15.87
Unidentified small mammal 2 3.17 a 0 2 3.17
Bird 3 4.76 0 a 3 4.76
Fish a a a a a 0
Vegetation 2 3.17 a a 2 3.17
Unknown 5 7.94 a a 5 7.94
Totals 63 100.00 0 a 63 99.9
Grouped Data for 41 Scats
Food Item /I %
Ungulate 27 42.86
Small Mannnal 26 41.27
Other 5 7.94
Unknown 5 7.94
Total 63 100.00
48
-
-.
~I
..."
-
"""'
Susitna-Sinona Pack -During 1980 and 1981 members of this pack
were only observed on 5 kills (Table 12)because they were only
periodically monitored.The observed kills were comprised of 2
caribou,2 moose and 1 snowshoe hare.Prior to the establishment
of this pack in spring 1979 the area had been occupied by another
pack referred to as the original Sinona Pack.Interestingly,the
original Sinona Pack was observed on only one (2.6%)caribou out
of 39 kills (Ballard et a1.1981).Again,the importance of
caribou as a prey item is probably dependent on both ~aribou dis-
tribution and population status.Although caribou migrated
through this area prior to 1980,very few overwintered in the
area as they did in 1980 and 1981.Although this pack denned in
1980,we were unable to locate a den site and therefore no wolf
scats were collected for summer food habi ts studies.
Tolsona Pack -From 6 January 1980 through October 1981 members
of the Tolsona Pack were observed on 19 kills (Table 13).Moose
accounted for 9 kills (47%)and caribou for 7 kills (37%),while
bear and snowshoe hare comprised 3 kills.In 1979,however,
caribou comprised 8 (67%)of 12 ki lIs.The difference in the
importance of caribou to the Tolsona Pack again appears to be
related to caribou distribution.
Summer food habits were determined by examination of scats col-
lected at den and rendezvous sites in 1980 and 1981 (Tables 14,
15 and 16).In 1980 (Table 14)ungulate prey comprised 79 per-
cent of the prey remains with calf moose comprising almost half
(48%)of the identified prey items,while caribou of all ages
comprised 8 percent of the items.In 1981,ungulate prey were
also the most important prey item (54%of total combining den and
rendezvous scats),however,caribou comprised 36 percent of the
food items.Small mammals were an important prey item in both
years;33 percent in 1980 and 42 percent in 1981.
Tyone Creek Pack During 1980 thi s pack was observed on 11
kills (Table 17),10 (91%)of which were moose.Following the
loss of the adult male and female in early spring 1980 remaining
49
Table 12.Chronological summary of kills at which the Susitna-Sinona wolf pack was
observed from mid-April 1980 through August 1981.1/
Date of Species and Age Percent Estimated Date of
Observation Time Consumed Kill or Comments
1/9/80 11:40am Caribou -?75%1/8 2 wolves obs.
1/29/80 10:30am Moose -calf 75%1/28 3 wolves
9/26/80 9:05am Moose -adult F?100%?3 wolves
10/14/80 9:15am Probe snowshoe hare 10/14 4-5 wolves
3/16/81 4:30pm Caribou - ?100%1 wolf
1/Wolf 313 joined Tolsona Pack by mid-July 1981.
50
~I
Table 13.Chronological summary of kills at which the Tolsona wolf
pack was observed from 6 January through 17 February 1980
and July 1980 through 8 September 1981 in Game Management
Unit 13 of southcentral Alaska.
Date of
Observation Time Species and Age
Percent Estimated Date of
Consumed Kill or Comments
15 wolves
15 wolves
7+wolves
15 wolves
5/23/81 1 wolf
Kill appeared to have
been buried and dug
up -possible bear kill?
1 wolf present
1 wolf
same wolf present at kill
12 wolves
1/5 4 wolves
1/8 11 wolves
1/24 12 wolves
1/26-27 12 wolves
1/28-29 12 Wolves
2/7 12 wolves
2/8 12 wolves
Still on kill
of 2/9 11 wolves
2/14 8 wolves
2/15-16 11 wolves
Still on kill
of 2/16 11 wolves
8/26-27 4 wolves
10/16 15 wolves
15%
75%
80%
95%
90%
90%
75%
95%
100%
calf moose
100%
95%
75%1/20/81
Leaving kill
95%
<5%
80-90%
Caribou -?
Moose-newborn calf
Moose -?
Caribou -Adult
Caribou -?
Moose -Calf
Caribou -?
Moose -Calf
River Otter
Moose - 2 yr.
am
7:00pm
7:18pm Moose -newborn calf 50%
6:35pm
1:45pm Caribou -adult
9:35pm Unk.-poss.beaver
1:20pm Moose -Calf
12:05am
3:30pm Caribou -Ad M
11:00am Snowshoe hare
10:00pmm Attempting to kill
11:20pm Moose -adult
10:02am Caribou -?
10:15am Moose -calf
10:00am
10:25am
2:45pm
10:45am
10:06am
10:54am
11:15am
3:36pm
11 :05pm
10:20am
1/6/80
1/9/80
1/25/80
1/27/80
1/29/80
2/7 /80
2/9/80
2/10/80
2/14/80
2/16/80
2/17/80
8/27/80
10/16/80
1/09/81
1/13/81
1/16/81
-1/21/81
1/22/81
1/24/81
5/23/81
6/2/81
6/4/81
6/5/81
8/20/81
-.
51
Table 14.Incidence of food remains in wolf scats collected at the Tolsona wolf
rendezvous site (old Mendelta 2nd den)utilized by pack members during
summer 1980 in GMU-13 of southcentral Alaska.
Item in 113 Items in 104 Items in 217
Food Item Adult Scats Pup Scats Combined Scats
fl %IF %#%
Moose,adult 32 19.63 1 0.85 33 11.74
Moose,calf 62 38.04 74 62.71 136 48.40
l'loose,age unknown 4 2.45 1 0.85 5 1.78
Caribou,adult 15 9.20 3 2.54 18 6.41
Caribou,calf 5 3.07 4 3.39 9 3.20
Caribpu,age unknown 3 1.84 2 1.69 5 1.78
Moose or caribou 3 1.84 9 7.63 12 4.27
Beaver 5 3.07 0 0 5 1.78
Muskrat 4 2.45 0 0 4 1.42
Snowshoe hare 4 2.45 6 5.08 10 3.56
Microtine 7 4.29 4 3.39 11 3.91
Unidentified small mammal 7 4.29 0 0 7 2.49
Bird 5 3.07 1 0.85 6 2.14
Fish 1 0.61 0 0 1 0.36
Vegetation 0 2 1.69 2 0.72
Unknown 6 3.68 11 9.32 17 6.05
Totals 163 100.00 118 100.00 281 100.11
Grouped Data for 217 Scats
Food Item
Ungulate
Small Mammal
Other
Unknown
Total
fl
218
37
9
17
281
%
77 .58
13.17
3.20
6.05
100.00
52
~,
-
Table 15.Incidence of food remains in wolf scats collected at the Tolsona wolf den
~site (old Mendelta rendezvous site)utilized by pack members during late
spring and early summer 1981 in GMU-13 of southcentral Alaska.
.-Item in 59 Items in 5 Items in 64
Food Item Adult Scats Pup Scats Combined Scats
it %If %1F %
Moose.adult 4 4.82 0 0
-4-4.44
PDt Moose.calf 18 21.69 2 28.57 20 22.22
Moose.age unknown e 0 0 0 0
Caribou.adult 9 10.84 2 28.57 11 12.22-Caribou.calf 12 14.46 0 0 12 13.33
Caribou.age unknown 0 0 0 0 0 0
Moose or caribou 7 8.43 0 0 7 7.78
r"""'Beaver 6 7.23 0 0 6 6.67
Muskrat 4 4.82 1 14.29 5 5.56
Snowshoe hare 7 8.43 0 0 7 7.78
Microtine 6 7.23 2 28.57 8 8.89
Unidentified small mammal 4 4.82 0 0 4 4.44
Bird 0 0 0 0 0 0
Fish 2 2.41 0 0 2 2.22
".,..Vegetation 0 0 0 0 0 0
Wolf 0 0 0 0 0 0
Unknown 4 4.82 0 a 4 4.44
Totals 83 100.00 7 100.00 90 .99.99
Grouped Data for 64 Scats
,~
Food Item iF %
Ungulate 54 60.00
Small Mammal 30 33.33
Other 2 2.22
Unknown 4 4.44
Total 90 99.99
-
.-.
53
.....
~~
Table 16.Incidence of food remains in wolf scats collected at the Tolsona wolf
rendezvous site (old Mendelta 2nd den)occupied summer 1981.
Item in 16 Items in 15 Items in 31
Food Item Adult Scats Pup Scats Combined Scats
if %II %II %-Moose,calf 0 0 1 4.35 1 2.33
Moose,age unknown 4 20.00 3 13.04 7 16.28
Caribou,adult 6 30.00 0 0 6 13.95
~
Caribou,calf 1 5.00 1 4.35 2 4.65
Caribou,age unknown 1 5.00 0 0 1 2.33
Moose or caribou 0 0 1 4.35 1 2.33
Beaver 2 10.00 0 0 2 4.65 ""'"
Muskrat 0 0 1 4.35 1 2.33
Snowshoe hare 1 5.00 1 4.35 2 4.65
Microtine 3 15.00 3 13.04 6 13.95 -Unidentified small mammal 2 10.00 5 21.74 7 16.28
Bird 0 0 1 4.35 1 2.33
Unknown 0 0 6 26.09 6 13 .95 ~
Totals 20 100.00 23 100.00 43 100.00
Grouped Data for 31 Scats ~
Food Item II %
Ungulate 18 41.86
Small Mammal 18 41.86
Other 1 2.33
Unknown 6 13.95
Total 43 100.00
-
-
54
-
Table 17.Chronological summary of kills at which the Tyone Creek wolf
pack (2 adults,6 pups)was observed from January through
~December 1980 in Game Management Unit 13 of southcentral
Alaska.
Kill Percent Estimated Date
Made Species and Age Consumed of Kill or Comments
Date of
Observation Time
95%1/7-8
Still on kill of 1/9
Caribou -adult M
1/28
Dug up old kill
Visited caribou kill
of 1/25/80
Visited old unidenti-
fied kill
2/11 or 2/12
still on kill of 2/12
2/19
2t23/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 two ptarmigan
3/8
Still at kill of 3/8
Still at kill of 3/8
Still at kill of 3/8
3/13
3/16
Return to kill of 2/24 2/
Return to old kill 2/-
Return to kill of 3/14 2/
Return to kill of 3/8 2/
Return to kill of 3/14 2/
On old moose kill l/
>5%
5%
25%
60%
60%
60%
95%
100%
100%
100%
Moose -calf 95%
Moose -calf
Moose -adult F 75%
Moose -calf 50-70%
100%
Moose -calf
Moose -calf
Moose -adult
Moose -calf
Moose -calf
Moose -adult F
10:50am No
12:20pm Yes
11 :30am No
11:30am No
10:35am Yes
11:51am No
11:25am Yes
2:06pm No
10:56am No
10:00am No
3:05pm No
2:20pm Yes
10:30am No
5:22pm No
1:00pm No
10:10am No
8:50am Yes
No
12:00 m Yes
2:30pm No
9:15am No
No
No
10:15am No
4:30pm No
11 :OOam Yes
10:30am Yes
11 :45am No
9:45am No
10:00am No
11:00am Yes
1:00pm Yes
9:30am No
9:05am No
2:27pm No
12:55pm No
11:13am No
10:04am No
1/9/80
1/11/80
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
3/27/80
3/29/80
3/31/80
4/4/80
4/6/80
4/8/80
-
2/Pack of four pups.
55
pack members were not observed on a fresh ungulate kill until the
--------.
following year after the four remaining pack members had become
integrated into other packs.No scats were collected in 1980 or
1981 because the pack did not den.
Watana Pack During 1980 and 1981 moose comprised a majority
(74%)of the 19 kills observed (Table 18).Calf moose comprised
29 percent of the moose kills,however,moose of unknown age and
adults each comprised 36 percent of the moose kills.
Summer food habits of the Watana Pack durin~1980 and 1981 as
determined from analysis of scats collected at den and rendezvous
si tes are presented in Tables 19 through 21.At the den site,
calf moose comprised the single most important food items during
both 1980 and 1981,47.5 and 28.9 percent,respectively.Occur-
rence of adult moose increased from 18.4 to 38.7 percent from the
1980 den to the 1980 rendezvous site,suggesting that movement to
the latter area may have been related to moose distribution.No
scat data were available from the 1981 rendezvous site.Regard-
less,ungulates comprised the bulk of the food occurrences;71
and 52 percent for 1980 and 1981,respectively.Small mammals
appeared to be more important in 1981 since their occurrence rose
from 9.2 to 31.1 percent.
Summary of Food Habits
Table 22 summarizes summer food habits of all packs studied as
determined from analysis of scats collected at den and rendezvous
sites during 1980 and 1981.Moose of all ages were the most im-
portant summer food item during both years of study.However,
the importance of calf moose is believed to be over-emphasized by
scat analysis (Ballard et al.1981),perhaps because of the
higher of surface to volume ratio of smaller animals (Floyd et
al.1978).
Studies of wolf food habits in GMU-13 since 1975 have suggested
that moose are the single most important food item (Ballard et
56
-
Table 18.Chronological summary of kills at which the Watana wolf
pack was observed from mid-April 1980 through 30 October 1981
-in Game Management Unit 13 of southcentral Alaska.
-Date of
Observation
Kill
Time Made Species and Age
Percent
Consumed
Estimated Date
of Kill or Comments
1:16pm Moose ~adult
2+wolves
1 wolf (310)1../
7 wolves
1-2 weeks old?3 wolves !/
3 wolves 1/
3 wolves 1/
Old kill -3 wolves l/
3 wolves 1/
2 wolves -
1 wolf
2 wolves
?Could have been
a tagging mort.
At least 1 wolf
present.
Return to kill
of 5/13.
6/1 4 wolves
Old kill from past winter.
9/9
9/15-16
from
50%
100%
25%
50%
75%
100%
50%
60-75%
50%
90+%
75%
50%
cornered
Caribou -adult
Moose -?
Moose -adult?
Caribou-?
5 wolves
Old kill made by black
bear several days earlier
Moose -?Probable kill 1 wolf
Unknown sp.Kill in thick alder 2 wolves
Wolves in same location at pm on 6/13.Bear within 35 m.
Wolves in same location as 6/13.Bear close by.
1 wolf returned to location of 6/13.
Moose -calf 50%Wolves had cow surrounded
Moose -adult 100%1 wolf
Moose -adult
Moose -?
Moose -adult ?
Moose -adult
Had cow w/2 yearlings
Moose -newborn calf
Moose -adult
Moose -yearling
Moose -calf
Caribou -adult
Still on ill
Suspect wolves departing
kill made by black bear
Caribou -?8:50pm
5:20pm
6:20pm
3:35pm
9:25am
7:45pm
12:50pm
7:30pm
9:00am
12:59pm
10:43am
12:55pm
10:30pm
4:30pm
12:50pm
4:50pm
2:30pm
12:llpm
9:15am
2:30pm
8:55am
2:15pm
5/13/80
5/14/80
6/2/80
6/13/80
9/10/80
9/16/80
1/17 /80
3/6/81
3/7 /81
5/14/81
5/17/81
5/23/81
5/24/81
5/27/81
6/1/81
6/2/81
6/3/81
6/5/81
6/6/81
6/7 /81
6/13/81
6/14/81
6/17/81
6/19/81
6/19/81
6/21/81
-
-
!/Wolves 310,344 and 345 had dispersed from Watana Pack.
~57
Table 19.Incidence of food remains in wolf scats collected from the Watana pack den
occupied during late spring and early summer 1980 in GMU-13 of southcentral
Alaska.
Item in 33
Food Item
Moose,adult
Moose,calf
Moose,age unknown
Beaver
Snowshoe hare
Bird
Vegetation
Wolf
Unknown
Totals
Adult
It
12
16
o
o
o
3
o
o
4
35
Scats
%
34.29
45.71
o
o
o
8.57
o
o
11.43
100.00
Items in 104
Pup Scats
#%
14 13.21
51 48.11
7 6.60
9 8.49
4 3.77
o 0
4 3.77
3 2.83
14 13.21
106 100.00
Items in
Combined
if
26
67
7
9
4
3
4
3
18
141
137
Scats
%
18.44
47.52
4.96
6.38
2.84
2.13
2.84
2.13
12.77
100.00
Grouped Data for 137 Scats
Food Item
Ungulate
Small Mammal
Other
Unknown
Total
It
100
13
10
18
141
%
70.92
9.22
7.09
12.77
100.00
58
-
-
-
Table 20.Incidence of food remains in wolf scats collected at the Watana pack
rendezvous site which was used by the Watana pack during summer 1980
i'in GMU-13 of southcentral Alaska.
Item in 33 Items in 23 Items in 56
Food Item Adult Scats Pup Scats Combined Scats
It %It %It %
Moose,adult 17 48.57 7 25.93 24 38.71
Moose,calf 8 22.86 5 18.52 13 20.97
Moose,age unknown 0 0 3 11.11 3 4.84
Moose or caribou 2 5.71 4 14.81 6 9.68
Beaver 2 5.71 0 0 2 3.23
Vegetation 1 2.86 4 14.81 5 8.06
\Jolf 1 2.86 0 0 1 1.61
Unknown 4 11.43 4 14.81 8 12.90
Totals 35 100.00 27 100.00 62 100.00
Grouped Data for 56 Scats
Food Item
Ungulate
Small Mammal
Other
Unknown
Total
It
46
2
6
8
62
%
74.19
3.23
9.68
12.90
100.00
59
Table 21.Incidence of food remains in wolf scats collected at the Watana pack
den occupied during late spring and summer 1981 in GMU-13 of southcentral
Alaska.
Item in 73 Items in 31 Items in 104
Food Item Adult Scats Pup Scats Combined Scats
1/%11 %11 %
Moose,adult 15 15.46 -0-0 15 11.11
Moose,calf 26 26.80 13 34.21 39 28.89
Moose,age unknown 8 8.25 3 7.89 11 8.15
Caribou,adult 3 3.09 1 2.63 4 2.96
Caribou,calf 1 1.03 0 a 1 0.74
Beaver 8 8.25 4 10.53 12 8.89
l1uskrat 4 4.12 4 10.53 8 5.93
Snowshoe hare 5 5.15 3 7.89 8 5.93
Mircotine 8 8.25 2 5.26 10 7.41
Unidentified small mammal 4 4.12 0 0 4 2.96
Bird 1 1.03 0 0 1 0.74
Vegetation 1 1.03 1 2.63 2 1.48
Unknown 13 13.40 7 18.42 20 14.81
Totals 97 99.98 38 100.00 135 100.00
Grouped Data for 104 Scats
~,
-
Food Item
Ungulate
Small Mammal
Other
Unknown
Total
70
42
3
20
135
%
51.85
31.11
2.22
14.81
99.99
60
~
I
-
Table 22.Comparison between years of food remains in wolf scats collected at
den and rendezvous sites in 1980 and 1981 from GMU-13 of
southcentra1 Alaska.
Food Items 1980 1981
,~727 Scats·290 Scats
It Items %Occurrences it Items %Occurrence
Adult Moose 105 12.00 24 6.15
Calf Moose 369 42.17 87 22.31
Moose,Age Unknown 22 2.51 21 5.38
Adult Caribou 30 3.43 31 7.95
Calf Caribou 13 1.49 19 4.87
Caribou,Age Unknown 8 0.91 5 1.28
Moose or Caribou 31 3.54 9 2.31
Beaver 48 5.49 37 9.49
Muskrat 26 2.97 24 6.15
Snowshoe Hare 55 6.29 21 5.38
Micro tine 40 4.57 37 9.49
Unid~ntified Small Mammal 15 1.71 20 5.13
Bird 16 1.83 8 2.05
Fish 1 0.11 2 0.51
Vegetation 22 2.51 5 1.28
Wolf 4 0.46 1 0.26
Unknown 70 8.00 39 10.00
"""I
Total 875 100.00 390 100.00
N!ilIlM',
61
al.1981).These studies suggested that wolves were selecting
adult moose during half the year except for the months of January
through July,when short and long yearling moose comprised a dis-
proportionate percentage of the kill (op.cit.).This trend ap-
pears to have continued in 1980 and 1981 as well,except that
caribou appear to have increased in importance as a prey item.
Table 23 compares the percentage of moose,caribou,and miscel-
laneous prey observed annually for different radio-collared wolf
packs studied in GMU-13 from 1975 through 1981.According to
this comparison,the annual percentage of observed caribou kills
has varied from 4 to 30 percent.Excluding 1978 when the main
body of the Nelchina caribou herd wintered in the Wrangell
Mountains,and few were available to GMU-13 wolves during winter,
the importance of caribou in the diet of wolves appears to have
increased since 1978;average of 18 percent caribou for 1975
through 1977 in comparison to 26 percent caribou for 1979 through
1981.Some of the annual difference in percent occurrence of
caribou could be attributed to the studying of different packs
where caribou have always been relatively more available to
wolves.However,caribou distribution is probably at least in
part a function of thei r density.The Nelchina herd reached a
record low of approximately 10,000 in 1973 (Pitcher,pers.
corom.) .By 1981 the population had increased to approximately
22,000 (op.cit.).We suspect the increase in the caribou popu-
lation generally has made caribou more available to wolves
throughout the Unit.If true,this would suggest that as the
herd grows larger,caribou will also become more important as
wolf prey.Assuming wolf populations in GMU-13 increase slightly
or remain stable,a larger caribou population may have some posi-
tive benefits for moose in that a larger percentage of the may
will be caribou,relieving the moose population from some pre-
dation mortali ty.
62
~I
1]
Table 23.Comparison by wolf pack of the annual percentage of observed wolf prey from 1975 through 1981 in the Susitna and Nelchina River Basins of
southcentral Alaska.
%Moose Kills %Caribou Kills %other kills Totql Kills
Year 1975 1976 1977 1978 1979 1980 1981 1975 1976 1977 1978 1979 1980 1981 1975 1976 1977 1978 1979 1980 1981 1975 1976 1977 1978 1979 1980 1981
Pack
Brushkana 100 -- --- -
0 ---- - -
0 - --- -
-4
Butte Lake 75 100 -----25 0 --- - -
0 0 - - ---4
'Deadman 100 - ---- -0 ------0 - - - -- -
3
Deep Lake -50 33 60 ----50 67 20 0 0 0 -0 0 20 0 0 0 -6 3 10
Delta 100 - --- - -0 --- ---0 ------2
Ewan 42 33 -- - --50 67 -- ---8 0 -----12 6
Fish Lake ---- - -
29 - -----43 ------29 -- - - - -7
Hogan Hill 100 89 100 90 ---0 11 0 0 - --0 0 0 10 - --12 9 3 10
Jay Creek -50 100 100 - - --50 0 0 -- --0 0 0 --- -2 4 4
Key Creek 60 82 -100 - - -10 18 -0 ---0 0 -0 ---10 11 -7
Maclaren 82 100 -----18 0 - ----0 0 - ----11 1
0'Mendeltna -57 55 -----14 28 --- --29 17 -- -- -7 29
W Middle Fork --100 - --- --0 - - ----0 ------5
Saint Anne's - - -
75 67 100 -- -
0 0 0 0 - --20 25 33 0 -- -
5 8 3 6
Sinona 100 100 80 75 ---0 0 0 0 - --0 0 20 25 - --11 19 5 4
Susitna -- --86 41 67 - ---9 53 17 - -- -
5 6 17 -- - -
22 17 6
Sinona-Susitna - - -- -
50 0 - -- -
-25 100 - ----25 0 -----4 1
Tolsona - --100 17 45 67 - --0 67 27 33 ---0 17 27 --- -3 12 11 9
Tyone --80 100 100 91 --,-0 0 0 9 ---20 0 0 0 - - -
5 1 11 11
Watana - --- -
60 80 ----40 13 - - ---0 7 -----5 15
Totals 80 77 69 83 71 61 63 19 19 17 4 21 30 26 1 3 14 13 8 9 11 69 62 59 47 48 54 38
(
Table 24 lists the species,sex,age,condition,and cause of
mortali ty for kills examined in situ during 1980.These data
were combined with those collected in GMU-13 from 1970-72 and
1975 through 1979 in an effort to assess the age and condition of
prey taken by wolves.Analysis of this data was provided by
Ballard et ala (1981).Generally,it was concluded that wolves
were preying upon relatively heal thy calf and short yearling
moose.During severe winters wolves preyed upon relatively
healthy adult moose in proportion to the occurrence of their age
class in the population while during average or mild winters
older adult moose were being preyed upon.
During 1980 both the mandible and longbone were collected from
moose and caribou dying from various causes.These data were
jointly analyzed with those collected from 1978 through 1979 to
determine the relationship in marrow fat between the two bones.
A technical paper comparing mandible marrow fat with longbone
marrow fat was prepared and presented at the 18th North American
Moose Conference and Workshop held in Thunder Bay,Ontario.A
c'opy of the paper is presented in Appendix C.
Predation
Winter Rates
The most serious impact of the proposed project on wolves could
be reduction in prey numbers.In an effort to evaluate the af-
fects of prey reductions on wolves it is necessary to determine
the numbers and types of prey consumed.
Winter predation rates were estimated for three packs by inten-
sive radio monitoring and back tracking.A detailed discussion
follows:
64
-
]J "1 "'1 '~l "J
Table 24.Age,sex,condition (as determined by percent fat)and cause of mortality of
moose and caribou kills examined in GMU 13 of southcentral Alaska during 1980.
J 1 I
Accession Date of Approximate Percent Fat Marrow
Number Age Sex Collection Location Longbone Ramus Color Cause of Death
Species Moose
120601 2 F 01/28/80 Kenny Lake 90.4 68.1 Pink Accident
120602 Calf ?02/21/80 Susitna Lake 24.0 -_.Pink Wolf predation
120603 Adult calf ?02/21/80 Oshetna River 46.5 --Pink Wolf predation
120604 Calf ?01/12/80 Sinona Creek 89.7 --Pink Wolf predation
120605 Adult 02/20/80 Black River 84.5 61.6 Pink Wolf predation
120606 Calf ?02/21/80 Little Nelchina R.53.3 41.0 Pink Wolf predation
120607 2 ?02/21/80 Old Man Lake 84.1 65.2 Pink Wolf predation
120608 Calf ?02/21/80 Little Nelchina R.Wolf predation
0'1 120609 10 F 02/21/80 Grayling Lake 88.1 72.6 Red Unknown
VI 120610 Adult F 02/04/80 Eureka 60.8 --Pink Potlach kill
120611 Adult F 04/01/80 Kenny Lake 98.1 --White Shot
120612 Adult F 03/18/80 Tyone Mountains 81.2 --Pink Wolf predation
120613 14 F 03/13/80 Black River 83.7 67.2 White-Wolf predation
red
120614 Calf ?03/25/80 Squaw Lake 57.1 44.9 Red Wolf predation
120615 Calf ?03/26/80 Black River 61.2 60.0 Pink Wolf predation
120616 12 F 03/26/80 Oshetna River 87.3 65.8 Pink-Wolf predation
red
120620 Adult F 04/22/80 Watana Creek 83.9 54.6 Pink Tagging
120657 Calf F 04/30/80 Glenn Highway 5.9 9.1 Pink Winter kill
120658 16 F 03/08/80 Sinona Creek 85.6 77 .1 Pink-Wolf predation
red
120659 Calf F 03/08/80 Sinona Creek 33.9 --Pink Wolf predation
120660 Adult F OS/21/80 Glennallen 15.0 --Pink Winter kill
Species Caribou
55121 Adult F 01/27/80 Susitna Lake 90.5 --Pink Wolf predation
55122 Calf ??02/03/80 Minnesota Lake 75.6 --Pink Wolf predation
55123 Adult M 02/21/80 Tolsona Creek 29.4 --Pink Wolf predation
Susi tna Pack
From 23 January through 27 March 1980 members of the Susitna pack
were observed on nine kills.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 (three adults,two yearlings and two pups).During this
interval they preyed upon four caribou and one adult moose for a
kill rate of 1/4.2 days.Caribou comprised 80 percent of the
kills in 1980 while in 1979,all of the observed prey were moose.
Differences appeared to be related to the availability of prey
because in 1979 few,if any,caribou had been available to this
pack while in 1980 relatively large numbers of caribou overwin-
tered in thi s pack 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
Susitna Lake.Based upon kills observed during this time period,
this pack of seven wolves had 5.3 kg of available food/wolf/day.
During the second sampling period from 12 March through 27 March
1980 this pack numbered four wolves,providing an opportunity to
compare kill rates for the pack when at a lower number.Kills
were comprised of one adult moose,one calf moose,and two adult
caribou.The kill rate was 1/4.0 days,which provided 5.7 kgs of
available food/wolf/day,which was fairly close to the rate of
kill observed when the pack included seven members.
In an effort to roughly determine possible impacts of this wolf
pack on moose,a moose survey in the pack area was conducted in
late March.Four and one half hours of flight time (0.59 min-
utes/mi 2 )were spent surveying this 462 mi 2 area.A total of 51
moose were counted:43 adults and eight calves (15.7%).The
observer subjectively estimated that he may have observed 25 per-
66
-
""""
-
-
.....
--
-
r-
!
cent 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 per-
cent of the calf moose counted in March after most of the pre-
dation had occurred.If the observer had indeed counted only 25
percent of the moose and if we include 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 appeared to be con-
tributing to high mo'rtali ty of calf (short yearling)moose in the
Susi tna pack terri tory.
Tyone Creek Pack
During early 1980 weather and tracking conditions were excellent,
allowing this pack of two adults and six pups to be monitored
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 17).The prey species used by this pack
were similar to those observed in 1979 when they were comprised
of two adults.However,in 1979 when the pack was comprised of
two adults,calf moose (short yearlings)comprised only 29 per-
cent of the kills while in 1980 when the pack numbered eight
wolves,calves comprised 64 percent of the kill,possibly indi-
cating 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
wi th an estimated 4.9 kgs of food available/wolf/day.
As with the terri tory of the Susi tna pack,moose in this pack
area were counted in early March 1980.Four and one half hours
(0.89 minutes/mi 2 )were spent surveying the 302 mi 2 pack area.A
total of 266 moose were counted:221 adults and 45 calves (17%).
The observer subjectively estimated that he had observed 50 per-
cent of the moose present.The observed moose kill rate was ex-
trapolated for the months of December through April,yielding an
estimated ki 11 of eight adult moose and 20 calf moose.These
67
projections comprised 4 percent of the adult moose and 44 percent
of the calf moose observed during the survey.Assuming only 50
percent of the moose were observed during the survey and adding
the projected kills to 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 calves (short yearling)in this area was a
significant mortality factor.
Tolsona Pack From 9 January through 26 January 1981 we at-
tempted to determ.ine the predation rate of the Tolsona Pack on
caribou by attempting to locate the pack every other day.During
this period the pack numbered 14 to 15 wolves and appeared to be
feeding quite heavily on caribou.A total of four kills were
observed (Table 25)i 2 adult caribou,1 adult moose and 1 calf
moose.Unfortunately we were unable to locate the wolves at the
intensi ty we desired because of inclement weather.Based upon
these data,this pack of 15 wolves preyed upon a moose or caribou
at the rate of 1/4.5 days,and had 39 kgs.of edible prey avail-
able per day at a rate of 2.6 kgs./wolf/day.Comparison of this
rate with other predation data (this study and Ballard et al.
1981)reveals that the rate was well below the rate believed nec-
essary for the pack to maintain its size and productivi ty (3.6
kg/wolf/daYi from Mech cited in Peterson 1977).Therefore,we
believe that several kills were missed during the study period.
Our observation between 23 and 24 January 1981 when the pack had
killed,consumed,and traveled a minimum of 2 miles in a 24 hour
period,suggests that a monitoring intensity of every other day
is too infrequent for determining predation rates for packs which
rely heavi lyon caribou.In such cases,we recommend that packs
be moni tored twice dai ly if possible.
Summer Rate
Watana Pack -During May and June 1981 an attempt was made to
locate members of the Watana wolf pack daily in an effort to as-
ses's summer food habits and predation rates.At the ini ti ation
68
....
c ~-~]'--1 ']1 ]
Table 25.Daily movements.observed prey and location of the Tolsona wolf pack from 3 through 26 January 1981
in GMU-13 of southcentral Alaska.
Table 25.(cont'd)
1.D.of
Radioed
Distance between obs.&Final Wolves il Wolves Kill Species Misc.
Dates and Time AEpx,direction traveled Location Present Observed Made of Prey Notes
220
1/17/81 -1/18-81 2 mi.NE Marsh 316
0919 12:40 4 mL NW Lake 316 14 No
1/18/81 -1/20/81 220
12:40 1210 7 mL S Mendelta Sp.315 14 No
316
1/20/81 -1/21/81 220
12:10 1015 3 mi.E Old Man Lake 315 7 Yes Calf Est.75%
316 Moose consumed
1/21/81 -1/22/81 220
1015 1000 2 mL N.2 mL N.of 315 15 No......1/21 location 3160
1/22/81 -1/23/81 220
1000 0948 10 mi.NE 2 mi.SW upper 315 6 No Unable to back
Tolsona Ck.316 track
1/23/81 -1/24/81 220 Adult Kill made 4 mi.
0948 1025 6 mL ESE Tolsona Ck.315 12-13 Yes caribou east of the
316 1/23.Kill est.
consumed.
1/24/81 -1/26/81 220
1025 1445 3 mL W W.of 1/24 315 12 No
location 316
J ;)J J )J 1 J J 1 J ))
of the study 3 members (#310,344 and 345-see Watana Wolf Pack
History section of this report)split off from the pack,leaving
the main pack wi th 8 members.
From 10 May through 23 June 1981,the eight members of the main
Watana Pack were observed on only 6 kills (see Table.16 and 32).
~
The kills were comprised of 2 adult caribou (1 was unclassified
and assumed to be an adult),2 calf moose,1 adult moose,and 1
unknown species.In addition to the above,pack members were
known to have twice visited an adult caribou which had been
killed by a black bear,revisited one old moose kill once,and
visi ted the unknown species kill on three separate occasions.
Using the live weights and estimated consumable flesh given by
Peterson (1977)and Ballard (1982)a gross estimate of the kilo-
grams of edible prey/wolf/day was derived based on the observed
kill rates.Based upon the assumption that the unidentified kill
was an adult moose and that the wolves were able to derive 10
percent of the edible flesh from the old revisited kills,this
pack had 946 kg of prey available at 21.01 kg/day for a 45 day
j"period.The derived consumption rate of 2.6 kg/wolf/day is well
below that necessary to sustain adult denning wolves and their
pups.Also,the observed kill rate of 1 kill/7.5 days is well
below the winter rate of 1/4-5 days.
Peterson (1981)believed that summer wolf predation rates were
lower than those of winter.Studies of two wolf packs elsewhere
in GMU-13 suggested that predation rates were equal or greater
than those observed during winter.We suggest that the low sum-
mer rate for the Watana Pack was the result of poor visibility
and not having radio-contact wi th the alpha male.
During the 45-day study,radio-collared wolves from the main
Watana Pack were only observed on 55 percent of the occasions
they were located away from the den site (presumed hunting).
Therefore,on the 45 percent of the occasions when the wolves
were unobservable,they could have been present at a kill which
in all likelihood also would not have been unobservable.Unfor-
71
tunately as the study proceeded,it became obvious that the alpha
male was not radio-collared.During earlier studies (Ballard et
al.1981a,b)it had been determined that whenever a large ungu-
late was killed during the denning season,the alpha male was
always·present.Therefore we believe the kill data grossly
under-represents the food habits of the Watana Pack.Poor ob-
servability and lack of having enough members radio-collared did
not,however,hamper efforts at determining the food habits and
predation rates for a non-denning pack.
From 10 May through 28 May 1981,wolves 310 (adult male),;344
(yearling male)and 345 (yearling male)functioned as a distinct
pack away from the main WatanaPack from which they had dispersed
(Table 25).Prior to wolf 310 visiting the Susi tna den,the
three adult wolves were observed on a total of 4 kills over a 19
day period;kill rate of 1/4.8 days.The kills were comprised of
2 adult moose,1 yearling moose,and 1 calf moose.In addition,
the wolves were observed attacking a cow moose with twin year-
lings,but were unable to confirm whether they in fact made the
kill.Based upon the observed kill rate,this pack of three
adults had available 819.5 kgs of edible prey or 43 kgs/day (14.4
kg/wolf/day).This particular set of data do not support the
hypothesis of smaller packs having lower predation rates.
A controversy exists in the literature concerning the predation
rates of packs of various size.Peterson (1977)maintains that
predation rates for packs of 9 or more individuals remain con-
stant.Ballard et al.(1981)demonstrated that pack size of from
2 to 9 individuals did have differing predation rates.If large
packs (15 individuals)do in fact have higher rates of predation
than smaller packs,the significance of wolf preation to an ungu-
late population could differ from pack to pack and could have a
pronounced effect on how managers manipulate wolf popUlations.
More predation rate data on packs of various sizes is needed be-
fore this controversy can be resolved.For purposes of this re-
port,we assumed a year-round predation rate of 1 kill/5.0 days.
72
~I
-
Wolf Territories,Population Numbers,and Numbers to be Impacted
--~-------~---------
F'"by the Project.
For the purposes of this report Etkins (1964)definition of ter-
ritorality was used;"any behavior on the part of an animal which
tends to confine ...its movements to a particular location."Most
~
defini tions of terri torali ty assume that the terri tory is de-
fended against intruders.Although wolves in the Susitna and
Nelchina River Basins apparently do at times defend their area
against other wolves,intrusions into a neighboring terri tory
often occur when the home pack is not using that portion of the
area.
Table 26 summarizes territory sizes for the six wolf packs which
have been intensively investigated for Susi tna hydroelectric
studies.Terri tory sizes for the six packs averaged 545 mi 2
(1414 km 2 )which was almost identical with sizes determined for
other wolf packs in GMU 13 (Ballard et aL 1981).
-i
1
I
If"'"
1
:1
'F'"
'I'I
j
r
i
I
Figure 4 depicts the spatial arrangement of known and suspected
wolf terri tory boundaries in the proj ect area during 1980 and
1981.Based upon track counts,public sightings,and radio lo-
cations of radio-collared packs and previous studies,at least
six and perhaps seven wolf packs occupy portions of the Susi tna
River which would be directly impacted by the Devil Canyon or
Watana impoundments.The packs are Portage Creek Pack,Watana
Pack,Upper Talkeetna or Stephan Lake Pack,Jay Creek Pack,Tyone
Creek Pack,Talkeetna River Pack and the Devil's Canyon Pack.
Additionally,two packs (Fish Lake pack and the Susitna-Sinona
pack)were in part the result of dispersal from the aforemen-
tioned packs.Had these packs been at reduced densities,these
dispersals may not have occurred and these pack areas could have
remained vacant.It is interesting to note that even under the
intensive harvest pressure exerted by ADF&G personnel from 1976-
78,not all wolves were removed from along the Susitna River.At
that time it was suggested that wolves occupying the Susi tna
73
Table 26.Summary of territory sizes for wolf packs studied as
part of the Susitna Hydroelectric Project studies during
1980 and 1981 in southcentral Alaska.
Area Territory Size
mi 1:/(km '!:.-/)
Fish Lake 346 943
Susitna 561 1,453
Susitna-Sinona 466 1,208
Tolsona 981 2,541
Tyone Creek 364 943
Watana 534 1,383 """
x 545 1,412
s.D.209 541
~
74
1
e·uepected wolf
paok.and
ooncentratlon area
ttl
Figure 4.Suspected locations and territorial boundaries of wolf packs Inhabltatlng the Susltna Hydroelectric
Project area during 1880 ,and 1881.
.......
111
Ri ver bottomlands provided a reserve population for emigration
--~-- -----------
into areas where harvesting either eliminates packs or severely
reduces population densi ty (Ballard and Spraker 1979:57).
Wolf territories were essentially nonoverlapping during the
course of any particular year (Ballard et ale 1981).What over-
lap did occur was either seasonal in nature or was the result of
the manner terri tories were plotted.
Numbers of wolves estimated to occur in 13 wolf packs known to
occur in the study area are presented in Table 27.Spring 1980
and 1981 estimates represent the post-hunting population while
those in fall represent gains due to reproduction and dispersal
prior to hunting and trapping losses.Because the Susitna River
frequently serves as a travel corridor and a concentration area
for wolf packs,inundation will probably cause a drastic reduc-
tion in wolf numbers due to both direct loss of prey,loss of
escape cover from hunting and trapping acti vi ties,and loss of
habitat.Six to seven wolf packs would be directly impacted by
the proj ect due to those factors.
Figure 5 compares the distribution of known and suspected wolf
packs occupying the Susi tna and Nelchina River Basins with the
documented locations of radio-collared caribou from the main body
of the Nelchina herd,located during 1980 and 1981.(Note:
boundaries and suspected existence of wolf packs west of Kosina
and Deadman Creeks are the least reliable of data presented).
Since wolves depend upon caribou for from 20 to 30 percent of
their annual diet,any disruption of caribou migration or reduc-
tions in herd size due to the proposed project potentially could
severely impact wolves in most of Game Management Unit 13 and in
the west half of Game Management Unit 11.Of the 24-26 packs
known to be present in GMU-13 in fall 1980,6 to 7 packs would be
directly impacted and an additional 15 packs could be adversely
affected by reductions in either caribou numbers or migration of
the main Nelchina herd as a result of the proj ect.If caribou
76
--
Table 27.Estimate of numbers of wolves by individual pack inhabiting the Susitna
Hydroelectric study area in spring and fall 1980 and 1981.
f"'"
I
r
.....
Spring 1980
Pack Area (Post Hunt)
Butte Lake 3-41
Fish Lake 1
Jay Creek 6
Keg Creek 1
Maclaren River 2
Portage Creek 1
Stephan Lake 2+
Susitna 4
Susitna-Sinona 4
Tolsona 9
Tyone Creek 4
Upper Talkeetna River ?
Watana,5
Total 40
Fall 1980
(Prehunt)
3-4+
2
7-81
1
4-5
1
11
10
4-5
16
2
1
14
77
77
Spring
1981
3
9
1
2-3
1
1
1
·5
2
13
o
?
8
42-43
Fall
1981
5
12+
10
2-3
2-3
6
1
4
1
15
?
2
14
72-74
"'-I
~
.
).
J
~~?JI~~auapected wolf...:....:...pack and
concentration area
*location of radlo-
collared caribou
Figure 6.Distribution of main Nelchlna radio-collared caribou,14 April 1980 through 29 September,1981 (from Pitcher
1981)In relallon to known and auspected wolf packs and concentration areaa within the Susltna and Nelchlna River
Baalna of aouthcentral Alaaka <from this ~tudy and Ballard et al.1981).
1 ))J J I ~.~l I ~1 j J .~J
r
!
.-..
I
I
r
[
r
~
I
I
subherds were severely impacted by the proj ect,the approximate
number of wolf packs which could be impacted can be determined
from Figure 5.
Migration and movement patterns of moose are compared with known
and suspected wolf territories in Figure 6.In addition to the
six or seven packs which would directly be impacted by the pro-
j ect,a minimum of 7 to 8 additional wolf packs would be in-
directly affected by either disruption of moose movements and mi-
gration and/or reductions in moose density.These impacts could
be severe since wolves depend on moose for 60 to 70 percent of
their annual diet.In summary,6 to 7 wolf packs would be
directly impacted by the proj ect,an additional 7-8 packs would
be indirectly impacted by either reductions in density or move-
ments of ei thermoose or caribou,and an additional 7-8 packs
could be indirectly impacted by reductions in numbers or distri-
bution of caribou.
Impacts of Wolves on Moose and Caribou
Based upon an intensive census of the study area in fall 1980
(Figure 7)we estimate that a portion of the wolf study area con-
tained 1,985 moose (see Upstream Moose Report).They are cen-
sused roughly corresponds'to an area which would be occupied by
five wolf packs.Using the census and stratification moose data
and our estimate of five wolf packs,we attempted to assess the
importance of wolf predation to the study area moose population.
We assumed that each pack made an ungulate kill once every 5 days
and that from 60 to 70 percent of the kills were comprised of
moose,32 percent of which were calves.Based upon these assump-
tions we estimated that wolves were annually preying upon 11 to
13 percent of the study area's fall moose population.Percent
mortality of calves present in fall ranged from 16 to 18 percent
while mortality on adults of both sexes ranged from 10 to 11 per-
cent.It ,should be pointed out that these calculations are based
on a prey base present in November and thus the mortality figures
are slightly inflated.
79
00o
.
)..)
J.' . ',~~~\~.~~~\~.
rr!!:.suspected wolf........;...pack and
concentration area
j
Figure 8.Migration routes and movement patterns of radio-collared moose In the Sualtna and Nelchlna River Baalns
(from Ballard and Taylor 1980;Ballard and Gardner 1881.1981a;Ba;lard et al.1881)In relation to known and suspected
wolf territories (from this study and Ballard et $1.1981 a).'
j II 1 I J I J I J J:I _J
.
I
j
_-l~__
Figure 7.Relative densities of moose as determined
from stratification and census flights
made on 2 through4~8 and 29 November '1980
in the Sus1tna River Hydroelectric Project '
,study area of southcentral A1ask~(from
Ballard et a1.1981 ...upstrea,m moose report).
,.
.,'
MEA,ALAlICA
OCTOBER 1111
UPPER SUSITNA RIVER BASIN
TWO 1W.t.PLAN
u.s.DEPAftTUENf ""1"£RIOA
fISH NG WI.DLJFE SERVICE
,
I
\......~<"'-1/~
)
\\.
l,
-.:?J".....;-
,--",.,
_J
.I
-=""\
\,,
J+,a
(-v1
\
)-':>
"...._..a-..,.....-..-
r
I
I
,,~~--t,.".'"'-..J "\-./.'1,
I
~~
l 1\
IIVU "UI ........
0---------.10 •ZOWlI..
o ~EN~tlTY
LO!l r:'ENSITY,
{
CENSUS AREA
_..MEDIUI\DEN~ITY
~HIGH D~NSITY
SCALE
-
-0
m····...........
00
~
Based upon calf and yearling mortality studies conducted in and
adjacent to the Susitna Study Area,it was estimated that between
9 to 24 percent of the fall calves were succumbing to wolf pre-
dation.This estimate obviously falls wi thin that range.
Appendix D contains a copy of the yearling moose mortality study
which provided an estimate of first year calf mortali ty ranging
from 22 to 84 percent depending upon the magnitude of bear pre-
dation,wolf predation and winter kill.
Determining the level of wolf predation on caribou for the study
area required a slightly different approach because of the sea-
sonalnature of caribou distribution.Caribou population numbers
between 1973 and 1981 were extrapolated by assuming a linear re-
lationship by year from a low of 10,000 animals in 1973 to a high
of 22,000 animals in 1981 (Pitcher,pers.comm.).We estimated
the impact of wolf predation on caribou by assuming that 25 wolf
packs occur within the range of the main Nelchina herd and its
subpopulations in 1981.We also estimated a mortali ty rate for
1973 and 1976 when the Nelchina herd numbered approximately
10,000 and 14,500 animals,respectively and assumed that approxi-
mately 45 wolf packs occurred in the Uni t during those years.
Further,we continued to assume a year-round predation rate of 1
kill/5.0 days and that caribou comprised 20 to 30 percent of the
annual diet.No separation between calf and adult caribou was
made because existing wolf data do not suggest selection of the
calf age class.Based upon these assumptions,we estimated that
in 1973 when the Nelchina herd reached a record low of approxi-
mately 10,000 animals and approximately 45 wolf packs occurred in
the Unit,wolf predation was estimated to account for from 7 to
10 percent annual mortality.In 1976 when the caribou population
numbered approximately 14,500 and assuming 45 wolf packs,caribou
losses due to wolf predation were estimated at 5 to 7 percent.
If only 25 packs had been present in the 1976 such as in 1981,
total caribou annual mortality due to wolf predation would have
ranged from 5 to 7 percent depending upon weather caribou com-
prised 20 or 30 percent of the wolf diet.
82
-
-
In 1981 with the herd at approximately 22,000 animals and with 25
~--_.---_..------
~wolf packs present in its range,we estimate current caribou an-
I
I
I nual mortality due to wolf predation to range from 2 to 3 per-
cent.
Wolf Hunting-Trapping Mortali ty
r
I,I
r
'I,I
Table 28 summarizes wolf harvests for GMU-13 from 1971-72 through
1980-81 seasons.During this period,annual harvests ranged from
46 to 128 wolves.Method of harvest was classified into four
categories:(1)trapped (snared or leg-hold trap),(2)ground
shot which usually involves landing aircraft and shooting same
day airborne,(3)experimental removal by ADF&G from 1976-1979,
and (4)other-which includes miscellaneous forms of mortality
such as auto collision and natural mortality.Aerial hunting
(shooting from fixed-wing aircraft)was legal only in 1971-72.
From 1971-72 through 1975-76 ground trapping was the most common
method of harvesting wolves in GMU-13,accounting for 59 percent
of the total harvest.From 1976-77 through 1978-79 ground shoot-
ing (primarily hunting from aircraft)was the most common method
of harvest.In 1980-81 trapping again was the most prevalent
harvest method due to both poor snow conditions which did not
allow wolves to be tracked from airplane and reduced wolf den-
sities.
Figures 8 through 12 depict the kill locations of wolves har-
vested from regulatory year 1976-77 through 1980-81 for both
GOOD's 13 and 11.For all packs which would be directly influ-
enced by inundation of portions of wolf pack terri tories,the
harvests by year were as follows:1976-77 -16,1977-78 -21,
1978-79 -I,1979-80 -II,1980-81 -5.
Den Sites
General location of both den and rendezvous sites located from
1975 through 1981 in GMU-13 are depicted in Fig.13.Because
83
Table 28.Summary of reported wolf harvests for Game Management Unit 13 of southcentral Alaska by method of take
from 1971 through 1980-81 season.
GMU 13 Subunits
Harvest Subunits Sub-
year Method of take 13A lJB 13C 13D lJE unreported totals Totals
1971-72 Trapped 43
Ground shot 22
Aerial shooting 46
Other 0 111
1972-73 Trapped 11 4 20 ,16 6 57
Ground shot 4 4 4 3 4 1 20
Other 1 0 2 0 0 3 80
1973-74 Trapped 15 10 8 9 11 53
Ground shot 4 4 4
6 4 22
Other 0 0
0 0 0 0 75
OJ
.I:'-1974-75 Trapped 25 5 3 21 6 60
Ground shot 7 11 3 9 11 41
Other 0 1 0 1 0 2 103
1975-76 Trapped 16 9 8 12 3 48
Ground shot 7 3 2 9 15 36
Exp.removal 6 2 0 0 17 25
Other 0 0 1 0 0 1 110
1976-77 Trapped 5 3 9 7 1 25
Ground shot 2 9 21 6 11 49
Exp.removal a 14 0 0 10 24
Other 1 1 0 1 1 4 102
1977-78 Trapped 8 2 16 20 5 51
Ground shot 27 7 20 6 8 68
Exp.removal 3 1 0 0 3 7
Other 2 0 0 0 0 2 128
1978-79 Trapped 5 2 3 6 4 20
Ground shot 0 12 30 4 1 47
Exp.removal 1 1 0 0 0 2
Other 0 0 0 0 0 0 69
J J J I ~I B ~,j I I I J »)J
1 ~4
Table 28.(cont1d)
GMU 13 Subunits
Harvest Subunits
y~a~Method of takeUA 13B Be 130 13E unreported
1979-80 Trapped 11 1 10 9 4
Ground shot 1 3 2 3 11
Other 2 0 0 0 0
Sub-
totals
35
·10
2
Totals
57
i
CXl
VI
1980-81 Trapped
Ground shot
Other
7
3
o
2
1
o
5
2
o
9
1
o
7
9
o
30
16
o 45
ex>
0\
••-,......,.,.".1'
.....~I
~Yj~L ..!,\
Diagram of hunting-trapping location of wolf harvests
1976-77 regulatory year.
,.
J ~J D 1 J I ~!•~J )I J S )J
Figure 9.
C--C-l
_ '-t~
Diagram of hunting-trapping location of wolf harvests in GMU-ll and 13 of Southcentral Alaska for
1977-78 regulatory year.
j
OJ
OJ
Figure 10.,Diagram of hunting-trapping location of wolf harvests in GMU-11 and
,197~-19 .regu1at~~y.y~ar.
J J .~_J J J I J J ))J J )J ,!
I J -1 '--'4
"
--1 "-'----'1 I
~
ex>
""
Diagram of hunting-trapping location of wolf harvests in GMU-l1 and 13 of Southcentra1 Alaska for
1979-80 regulatory year.
\
,.!
\0o
i
I
-_.""_.---=--~.._...~..~:.
Figure 12.Diagram of hunting-trapping location of wolf harvests in GMU-11 and
1980-81 regulatory year..If \)\ \1 't ,....bcd·uti .H f
1 1 J J ~,j 1 J j J )~j !I J J I
.~"'"....
1
..
~---"=1
I
HOGAN
HILLDEN1975
\..
a.,...
oc;T081:A 11ft
•HOGAN
HILL
DEN
1918
fillt NII .....n al"VI".
UPPER SUSlTN4 RIVER a.a
TWO DAM PLAN I
u.I.DEPAIITM(H1'W ..nRtO
1GIIf1lCPl1llJA.."IlM:L'M.",'"
(
.-~(-~
)
0'
\
l
\-'J--'/
r-",..,
.-J·KEG ··
_.I .CREEK
:\DEN 1915
P\,,1916,1917"
1"&EJ'
I a.:=c f,
TULSONA DEN-~"L SU81TNA RENDEZVOUS
o
D 1880,1981 &.\SITES-U80
.MENDEL.TNA .'"!O',
RENDEZVOUS ••oIftl:--~=a...,.....pUSITN~DEN-
.....1971 '1880
................--................--
-,L-WATANA
,OAMSITE
~'"\
OJ}.........,
;~'h '....\
"'0 (~,
sITe -18 10 .
SITES -1817
UPPER IUSITHA RIVER PROFILE
lINt.ilL....-...
SUSPECTED STEPHAN
LAKE DEN -1lUO'
".:.,."
TOUONA RENDEZVOUS SITIl-1ItBO
a MENDELTNA DEN -1877
MENDELTNA .'RENDEZVOUS·
ME~E"'~NA RENDEZVOUS
rAUlrUHA
~IIlI ':"'\_~...
••
~~--~-~·'L·ro!.---r-e:'l_10 Ii i~~l1'~_'m __UH •.l I ..~::...a-.~
Figure 13,General location and year of use of observed wolf .,r_~,
den and rendezvous istes discovered in the Susitna ....'M~NDELTNA DEN-~
Hydroelectric Project area from 1975 through 1981.'187.'1877
"
I,Q
f-'
,,'
some misinformed individuals believe wolf pups make desirable
----
pets,more specific descriptions of den and rendezvous site lo-
cations will not be provided in these reports.This information
will be retained on file in the Glennallen office and will be
provided to proj ect personnel upon request.
Many of the sites depicted in Fig.13 have not been examined on
the ground.Specific site descriptions are presented in Tables
29 through 31.
Den and rendezvous site usage by wolves in GMU-13 was fairly tra-
ditional.Of the 23 sites examined at least 6 have been used a
minimum of 3 seasons.Several have been used during two seasons
since 1975.The average elevation of the 23 sites was 2,550 feet
('S.D.=360)ranging from 2,000 to 3,600 foot elevation.Most den
sitea were old red fox (Vulpes vulpes)dens which had been dug
out by wolves.Most sites consisted of 3 to 4 large holes which
generally measure from 33 to 54 em high by 39 to 60 em wide.
Most sites are also characterized by the presence of several
small holes (measuring from 15 to 29 em wide)which are commonly
referred to as pup holes but which are rarely,if ever,used.
Most sites were located on slightly elevated areas with sandy
soil providing good drainage.Although we found that holes were
oriented towards all ordinates of the compass,most (x%-16.50,
p<.01)were found on south to and/or east exposures.
Thirteen of the sites we examined contained what we termed
"Whelping chambers ll which were usually located at least several
feet back from the main entrance and generally were 30 to 60 em
high,76 to 152 em wide and 89 x 152 em deep.None of the cham-
bers were lined as described by Haber (1968).
Table 32 summarizes the distances of discovered den and rendez-
vous sites from the upper water level of both the Devil's Canyon
and Watana impoundments.Four den and 1 rendezvous sites occur
within 5 miles of the average pool level of the reservoirs.The
figure contained in Table 21 must be considered absolute minimums
92
-
.....
-
'--'--C'--l
Table 29.External characteristics of wolf den and rendezvous'sites examined in GMU-13 of southcentral Alaska from 1976 through summer 1981.
Year iJ Animals Years Site Proximity
Site Site in Pack H Holes Known to be Orientation Elev.Soil to Water
Pack Visited Type Adults Pups Lg.Sml.Used of Holes (ft.)Type (m)Comments
Mendeltna 1976 Rendellvous 5 6 3 0 1976,1977 N NE 2,700 Sand Pothole
(Old Man L.)Nearby
Mendeltna 1976 Den (Old Man Lake)5 6 4 1 1976,1977 S 2,800 Sand,clay,400
1977 Den (lst)4 7 4 1 gravel (tr.)
Mendeltna 1977 Rendellvous 4 7 4 4 1977 N NE 2,900 Sand.183
(Kelly L.)clay tr.)
Mendeltna 1977 (Nicholson Lake)2 2 2 4 1977.1980.1981 SW 2,900 Sand with 69 (Pond)Elevated.
Rendezvous some gravel 274 (lake)flat.open
area probably
for sunning.
were evident
above each
\0 hole.W
Mendeltna 1977 Den (2nd)3 2 4 3 1977.1980.1981 E 2.850 Sand clay 50
gravel (tr.)
Tyone 1976 Den 7 3 1 5 1976 S (5)2.700 Sand NRa
(Clearwater)N (1 pup hole)
Susitna 1980 Den 4 6 3 4 1979. 1980.1981 . S 2.500 Sand 274 (pond)
Susitna 1980 Rendezvous 4 6 3 6 1980 S 2.475 NR 46 (swamp)
183 (lake)
Watana 1980 Den 3 6 1 1 1980,1981 S 2,300 Sand 15 (standing
(lower holes)(+2 unused water)
fox holes)
Watana 1980 Rendezvous 3 6 2 0 1980 E 2.450 Sand 15 (standing
water)
"''-.
_I J J J I •I,j I I I I J J I J D t J
U Animals Years Site
Site in Pack II Holes Known to be Orientation Elev.Soil
Type Adults Pups Lg.Sm!.U..ed of Holes (ft.)Type
Table 29.'(cont'd)
Year
Site
Pack Visited
Sinona 1978 Den
(2nd Gakona)
4
~J
6 2
=~1
3 1978
~
SW
'-'~-~i
2,000 NR
CCC4
Proximity
to Water
(m)
69 (stream)
-~,
Conunents
,/
"/
a NR =Not recorded
b Number of pups and adults not distinguished.
c Number of individuals at each site ~ot determined.
'-0
V1
(
/
Table 30.Dimensions of hole entrances and chambers for wolf den and rendezvous sites examined in GMU-I3 of southcentral Alaska from
1976 through 1981.
Pack
i Height (em)by x Width (em)
Year ,Animals in pack of Hole Entrances
Visited Site Type Adults Pups _~.Sm!.
Height (cm)
x Width (cm)
x Depth (cm)
of Chambers COlDments
Mendeltna
Mendeltna
Mendeltna
1976
1976
1977
1977
Rendezvous
(Old Man L.)
Den
(Old Man L.
Den (lst)
Rendezvous
(Kelly L.)
5
5
4
4
6
6
7
7
38 x 41
s ..2.9 4.0
n"3
49 x 60
s ..3 14.1
n ..4
37 x 44
s ..3.4 9.5
n ..4
No external opening
for any pup
tunnels
15 x 20
n ..
22 x 28
s ..6.4 3.6
n ..4
28 x 76 x 122
61 x NRa x 107
Nil.x 152 x Nil.
Holes lined with
wiregrass.
No change between
1976 &1977 for
dimensions or
number of holes.
Some tunnels and
chamber lined with
bunchgrass and
lowbush cranberry
(Vicc1nium
vitis-idea).
1.0
0"1 Mendeltna 1971 Rendezvous 2
(Nicholson L.)
2 42 x 39
s ..4.9 4.2
n"2
17.5 x 24
s ..3.7 11.2
n ..4
Nil.Holes partially
lined with wheat-
grass.dwarf birch
and fescue.
Mendeltna
Susitna
1977
1980
Den (2nd)
Den
3
4
2
6
46 x 43
14.1 17.1
n ..4
48 x 49
s ..2.0 7.0
n ..3
25 x 33
s ..12.0 3.0
n ..3
24 x 29
s ..3.8 5.8
n ..4
Nil.
Nil.x 122 x 135
Two small holes
not used.Two 2
small holes and
2 large holes
lined with fescue.
lowbush.cran-
berry &birch.
Two slDall chambers
(48 em high x 46
cm wide x 61 em
deep also located
in tunnel system.
J J J J J I J 1 )I ],-J I J J J
/
Table 30.(cont'd)
i Height (cm)by i Width (cm)
Height (cm)
x Width (cm)
Year ,Animals in Pack of Hole Entrancea x Depth (cm)
Pack Visited Site Type Adults Pups Lg.Sml.of Chsmbers Comments
Sus1tna 1980 Rendezvous 4 6 Nil.NR 48 x 86 x 89
Watana 1980 Rendezvous 3 6 35 x 57 0 66 x 128 x 99 Chambers not inter~
s ~6.0 34.9 connected;both
n • 3 67 x 116 x 145 chambers were used to
though impossible
to identify
msin chamber.
Keg Creek 1976 Den 3 5 53 x 50 28 x 38 Nil.x 112 x 69 Both chambers located
a ~6.0 7.6 s •3.5 11.3 Nil.x 142 x 69 within one·complex of
n·3 n • 2 3 holes.Complex of
5 holes did not
\0 include a chamber •......
44.7 x 47.3 30.0 x 237.2 24 x 40 x Nil.
a ..3.2 12.8 s •2.4 1.5 Nil.x Nil.x 54
n ~3 n ~4
Watana 1980 Den 3 6
Tolsona 1978 Den 3 8 36 x 51 21.6 x 20.2 Nil.Two large holes
s ~0 7.1 s ..2.3 2.9 thought to inter-
n ~2 n ..1 connect.
Hogan Hill 1978 Dell 7 5 30 x 68 20 x 25 46 x 152 x Nil.
Main Den s ..0 7.1 n ..1
.n •2
Hogan Hill 1978 Rendezvous 7 5 36 x 46 25 x 30 Nil.One "amall"hole
n ~1 s ..5.0 5.5 though.to be 1 of
n ~3 2 "main"holes.
St.Anne's 1977 Den II 6 44 x 42 46 x 152 x 122 Two holea inter-
1978 (Long Lake)
7 tot.b s ..11.0 16.0 0 connect at a point
n ~2 thought to be the
chamber.Interior /of den not lined.
Table 30.(cont'd)
",
Pack
x Height (em)by x Width (em)
Year 1/Animale in Pack of Hole Entrances
Visited Site Type Adult~~I;._].g.Sm!.
Height (em)
x Width (em)
x Depth (cm)
of Chambers Comments
St.Anne's
St.Anne's
1981
1981
Den 3 6 33 x 5 0
NiCkel Crk.s =3.8 14.1
(main)n =4
unltnuWD 3 6 54 x 54 0
s a 2.1 10.6
n a 2
33 x 104 x 152
41 x 109 x 91
36 x 122 x 122
41 x 142 x 11
61 x 91 x 61
Holes not lined.
Large ruots
bitten off.
Holes inter-
connected chambers
separated by s
large pillar of
earth.Small
chamber separated
from one of larger
chambers by small
pillsr of earth.
Holes not lined.
Sinuna 1976 Den (let)33 x 53 0 33 x 229 x NR
n -1
~
CO
Sinona 1976 Den (2nd)lid 9d 0 30 x 30 NR x 91 c 122
n a 1
Sinuna 1977 Den 2 6 39 x 52 28 x 33 NR
(1st Gakona)a -1.1 5.1 n -1
n -3
Sinona 1978 Den 4 6 36 x 54 18 x 28 NR
(2nd Gakona)8 a 7.8 9.2 s a 7.8 9.2
n a 2 n =3
a NR -Nut Recorded
b Number of pups and adults not distinguished.
c This chamber probably was not the whelping chamber though it was used during 1981.
d Number of wolves at each den site not distinguished.
Pillar of earth
located in center
of chamber.
Porcupine scats in
all holes.
,/
J J _J •J J J J I .J J I J I J ~J
~
/
Table 31.Cha~acte~lstlcs of vegetation of wolf den and ~endezvous sites examined in GMU 13 of southccntrsl Alaska from 1976 through 1981.
".
Pack
Year Pack Sbe %Cove~Overstory %Cover Understory
Vidted Site Type _./\dults Pups Ove.rst()ry Composition Understory Compoaition
Mendeltna
Hendeltna
1976
1976
Rendezvous
Den
(Old Man L.)
5
5
6
6
NRa
95
Black Spruce
(Plcea mariana)
(100%)
Balsam Popler
(Populus
balsamifera)
(90%)
White spruce
(Picea glaucs)
(lOX)
NR
100
Willows (Salix spp.)
Dwarf Birch (Betula nan a and!.glandulos..)
Graaaes (Posceae)
Fireweed (Epilobium augustifolium)
Rose (Rose acicularis)
Indian cabbage (Heracleum lanatum)
Wild Celery (Rumex app.)---
0..0
'"
Mendeltna 1977 Den
(2nd)
4 7 95 Balsam Poplar (90)100
White Spruce (10)
Graslles
Fireweed
Rose
Indian Cabbage
Mendeltna
Mendeltna
1977
1977
Rendezvous 4
(Kelly L.)
Rendezvous 2
(Nickolson L.)
7
2
10
10
Spruce (l00)
Willows (100)
90
70
Fescue (Featucs sp.)(50)Dwarf ~irch (tr.)
Wormwood (Artemisia)
Rose (tr.)
Crowberry (Empetrum niarum)(20)
Lowbush Cranberry (Vaccinium vitis~idese)(20)
WilloW (tr.)
Grasses (10)Labrador tea (Ledum palustre)
Fescue (tr.)Rose (tr.)
Dwarf Birch (40)
Willows (20)
Lowbush cranberry
Mendeltna
Tyone
1977
1976
Den
Den
3
7
2
3
95
HR
Aspen (90)
Spruce (10)
NR
100
NR
Grasses (70)
Fireweed (t r •)
Dwarf Birch
liose (5)
Indian cabbage (20)
/
Table 31.(cont'd)
'lear Pack Size :I:Cover Overstory %Cover Understory
Pack Visited Site Type Adults Pups Overstory Composition Understory Composition
low aspen
Susitna 1980 Den 4 6 50 Aspen (40)Nil.Soapberry (Sheperdia canadeusis)(tr.)
Spruce (40)Rose (20)
Fireweed (20)
Grasses (15)
,Aspen
Highbush cranberry (Viburnum edule)
Crowberry
Susitna 1980 Rendezvous'4 6 Low Density Aspen Nil.Low Aspen (10:1:)Fir';weed (tr.)
Spruce (S5%)Soapberry (tr.)Peaa (Fabaceae)(tr.)
Rose (tr)Mallses (n.)
Grasses (25%)Crllnberry (25%)
Highbush Cranberry (lOX)Crowberry (25:1:)
'Lowbush Cranberry (tc.)Blueberry (Vsccinium
ul1ginosum
Watana 1980 Den --Low Density Spruce (l00%)Nil.Dwarf Birch (30X)Willows (tr.)
(upper holes)Lowbush Cranberry (15%)Rose (tr.)
Lichens/Sphagnum (lOX)Spruce Seedlings (tr.)
Crowberry (20X)Grllsses (tr.)
Bearberry (Arctostaphylos LlIbrlldor Tea (tr.)
spp.)
to-'Watana 1980 Den 3 6 Low Density Spruce (100%)Nil.Dwarf Birch (30%)Dunchgrass (tr.)a (lower holes)Blueberry (10%)Crowberry (20X)0 Lowbush Cranberry (15:1:)Lichens a Mosses (lOX)
Rose (tr.)Labrador Tell (tr.)
Willow (t r.)Birch (20%)
Bearberry
Watana 1980 Rendezvolls 3 6 Medium Spruce (lOOX)Nil.Dwarf Birch (20X)Fireweed (tr.)
Density Blueberry (20%)Lichens +Mosses (tr.)
Lowbush Cranberry (20%)Crowberry (2SZ)
Rose (tr.)Labrador Tea (tr.)
Grasses (tr.)
",
0'])J J J I J ]I I ,~J J ~J )1 :I
~=1
Table 31.(cont'd)
"
Pack
Year Pack Size %Cover Overstory %Cover Understory
Visited Site Type ~dults _Pup~Overstory Composition Unders!Q!y ~os~tion
Sinona
Sinons
Sinona
1976
1977
1978
Den '2
1st Gakona
Den
Den
(2nd Gakona)
11
2
4
9c
6
6
NR
90
NR
White Spruce
Aspen
Spruce (tr)
Aspen
Med.to
Dense
75
NR
Dwarf Birch
Highbush Cranberry (10)
Aspen (30)
Spruce (5)
110ss (tr.)
Willow (tr.)
NR
Lowbush Cranberry (30)
Rose (10)
Fireweed (10)
Grasses (5)
Blueberry (t r .)
t-'a
N
ab NR a Not Recorded
c Numbers of pups and adults of distinguished.
Number of individuals at each den site not determined.
/
J ]_J _J -D J J J J J B .1 -_.!)D J J
Table 32.Summary of wolf den and rendezvous sites discovered from 1975 through 1981
occurring within a 50 mile radius of the proposed Susitna Hydroelectric
Project in southcentra1 Alaska.
Site
Miles from upper
water-level
5
15
20
25
30
35
40
50 +
Pack Type Year of
Name -Site -Documented use
Watana Rendezvous 80,81
Watana Den 80,81
Deadman Den 75
Jay Creek Den 78
Stephan Lake Den 76
Tyone Creek Den 79
Susitna Rendezvous 80
Susitna Rendezvous 80
BrushkanaDen 75
Mende1tna Rendezvous 76
Susitna Den 79 &80
Keg Creek Den 75,76 &78
Clearwater Den 76
To1sona Den 80 &Mend.Rend.77
To1sona Rendezvous 80 &Mend.Den 77
Mende1tna Rendezvous 77
Mende1tna Rendezvous 77
Mende1tna Den 76,77
Hogan Hill Den 78
Hogan Hill Den 75
Sinona Den 78
Sinona Den 77
103
because they pertain primarily to the area lying east of Deadman
----'~-----,
and Kosina Creeks.Further study effort is needed to locate po-
tential and existing den sites in areas west of the above-men-
tioned creeks.
Most den sites discovered in GMU-13 were roughly centered within
the observed territory,but several were located near territorial
boundaries.Table 33 summarizes the annual distances between
contiguous wolf dens discovered from 1975 through summer 1981.
Average distances between den sites between years were quite sim-
ilar from 1975 through 1981.Overall,for the years studied,the
average distance between 38 natal dens was 28.1 miles (S.D.=8.4).
This average distance compares favorably with the average dis-
tance of 25 miles computed by Stephenson and Johnson (1973)for
the northcentral Brooks Range,Alaska.
Based upon the characteristics of den and rendezvous sites exa-
mined from 1976 through 1981 and the apparent abundance of such
si tes in the Unit,we do not believe that wolf den sites will
ever become a limiting factor in the foreseeable future.Al-
though an undetermined number of actual and potential wolf den
and rendezvous sites will inundated as a result of the project,
we believe that this impact to be of minor significance relative
to the other identified impacts.
Summer Acti vi ty Patterns
Susi tna Pack
From 1 May through 6 June activi ty patterns of the four adult
members (two adults and two yearlings)of the Susitna pack were
intensively monitored through a combination of ground observa-
tions,continuous 24 hour monitoring of radio signals from a per-
manent ground station,and periodical monitoring from fixedwing
aircraft.
104
~,
-
-
-
"I'"'"Table 33.Average distances between contiguous natal wolf dens from 1975 through
I 1981 in GMU-13 of southcentral Alaska.
!
Year of Name of Continued Distance between Natal den
Use Packs mi.km.
1975 Brushkana -Deadman 18.5 29.8
Keg Greek -Hogan Hill 32.0 51.5
Hogan Hill -Sinona 20.0 32.2
Hogan Hill -Ewan 36.5 58.7
x 26.8 43.0
s 8.9 17.5
1976 Clearwater -Keg Creek 19.0 30.6
Mendeltna -St.Anne's 34.0 54.7
Keg Creek -Deep Lake 21.0 33.8
Mendeltna -Deep Lake 36.0 57.9-27.5 44.2x
s 8.7 14.0
1971 Mendeltna -St.Anne's 34.0 54.7
1978 Keg Creek -Hogan Hill 17.0 27.4
St.Anne's -Toisona 22.0 35.4
Hogan Hill -Sinona 31.0 49.9
Keg Creek -Jay Creek 41.5 66.8-x 27.9 '44.9
s 10.8 17.3
1979 Susitna -Toisona 25.0 40.2
Tyone -Susitna 26.5 42.6
St.Anne's-Tolsona 22.0 35.4
x 24,.5 39.4
s 2.3 3.7
1980 Toisona -Susitna 18.5 29.8
Toisona -St.Anne's 40.0 64.4-29.2 47.1x
s 15.2 24.5
1981 Toisona -St.Anne's 40.0 64.4
Pooled Data x 28.1 45.3
S.D.8.4 13.6
N =19
-
105
Two hundred twenty-seven hours of ground observations were made
-------_._.
from a blind located on a ridge next to the den site.Presence
or absence of radio-collared wolves at the den site during obser-
vation periods,which usually occurred from 0800 hours to 1600
hours,was determined by both scanning with a hand held antenna
(Ballard et ale 1979)from the blind and by direct observation.
Figures 14 through 16 visually depict the various family associa-
tions which were present at the den site according to hand held
antenna locations.Of the three radio-collared wolves which were
radio tracked from the blind,the adult female (#295)was present
at the den site more often than either the adult male (#305)or
the yearling female (#306)(Fig.16).The most likely explana-
tion for wolf 295's disproportionate presence at the den is due
to nursing responsibilities.Ground observation suggested that
parturi tion had occurred on 1 or 2 May and thus the pups were
relying on female #295 for nouri shment.
Figure 15 provides additional support that the alpha female
(#295)occupied the den site more often than other pack members.
These data suggest that although the alpha female spends more
time at the den all adults share in spending time alone at the
den with pups.Figure 15 suggests that on some occasions no
wolves were present at the den.However,the radio signal from
wolf 302 could not be received and thus thi s wolf I s presence
alone at the den may have accounted for the time gaps.The pat-
tern for wolf 302 is expected to be similar to that of the other
yearling (#306).
These data also suggest that the adult male (#305)spent the
least amount of time at the den.Ballard et al.(1981)demon-
strated that during the denning period adult males (or in this
case the alpha males)were usually present when large ungulate
kills were made.It can be inferred since wolf 305 spent the
least time at the den,that much of his activity was associated
with providing food for the pups.This was supported by aerial
observations.
106
-
-
-
~'
""'"I
I
<-I
10
100
.....
'i 90,
I
I
I
r 80
I
w
~70-...
C'
Z 60-~.
CJ
4(
a:...50
Q
Z
~
0 40
a:
(:J
II.
0 30
'#.
20
r
I.,
305 295
WOLF NUMBER
306
Fig.14.Occurrence of three radio-collared wolves at the
Susitna wolf pack den studied from 1 May through 6 June
1980 in GMU 13 of southcentral Alaska.
107
Fig.15.Occurrence of lone adult wolves at the susitna wolf
den from 1 May through 6 June 1980 in GMtJ 13 of
southcentral Alaska.
50
40
w
Q ~
Z ...=0 30
a:<:I
<:I Z-
I&.~
0 (J 20
oC-a:...
10
3
305:295 306 NONE
PRESENT
-
-
WOLF NUMBER
~,
-
'""'"1
40
Q w
Z ~.
=t-O 30
a:<:I
<:I Z-I&.::ac::
0 (J 20
oC-a:
t-
10
Fig.16.Frequency of occurrence of adult wolf associations at
the Susitna wolf den from 1 May through 6 June '1980 in
GMU 13 of southcentral Alaska.
50
305 "
295
295 &305 &
306 306
WOLF NUMBERS
295,305
&306
108
/II""
I,
'1"'"
i
i
'i
I
The den site is the focal point of wolf activity during the pe-
riod of pup rearing.Fig.16 depicts the association of various
pack members while at the den.When more than one adult wolf was
present at the den site,most often all three and probably all
four adult wolves were present.Wolves 305 and 306 were most
often absent from the den site.These data and those presented
in Table 34 suggest that young adults were accompanying the adult
male on hunting forays.The time of day at which these activi-
ties occur is pertinent to potential development activities which
must occur close to den sites.
Continuous monitoring of signals from the permanent tracking
station was not initiated until 19 May 1980 and was continued
through 10 June;Comparison of ground observations with data
collected on the recorder suggests that behavior of individual
wolves was accurately displayed on the tape.However,ground
observations were too infrequent to accurately truth all wolf
activities.Also the design of the tapes does not lend itself to
transfer onto computer.Thus each data point would have to be
i:q.dividua11y transferred onto computer,requiring hundreds of
man-hours.Development of a tape which would allow easy transfer
of data onto computer,would provide an enormous amount of data
concerning wolf activi ty and could easi 1y be related to energy
budgets.Regardless ,the techniques did allow the determination
of the presence or absence of wolves at the den for 24 hour pe-
riods.
The presence or absence of the alpha female (#295)and male
(#306)at the den site from 19 May through 10 June is depicted in
Fig.17.As expected,the female spent more time at the den site
than did the male.These observations were also supported by
ground and aerial observations (Table 34).Male 305 exhibited a
pattern of remaining at the den site during mid-day hours but
away from the den during evening and early morning hours.Ground
observations suggested that wolf 306 exhibited a similar pattern
of den occupancy..Aerial observations indicated that when away
from the den,the pack was hunting.Similar to the pattern de-
109
110
-
Table 34.(cont'd).
'I"""Date Time (Ob.No.)liD Location Activity
!
June 4 11:50 129 295 At den Resting
129 305 At den Resting
129 302 At den Resting
129 306 At den One unidenti-
fied gray
observed trotting
away 600 yd west
June 6 10:21 143 295 At new den Resting
143 305 At new den Resting
143 302 At new den Resting
r 143 306 At new den Resting
June 7 12:15 144 305 Approx.8 mi SW Beaver hunting area
June 10 12:15 143 295 At new den Resting
143 302 At new den Resting
143 306 No signal
June 12 08:46 145 306 Approx.5 mi SW Traveling
r
111
-
-
,.Begin
tape ~--------------------~···············-······..·······..···············..····t -
-_.-.,_.-Tape r&D out
20
--~
~------------------------
~-------------------------------------....................................
.....
---------------------~---------------------~
---_..._-------------------------------------------------•4 ••••••••••••••••__,.
~~------------------_.tape:••••••••·.1
~----_.
....--------
...-------------_..---------..--1
----'-------------------------------~.....................................................................................
---------------------------~~305 killed cadbDu 18 mi.tD SW
2.
..
!C ua
of the
in GMO 13 of
1980.
of the two adult members
located in
through 10 June
Fig.17.Presence or absence
Susitna wol£pack at a den site
southcentral Alaska from 19 May
112
r
.1
:to
•All _lvee lITueot .
~-------------------------------~-~t········..··········..··········•········•..·······_..
~------~-----------------------------------~---_.
•Hale·pup 306 6 111..NW
-----------------,----~-----------~----t
Started
•maving PUliSt------,---------------------------
•All wolves present----------..-.._---------------------,---,---..t·················-···..····..·····.-··'..··..···--.
~------------------------------~--------~----------
..-_...
•302 and 306 1/2 1111.•.tI..---'-------------------------~•305 1d.l1ed beaver 12 1B1.SW ••
---------------------~-------------~...............................................................................................................
.•All wolves present--------------------------------------............................................................................................................_.
2
:I
4....
C
CI
5
IS..
Z
~..
7
•
•
-_··_-------------11·················..··..······1
Bellin tape
1--------.-------.Tape ran out
-------~--~---~-----~----~----
to
•302 at den
--.------------------..-.---------End of recording'•305 12 1B1.SW of den
11 1 ••5 •7 •t 10 11 1 4 ••7 ••10 l'12
I----------A.----------.....,It----------I'.-----------1
TI••Of'DAY
Fig.l?Continued.
113
scribed by Ballard et ale (1981),whenever a large ungulate kill
----- --_.-
had been made wolf 305 was usually present.
Wolf 295 remained at or close to the den site during most of the
study.By late May it appeared she began making short forays
away from the site,leaving yearlings to "babysi til the pups.
Watana Pack -During summer 1981 from 7 May through 23 June the
same procedures used the Susitna den study in 1980 were repeated
wi th the Watana Pack except that all radio-collars had been
color-coded to facilitate individual recognition from the blind.
Originally,we had planned on having eight of the eleven pack
members fitted with activity transmitters.However,because
three of the pack members remained away from the den site and
wolf 323 and 346's radio frequencies could not be received by the
special receiver,only 4 radioed wolves were available for study.
The four included wolf 308 -the adult alpha female,wolf 324 - a
yearling female,325 - a yearling male,345 - a yearling male,
and 346 - a yearling female.
From 7 May through 23 June 1981,a total of 369 hours of ground
observations were made from a blind located just south of the den
site.Ground observations were used to truth the reliability of
the recorder data.At the time this report was prepared ground
observations and recorder data were being transferred onto com-
puter sheets for comparison and analysis with aerial observations
and thus were not available for this report except for wolf 308.
Figure 18 depicts the attendance of wolf 308 (the alph~female)
at the Watana den during late spring and summer 1981.As ex-
pected,wolf 308 rarely left the den for any significant time
period.Ground observations suggested that the pups were born on
11 May.Comparison of den attendance by wolf 308 with wolf 295
of the Susitna Pack during 1980 (Fig.17)suggested that wolf 308
spent more time at the den (1.8 hours/day away from the den in
1981 versus 5.8 hours/day.in 1981 (P<0.05)than had wolf 295.We
suspected that this was the result of pack size in that in 1980
only four adults were present and therefore it was necessary for
114
~,
1
11
.,
1
---------11 11-------------------
Tape begins
----------:---------------,a -1....------
-----------------------1]1--
May 7
8
9
10
11
12
13
14
r 15
16
17
18
19
20
21
22
23
24
25
26 1
lU 27....co 28c
29
30
31
June 1
""'"2
!
)3
4
5
6
7
12 1 2 34.5 6 7 8 9 10 11 12 1 2 3 4 5 6 7 8 9 10 11 12
AM PM --------
Time of Day
=Wolf 308 (adult alpha
female)present.
Figure 18.Presence of wolf 308 (adult alpha female)at the Watana den site
from 7 May through 22 June 1981 in the Susitna River Basin of
southcentral Alaska (solid line indicates presence at denJ.
115
~!
""'"
1
11-----+1 l--
1
"1
11---------.~
11----------11 11-----
1---------------11
1
1
1
1
·1l-----~·1
,++ + + + + + + + + + + +Tape Out + + + + + + + + +-----
-----------------1"1 11-------------
8
9
10
11
12
13
14 -1
15
16
17
18
19
20
21
22 1
June
12 1 2 3 4 5 6 7 8 9 10 11 12 1 2 3 4 5 6 7 8 9 10 11 12
AM PM -------
Time of Day
1 1 =Wolf 308 (adult alpha
female)present.
Figure 18.Continued.
116
r
1
IF""
I
ii
the alpha female to hunt and help provide for pups.However,in
----,_.~,--~
1980 the study was not initiated until 19 May.We believed the
pups had been born on 1 May and thus the activity data from the
two sites were not comparable.We recomputed the time spent away
from the Watana den ~o that it corresponded with the timing se-
quence observed for the Susitna pack;19 days after birth.This
analysis suggested that there were no significant differences
between the two sites for comparable time periods;Watana wolf
308 absent for 4.9 hours/day in 1981 in comparison to Susi tna
wolf 295 in 1980 which was absent 5.8 hours/day,(t=O.54,
P<0.005).Therefore,the attendance of the alpha female at the
den site did not appear to be a function of pack size.
In 1981,pups were first observed outside the den on 7 June.
This observation corresponds with the alpha female beginning to
leave the den site more frequently and for longer time periods,
apparently beginning to participate in hunting forays.
Appendix E summarizes the daily movements,activity and associa-
tion of all eight Watana wolves as observed from aircraft from 6
May through 22 June 1981.Ground and aerial observations of wolf
308 were closely correlated with activity data provided by the
continuous moni toring system.
Daily movements of individual pack members were quite variable.
Yearling female wolves appeared to spend more time and remain
closer to the den site than yearling males.This higher affinity
for the den site by yearling females was also supported by the
dispersal of wolves 310,344 and 345,all of which were males
(two were yearlings).
Den site observations may provide a basis for recommending time
periods when Susi tna personnel should avoid the area.Wolves
were most frequently absent from the den site during the _early
morning and late evening hours,but some pack members were always
present.Therefore,in reality there probably are not time pe-
riods when project activities could occur without causing some
117
disruption and annoyment to the pack.However,
--~_.-
essential,the late evening -early morning hours
disruptive.
!fIIill!rn
if absolutely
~
would be less
~
During 1981 the Watana wolves were subjected to relatively fre-
quent disturbance by helicopters.Not only were our personnel
ferried to and from an area ~mile away from the site,but heli-
copter pilots unaware of their location in relation to the den
site flew directly over the den site on several occasions.Ini-
tially,the wolves were quite frightened by the helicopter and we
feared the den site might be abandoned.However,as the study
proceeded the wolves became relatively tolerant of helicopter
activity and no longer fled the scene when a helicopter approach-
ed.It was,however,always a source of at least minor annoy-
ment.From these observations it appears that denning wolves
eventually get accustomed to helicopter activities.The same is
not true for human di sturbance on the ground.
During the 1980 study it was apparent that the blind was a source
of annoyance to the Susitna pack.Between 4 and 6 June 1980 the
Susi tna pack moved to their first rendezvous site.At the
Tolsona pack den in both 1980 and 1981,a single visit by humans
caused the wolves to move the pups and abandon the site for the
remainder of the year.All three of these observations were the
earliest dates that wolves were observed moving from a den to a
rendezvous site,obviously in response to disturbance.Subse-
quent observations,however,indicated that pups had not suffered
any mortality as a result of this disturbance.From these obser-
vations we infer that if project personnel absolutely had to per-
form activities close to a den site,they could do so as early as
4 June without increasing pup mortality even though the si te
could be prematurely abandoned.For all other routine activities
we recommend that areas within a 1.5 mile radius (Chapman 1976)
of wolf dens be closed to proj ect personnel.
118
.....
-
r
.....
r
POTENTIAL IMPACTS OF SUSITNA HYDROELECTRIC PROJECT ON WOLVES
The most severe impact of the proposed project on wolves would
occur indirectly due to reductions or changes in the density,
distribution,sex-age composition,and/or physical condition of
prey.Reductions in moose or caribou density in the immediate
vicinity of the impoundments would probably cause reductions in
wolf densities for at least six to seven resident wolf packs
which currently occupy the area.Also any di sruption of moose
migrations and/or reductions in migratory moose densities may
also reduce wolf densi ties in areas where migratory moose reside.
Immediately following construction of the impoundments we antici-
pate temporary increases in wolf densities next to impoundment
areas due to the increased availability of moose and caribou
which would be displaced from the reservoirs.In turn,this may
amplify the effects of wolf predation on moose and caribou and
ultimately result in lower densities for all species.Increased
competition between bear and wolf could be expected which would
probably result in additional mortality to each species.
Aside from the indirect affects resulting from reductions in prey
density,the proposed impoundments would directly eliminate wolf
habitat by inundating den sites,rendezvous sites,travel corri-
dors,and feeding areas.Loss of habitat would force wolf packs
to readjust terri tory boundaries with neighboring packs which
probably would result in an undetermined amount of mortality due
to social strife.Lower wolf densities in the vicinity of the
impoundments may also result in lower densities elsewhere.If
populations reach low enough levels,wolves will no longer be
able to disperse from the impoundment area to territories vacated
by hunting,trapping and natural mortali ty.
Increases in human activity in the project area will probably
disrupt and in some cases cause wolves to abandon den and feeding
sites.Early den site abandonment could increase pup mortality.
Increased human activities may result in increased hunting and
119
trapping activities as the occurrence of different packs become
common knowledge to larger numbers of hunters and trappers and as
access into the proj ect area becomes more developed.
PROPOSED PHASE II STUDIES
Completion of Phase I studies documented a number of gaps in our
knowledge of Susitna River Basin wolves and the potential impacts
of the proposed proj ect on them.
First,we believe a continuation of all of the Phase I objectives
is warranted because poor snow conditions in 1980 and 1981 did
not allow us to capture and study packs known to occur in the
area.This is particularly true for the areas west of Kosina and
Deadman Creeks.In December 1981 while this report was in pre-
paration three new wolf packs were captured and radio-collared.
Also further effort should be directed towards determining winter
predation rates for large packs such as the Watana Creek Pack.
Emphasis of Phase II studies should be directed towards the in-
teraction of all predators to their prey.More specifically we
propose that a moose and caribou calf mortality study be initia-
ted to determine the impacts of wolves relative to brown and
black bears on ungulate survival.Efforts to mitigate for lost
moose habitat by manipulating vegetation could fail if ungulate
populations are already limited by predation.Also if predation
is already limiting recruitment into adj acent ungulate popula-
tion,displacement of predators from the impoundment areas could
cause further population declines.We also propose that consid-
eration be given to studying caribou mortality not only in the
main herd,but perhaps more importantly,in a subherd such as the
Butte Creek -Monohan flats herd where predation could be more of
a limiting factor on population growth.
Wolf studies should be expanded to include downstream areas.If
moose migration measures occur downstream an assessment of exist-
ing predator population should be made to determine if mitigation
120
'""",
...,
I
-I
.-
measures for moose would also serve as mitigation for predator
losses.Predator densities appear to be quite low in downstream
areas and thus moose mitigation measures may not result in miti-
gation for predators.
ACKNOWLEDGEMENTS
James Foster,Woodland Park Zoo,participated in and performed
the ground observations during the summer activities study as an
unpaid volunteer.ADF&G biologists Donald Cornelius,Sterling
Miller,Dennis McAllister and Enid Goodwin participated in tag-
ging programs.Special appreciation is expressed to Don
Cornelius for his assistance during the first year of the pro-
ject.
Vern Lofstedt,Kenai Air Service,piloted the helicopter and par-
ticipated in the processing of many of the immobilized animals.
Alfred Lee,Lee's Air Taxi,and Kenneth Bunch,Sportsman's Flying
Service,piloted fixed-wing aircraft both during tagging opera-
tions and during monitoring acti vi ties.The experience and help-
ful cooperation of these individuals contributed greatly to the
success of this project.
We also express appreciation to Grandville Cooley,Watana Camp
Manager,for his cooperation and assistance wi th field studies.
Karl Schneider,ADF&G,provided guidance and support throughout
the project and made a number of helpful suggestions for improv-
ing this report.
121
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131
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APPENDIX A.Copy of manuscript accepted by Journal of Wildlife
--_._-----------
Diseases.
COMPARISON AND ASSESSMENT OF DRUGS USED TO IMMOBILIZE ALASKAN
GRAY WOLVES (Canis lupus)FROM A HELICOPTER
WARREN B.BALLARD 1 and ALBERT W.FRANZMANN z
Abstract:One hundred three Alaskan gray wolves (Canis lupus)
were immobilized from a helicopter in the Nelchina and upper
Susi tna River Basins of southcentral Alaska between March 1977
and May 1981.Sixty-five wolves were immobilized with a mixture
of phencyclidine HCL and promazine HCl (PPjHC1)i 38 wolves were
immobilized with etorphine HC1.Phencyclidine HC1 is no longer
commercially available and an assessment of etorphine HC1 as a
replacement drug was made.Etorphine HC1 dosage of 2.5 mg/wolf
(.063 mg/kg)proved to be a suitable replacement for PP/HC1 for
immobilizing wolves.The advantages and disadvantages of the
drugs are di scus sed.
INTRODUCTION
Phencyclidine HC1 3 with promazine HC 4 (PP/HC1)has been "a widely
used drug combination to immobilize free-ranging gray wolves
(Canis lupus).Z,6 ,7,11 Recently phencyclidine HC1 was removed
from the commercial market,forcing biologists to seek suitable
alternatives.Fuller and Keith 5 reported successful immobiliza-
tion of 12 wolves with etorphine HC1 5 ,9 of which were caught
1 Alaska Department of Fish and Game,P.O.Box 47,Glennallen,
Alaska.
Z Moose Research Center,Alaska Department of Fish and Game,
Soldotna,Alaska.
3 Sernylan R,Bio-ceutic Laboratories,Inc.,St.Joseph,
Missouri.
4 Sparine R,Wyeth Laboratory,Inc.,Philadephia,Pennsylvania.
132
in steel traps.Darting wolves from a helicopter has become a
-~----- ----
common method for immobilizing Alaskan wolves7.and an assessment
of a replacement drug (etorphine HC1)for phencyclidine HC1 was
done.This paper compares responses of 65 wolves immobilized
with PP/HC1 with 38 wolves immobilized with etorphine HC1.All
wolves were darted from a helicopter.
MATERIALS AND METHODS
The study was conducted in the Nelchina and upper Susitna River
Basins of southcentral Alaska from March 1977 to May 1981.De-
scriptions of the area have been reported elsewhere.I,3,B,1 D
Wolves were located for radio-telemetry studies by searching the
edges of streams and lakes for wolf footprints from fixed-wing
aircraft when fresh snow depths generally exceeded 8 em.Foot-
prints were followed until wolves were observed,at which time
their location was relayed by radio to a crew in a nearby heli-
copter.5
Wolves were immobilized using a dart gun 7fired from the heli-
copter.To facilitate shooting,the helicopter was equipped with
a zippered canvas door which replaced the right rear window.
Generally from the time wolves were first spotted from the heli-
copter,1 to 5 minutes elapsed until a wolf was darted,requiring
an average of 3 shots for one hit with an experienced shooter.
Once darted,the wolf was observed at a distance from the heli-
copter until it became immobile.Usually the fixed-wing aircraft
continued to observe remaining pack members so that additional
5 00-99,The Lemon Co.,Inc.,Rockvi lie,Maryland.
6 Bell 206B,Bell Helicopter,Fort Worth,Texas.
7 Cap-Chur,Palmer Chemical Co.,Douglasville,Georgia.
133
~,
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r
ilI,
wolves could be captured.Induction time was measured with a
stopwatch and consisted of the time from injection to when the
animal was immobile enough to permi t handling.
From March 1977 to March 1979 a combination of 100 mg phencycli-
dine HC1 and 50 mg promazine HC1 (PPjHC1)was utilized to immobi-
lize 65 wolves of both sexes and all age groups.Recovery time
for this group was not measured;however,all were subsequently
observed in the wild usually wi thin 24 hours.From April 1980
through May 1981,2to 2.5 mg etorphine was used to immobilize 38
wolves of both sexes and all age classes.On this group once
processing was completed,an equivalent cc dosage of the antago-
nist diprenorphine HC1 Bwas administered either intravenously (IV)
in the radial vein or intramuscularly (IM)in the hip area.Re-
coverytime was measured with a stopwatch from when injection was
complete until the wolf regained mobi;Lity.All statistical com-
pari sons were by t-test.9
RESULTS AND DISCUSSION
Initially 2.0 -2.25 mg etorphineHC1 was tested on seven adult
wolves (x =0.50 mg/kg.Four of these wolves required additional
dosages of 00-99 to become immobile (from 0.4 to 2.0 mg addi-
tional).Average induction time was 17.3 minutes (S.D.=8.2).
Wi th the required additional drug the dosages (x =.064 mg/kg,
S.D.=.024)required for immobilization were considerably larger
than those recommended by Fuller and KeithSfor both trapped and
darted wolves.Subsequently,we began using 2.5 mg etorphine for
all wolves.
There was a significant reduction (P<.OOl)in induction time be-
tween initial injection of 2.5 mg (x =6.1 min)versus 2.0 mg (x
=17.3 min).This suggests that a larger initial dose results
B 00 50-50/The Lemon Co./Inc.Rockville,Maryland.
134
in quicker induction which was similar to that reported for moose
-------_.---
(Alces alces)immobilized with etorphine.4
Induction time for wolves immobilized with 2.5 mg etorphine were
significantly less (P<O.OOl)than those for wolves immobilized
wi th 100 mg Serny1an (Table 1).An average of .063 mg/kg 00-99
was used to successfully immobilize 25 wolves ranging in weight
from 20.9 to 50.9 kg.Numbers of wolves requiring additional
drug for no apparent reason after the first injection was similar
\
between the two drugs (etorphine HC1 - 2 of 30j PPjHC1 - 2 of 65.
Additional drug was required for four wolves (2 with PP/HC1 and 2
wi th etorphine HC1)due to malfunction of the dart.Although
initial durg costs for etorphine HC1 (approximately $82.00/20 m1
versus $11.00/10 m1)are substantially higher than that of
PP/HC1,a reduction of 40 percent induction time significantly
reduced costs per captured animal by reduced helicopter time.
Therefore,overall use of etorphine HC1 was no more expensive
than that of PP /HCl.
An average of .144 mg/kg of the antagonist diprenophine HC1 was
used to reverse immobilization (Table 1.)When administered IV
recovery to full mobility averaged 1.58 minutes.As expected,
there was a significant (P 0.001)increase in recovery time over
when'the antagonist was administered 1M (x =8.4 min).Because
no antagonist was available for PP/HC1,accurate comparison of
recovery times between PPjHC1 and etorphine HC1 was not possible.
However,observations of radio-collared animals immobilized with
PP /HC1 suggested that from 1 to 12 hours was required for full
recovery%.In one case,the prolonged recovery resulted in one
wolf drowning%whi1e no mortalities occurred with those immobi-
lized with etorphine RC1.Because of the quick recovery period,
etorphine HC1 is superior to PP/HC1 because it did not expose
immobile animals to prolonged extreme temperaturesSwhich are
common in habitats occupied by wolves,and it did not leave the
animal susceptible to accidental death or vulnerable to hunting
and trapping morta1i ty.
135
-
"'1
~~~1 =~~~1 ~-~]
f-'
W
0\
Table 1.Comparison of pehncyclidine hydrochloride to etorphine hydrocholoride for immobilizing gray wolves from
helicopter in the Nelchina and upper Susitna River Basins of southcentral Alaska between March 1977
and May 1981.
Dose Induction time Diprenorphine dose Recovery time
Sample mg/km (min.)(2 mg/ml)mg/kg (min.)
Drug size 1/i Range St.dev.it Range St.dev.it Range St.dev.i Range St.dev.
Sernylan
(lOO mg/ml)56 2.57 1.96 -6.12..75 10.0 3-25 5.39 1-10 hours
Sparine
{50 mg/ml
Etorphine 2/(l mg/ml.)25 .063 .049 -.120 .018 6.1 2.5-11 2.37 .144 .098-.282 .047 1.58-.8-3.5 .67
1/Sample size represents data only from animals where accurate timing was available.I/IV only.1M average was 8.4 (N =5).S.D.=4.34.Range 2-212 minutes.
Etorphine has two major disadvantages associated with its use:
- ----_..---
(1)its use requires a narcotics license and (2)it is extremely
toxic to humans.Al though there is no substitute for extreme
caution when handling etorphine;a human antidote naloxone ECl'
should be available at all times.
In conclusion;results of this comparison suggest that etorphine
is suitable and in some cases more advantageous than PPjHCl for
immobilizing wolves from a helicopter.We recommend that 2.5 mg
etorphine ECI be used for immobilizing wolves to be darted from
helicopter and that an equivalent ml dosage (2 mg/ml)of the an-
tagonist diprenorphine ECI be administered following processing.
The dosage of etorphine RCI was much higher (0.063 mg/kg versus
0.042 mg/kg)than previously reported 5 •
ACKNOWLEDGEMENTS
Appreciation is expressed to K.Schneider and G.Bos,both Alaska
Department of Fi sh and Game CADFScG)for reviewing the manuscript
and R.Tobey (ADFScG)for assistance in the field.This study was
supported in part by Alaska Federai Aid to Wildlife Restoration
Proj ects W-17-R with additional support provided by the Alaska
Power Authori ty.
,Narcan;Endo Laboratories;Inc.Garden Ci ty;New York.
137
-
-
LITERATURE CITED
Ballard,W.B.1981.Gray wolf-brown bear relationships in
the Nelchina Basin of southcentral Alaska.F.Harring-r
1.
ton and P.C.Paquet,Co.Eds.
Symp.Portland,OR.
Proc.Portland Wolf
2.R.o.Stephenson,and T.H.Spraker.1981 Nelchina
Basin wolf studies .Alaska Dept
Fish and Game.Final
P-R Rept.Juneau.201pp.
3.Bishop,R.H.and R.a.Rausch.1974.
fluctuations in Alaska,1950-1972.
593.
Moose population
Nat.Can.101:559-
4.Franzmann,A.W.and P.D.Arneson.1974.Immobilization
of Alaskan moose.J.Zoo Anim.Med.5(2):26-32.
5.Fuller,T.K.and L.B.Keith.1981.
wolves in winter with etorphine.
45(1):271-273.
Immobilization of
J.Wi Idl .Manage.
L.D.1974.Current techniques in the study of elu-
sive wilderness carnivores.Proc.IXth Intl.Cong.
Game Biol.11:315-322.
If'-6.Mech,
il
il
'I"'"
:1
7.Seal,U.S.,A.W.Erickson,and J.
immobilization of the carnivora.
10:157-170.
G.Mayo.1970.Drug
Int.Zoo.Yearbook.
-
8.Skoog,R.o.1968.Ecology of caribou (Rangifer tarandus
granti)in Alaska.Ph.D.Thesis,Univ.of California,
Berkeley,CA.699 pp.
138
9.Snedecor,
ods.
G.W.and W.G.Cochran.1973.
Iowa State Univ.Press.593pp.
statistical meth-
Moose movements and
River near Devils
10.Taylor,K.P.and W.B.Ballard.1979.
habitat use along with Susitna
Canyon.Proc.N.Am.Moose Conf.
186.
Workshop.15:169-
11.Van Ba11enberghe,V.,A.W.Erickson,and D.Byman.1975.
Ecology of the timber wolf in northeastern Minnesota.
Wildl.Monogr.43:1-43.
139
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i
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I
I
APPENDIX B.Abstract of report summarizing wolf research
activi ties in Game Management Unit 13 from
1975 through June 1980.
ALASKA DEPARTMENT OF FISH AND GAME
JUNEAU,ALASKA
STATE OF ALASKA
Jay S.Hammond,Governor
DEPARTMENT OF FISH AND GAME
Ronald O.Skoog,Commissioner
DIVISION OF GAME
Ronald J.Somerville,Director
Donald McKnight,Research Chief
NELCHINA BASIN WOLF STUDIES
By
Warren B.Ballard
Robert o.Stephenson
and
Ted H.Spraker
Volume III
Final Project Report
Federal Aid in Wildlife Restoration
Project,W-17-9 and W-17-10,Jobs l4.8R,14.9R and 14.10R
wi th Addi tional Support from the Alaska Power Authori ty
140
JOB PROGRESS REPORT (RESEARCH)
Alaska
Warren Ballard,Robert Stephenson,
Ted Spraker,Sterling Eide,James Foster,
Albert Franzmann,Art Flynn,Dan Holleman,
SuzAnne Miller and John Sch10tthauer
r
r
State:
Cooperators:
Project Nos.:W-17-9,W-17-10
and W-21-2
Project Title:Big Game
Investigations
Job No.:
Job No.:
Job No.:
14.8R
14.9R
14.10R
Job Title:
Job Title:
Job Title:
Wolf Populations
and Movements
in Relation
to Those of
Prey Species
Wolf Food Habits
Impact of Wolf
Predation Upon
Ungulate Populations
Period Covered:July 1,1975 to June 30,1980
SUMMARY
.....
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 (Ne1china Basin)of
southcentra1 Alaska.Cost and problems associated with heli-
copter darting were di scussed.
The 103 radio-collared wolves were individually located on 3,525
separate occasions resulting in 6,927 wolf sightings.Pack and
individual wolf histories were presented and discussed.
Wolf territories were for the most part non-overlapping.Over-
laps which did occur were seasonal in nature or due to the method
territories were plotted.There appeared to be changes in ter-
ritory boundaries from year to year.Territory sizes for 14 in-
141
-._._~----------------------_._-~-----
tensively studied packs ranged from 268 to 864 mi 2,averaging
537 mi 2 •Territory size appeared to increase for larger packs
and for those packs in areas of low moose densi ty.
Wolf den and rendezvous site usage was described.The earliest
that radio-collared wolves were observed at a natal den was
13 April.Wolves began visiting den sites in late April and
early May.Parturition appeared to occur throughout the month of
May.Natal dens were abandoned between 4 June and 1 August.
Pups were observed traveling with adults between 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 bear.Moose of varying ages comprised 72 percent of the
observed kills.Calf and short yearling moose comprised 20 per-
cent of the tot~l 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
a 5 year period.Overall,calf moose was the most frequent iden-
tified food item (44%).Percent occurrence of various prey items
in wolf scats were generally related to prey abundance.Occur-
rence 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 sug-
gested that wolves in GMU 13 were preying upon from 434 to 1,013
moose calves annually from mid-May through mid-July.
One hundred twenty five moose and 25 caribou kills were examined
in situ to determine both cause of death and age and physical
142
-
~.,
--
."1"""
I
II
r
1~
F""
t
I
-
"..,.
condi tion of prey taken by wolves.One instance of surplus
killing of caribou by wolves was reported.Percent fat of calf
and short yearling moose killed by wolves was significantly
higher than those of calves dying from both accidental causes and
winter kill.We concluded that wolves were preying upon rela-
ti vely heal thy 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.Over-
all,ages of adult moose killed by wolves were significantly
(P 0.05)different from·tagged moose,winterki lIed moose,and
moose dying from accidental causes and winter ki 11.
Age and condition of wolf-killed adult moose were compared with
those of moose tagged during the same year predation occurred.
We concluded that during severe winters wolves preyed upon adult
moose in proportion to their ratios in the moose population while
during average or mild winters older adult moose were being
preyed upon.
Marrow fat percent of wolf-killed adults was significantly
(P<O.OS)higher than moose dying from winter kill but not signif-
icantly (P>O.OS)different from those dying of accidental causes.
We concluded that wolves were preying upon relatively healthy
adu1 t moose during winter.
Qverall,17 wolf packs averaged a ki 11 every 4.9 days with a
range of 3.1 to 12.7 days per kill.Differences between packs
and problems associated wi th methods of calculating predation
rates were di scussed.
During the winters of 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 kil1/3.6 days for a pack of nine wolves.
Large wolf packs generally appeared to have a higher kill rate
than smaller wolf packs.
143
--------------------------
During the summers of 1977 and 1978 acti vi ty patterns of two
---~,-------~--
denning wolf packs were studied and are discussed.It was deter-
mined that adult males (presumed to be alpha males)were nearly
always present when ungulate ki lIs were made.
Wolf densities in GMU 13,excluding the wolf removal area,varied
from 1 wolf/37.6 mi Z in fall 1975 to 1 wolf/121.7 mi Z 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 were presented and discussed.Rates
of harvests from individual radio-collared wolf packs were exam-
ined in relation to productivity and ability 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
recommended that a hunting and trapping bag limit of seven wolves
be established in GMU 13.It was also suggested that a post
hunting-trapping wolf density of 1 wolf/100 miZmight be suitable
to keep wolf predation on moose to a minimum but yet maintain a
reasonable wolf 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 mor-
talities.
During this study at least 26 radio-collared wolves were known to
have dispersed from their original pack area.Sixty-eight per-
144
~,
-
-
cent of the di spersals were males.Approximate average ages of
dispersed males was 35 months,while females averaged 37 months.
Dispersal was most prevalent during months of April through June.
Average distance dispersed was at least 67.7 miles.The longest
documented movement was 460 miles,constituting a record movement
for this species.
The effects of wolf predation on moose calf survival was studied
in two areas of GMU 13.In one area,referred to as the Susi tna
River Study Area,wolf densities were lowered by Department per-
sonneL In the other area of GMU 13,(remainder of the Uni t
generally east of Talkeetna Mountains)wolves were intensively
studied to enumerate population densi ties and food habi ts.
From January 1976 through July 1978 a total of 60 wolves were
killed by Department personnel in an effort to test the hypoth-
esis that lowered wolf densities would improve moose calf sur-
vi val.Wolf Densities in the Susi tna River study area were
estimated at 1 wolf/98 mi 2 0 f habitat in spring 1975 to
1 wolf/232 mi of habitat in spring 1978.By spring 1980 wolf
densi ties had increased to wi thin 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 control had not been
conducted.Statistical analyses revealed no significant (P 0.05)
differences in either calf-cow ratios,moose observed per hour of
survey,nor in ratios of harvested moose.Had wolf control in-
creased moose calf survival we would have anticipated some sig-
nificant differences in these ratios.Results of wolf food
habits,moose calf mortality,and bear food habits studies in-
dicated that the rates of predation on moose calves by wolves
were far less than by brown bears.Thi s tended to explain the
lack of response by the moose population to reductions in wolf
densities.Results of the bear transplant on moose survival were
compared and di scu s sed with thi s study.
145
-
-
APPENDIX C.
Copy of paper presented at 17th North American Moose Conference
and Workshop held in Thunder Bay,Ontario.
USE OF MANDIBLE VERSUS LONGBONE TO EVALUATE PERCENT MARROW
FAT IN MOOSE AND CARIBOU
W.B.Ballard,C.L.Gardner,and J.H.Westlund
Alaska Department of Fish and Game
P.O.Box 47,Glennallen,AK 99588
and
S.M.Miller
Alaska Department of Fish and Game
333 Raspberry Road,Anchorage,AK 99502
146
Abstract:During winters 1977 --through-1980 the mandible and a
longbone were collected from moose (Alces alces)and caribou
(Rangifer tarandus)kills found while conducting a wolf (Canis
lupus)moose relationships study in southcentral Alaska.Percent
marrow fat for the paired samples was significantly correlated,
suggesting that mandibles could be used in lieu of longbones for
marrow fat analyses.Resul ts of the study were compared with
those obtained for Ontario moose and were combined for analysis.
Percent fat for the paired bones was significantly correlated for
both calf and adult moosej however,the slopes and intercepts for
the two age classes were different,suggesting differences in fat
mobilization by age class.
Marrow fat of longbones had been widely used as an index of phys-
ical condition of ungulates in North America.Prior to 1970 pro-
cedures for determining marrow fat content consi sted of either
crude visual estimates based upon marrow color and consistency,
or extraction procedures which were relatively expensive and time
consuming.Development of Neiland's (1970)dry weight for deter-
mining percent marrow fat in caribou allowed marrow fat content
to be quantified with relative ease and a relatively low cost per
sample.
Since 1970,Neiland's (1970)method of determining marrow fat
content has been widely used on a number of ungulate species for
assessing physical status.This type of information is of par-
ticular interest to students of predator-prey relationships be-
cause it allows inferences to be drawn about the physical con-
di tion of prey selected by predators.For compari son,samples
from nonpredator killed ungulates are needed to determine con-
dition of predator kills relative to the condition of other
members of the population.
The most widely used bone for determining percent marrow fat of
ungulates has been the femur (Cheatum 1949),although other long-
bones have been widely used also (Peterson 1977).Percent fat in
the mandibular cavity has also received some attention as an in-
dicator of physical condition (Baker and Lueth 1966,Purol et al.
1977 and Snider 1980).
147
While conducting a wolf-moose relationships study in Game Manage-
~--- ------_.-
ment Unit 13 of southcentral Alaska,we attempted to collect
longbones from moose and caribou dead from all sources of mor-
tali ty.Al though we strove for collection of femurs,we often
had to settle for metatarsals or metacarpals,and in many other
cases no bone was collected at all.Reasons for this varied de-
pending upon both the cause of mortality and the time available
for specimen collection.On both predator-and winter-killed
ungulates,which were only partially consumed,the flesh was
frozen and extraction of the femur was often time consuming and
expensive,particularly when kills were visited via helicopter.
On heavily consumed predator kills,often the ends of longbones
had been chewed and the marrow either eaten or exposed to the air
rendering the sample useless.In these latter cases,no speci-
mens were collected.On several predator kills the only remain-
ing intact bones suitable for marrow analysis were the mandibles.
Similar types of problems occurred with collection of samples of
road kills.Because of these problems and the presence of mand-
ibles at many heavily consumed predator kills,it appeared de-
sirable to determine if a relationship existed between percent
marrow fat estimated from longbones compared to that estimated
from mandibles.Since mandibles are relatively easy to extract
and are often collected routinely for aging purposes,establish-
ment of a fat relationship between the two bones would result in
a considerably larger sample size of condition data.The purpose
of this paper is to compare the percent marrow fat of mandibles
to that of longbones for moose and caribou killed primarily by
predators.
STUDY AREA
The study was conducted in Game Management Unit 13 of south-
central Alaska.Detai led descriptions of vege"tation,topography,
weather patterns,etc.,have been provided by Skoog (1968),
Rausch (1969),Bishop and Rausch (1974)and Ballard (1981).
148
..,.,
-
r
r
i
r
I
r""
,I
!
METHODS
During winters 1977 through 1980,58 paired mandible and longbone
samples were obtained from moose and caribou kills.Moose sam-
ples were compri sed of 21 calves and 24 adults of both sexes,
while the 13 caribou samples were adults of both sex.Ages of
moose were determined by incisor eruption and cementum annuli
counts according to methods described by Sergeant and Pim10tt
(1959).Caribou were aged on the basis of tooth wear described
by Skoog (1968).
Samples were collected on an opportunistic basis but most kills
were detected while making flights to monitor radio-collared wolf
packs.Causes of death for the paired samples were as follows:
for calf moose--8 wolf kills,7 winter kills and 6 road or acci-
dental kills;for adult moose--15 wolf kills,8 road or acci-
dental kills,and 1 from unknown causes;and for adult caribou-
12 wolf kills and 1 from unknown causes.
Procedures for determining percent fat of longbones were iden-
tical to those of Neiland (1970).Mandible marrow was extracted
by cutting a 10 cm longitudinal section of bone from the labial
side of the mandible beginning at the 2nd or 3rd premolar.The
section was cut with a bone saw and the resulting bone dust was
scraped from the marrow with a spatula.Later,we simplified
this process by ventrally splitting the left or right ramus with
a chisel and then extracting the entire section o'f marrow wi tha
spatula.This modified procedure also eliminated the need for
scraping off bone dust fragments.The remainder of the procedure
was identical to -that for the longbone,described by Neiland
(1970).
149
RESULTS AND DISCUSSION
Paired samples for both calf and adult moose were compared with
standard least squares regression techniques (Snedecor and
Cochran 1973).The best fit was by linear regression (Fig.I,
r=.92,P<0.05-)i however,the data appeared clumped according to
age class (calf versus adult).Analysi s of covariance for calf
and adult moose indicated that the variances were significantly
different (F=4.11,P<O.OOl)and,thus,comparison of slope and
intercept between the two age classes was not possible.We sub-
j ected each age class to polynomial regression techniques and
determined that the percent marrow fat relationship for adult
moose could be expressed as a 3rd order polynomial.The rela-
tionship was also significantly related linearly and thus we
chose it for adults because it allowed additional statistical
tests to be performed.
Figures 2 and 3 depict the relationship between percent fat for
longbones and mandibles for calf and adult moose separately.
Percent marrow fat in the two bones was significantly correlated
for both age classes (calves r=.88,P<0.05;adults r=.78,P<0.05)
suggesting that longbone fat could be estimated from mandible fat
for each age class.However,there was considerably more vari-
ation in the relationship for calves (mean square [ms]=127.8)
than for adults (ms=26.7).This may have been the result of
sample size since all of the adult moose were above 65%fat which
would have placed them in a relatively high condition class based
upon criteria established by Greer (1968)and Franzmann and
Arneson (1976).
Snider (1980)compared percent marrow fat for femurs and mand-
ibles for 29 moose from Ontario.He,like us,concluded that the
two variables were significantly correlated.He combined calf
150
-
"""
-
~~~-1
Jl
adults
Jl
I:•
o =calves
o
0.92
I:
r=
y=2.24 +l.16X
o
x
o 0
x
o
o
o
4'/....-.....;~,,)......-/
1:1:I:...
1.1/}"./...
...X /''''x_.o•••••..,,/•..../....JI.-...
.)11 •••../"x·...<'"•••..,....
""/J'~..,"~
.........,.........ft
X 0 ./"••••/'"0
..,-/.'..;
d '······/I""../...-
'.
6
,..-0
••••,iJ-"-'"//7'··......./'..0
...."....'••••r"
,,/,....~
/"......".
./,I.~,~#....••••.,...
./'"..-.../,.'~.,
.w'-.'./..
/'..1.1
,P~/••••••. 0
..'............""....
0/"0100
80
10
100
90
~70
~
~60.~50
~o~40
9
'30~
~20
~
Pol
I-'
U1
I-'
0 10 20 30 40 50 60 70 80 9U 100
PERCENT MANDIBLE MARROW FAT,
Figure L R,e1ationship between percent marrow fat for mandibles and longbones from calf and adult moose in
the Ne1china Basin,Alaska.
1/
80.I I I
100.I I I I I 7/I
~~
!6°,'1 r--1**~
......~
In @N~
...:I
~I *40.~~I /*I.I y=;"0.10+1 \07XPoi
*,I r=O.S'S
2e.
I /1
I )I*.~**
i·
e...40.0.20.60.80 ..100.
PERCENT MANDIBliE MARROW FAT.
Figure 2.Relationship between percent marrow fat of mandibles and longbones for calf moose
~,I in the Nelchina Basin.Alaska.J J J'I .1 I J
•
]1 J J )I .J J 1 1
100.80.60.40.20 •..
0·,
0.
20'.
80.
.
..,.,..""
,.'*
,,/-
,
.,/
y~.54.75 +O.43X
r=O.78
..,
,
.
•
100.
ell·;;:
~60.
~
~.
a
j:Q
0·~40.
~
t
~
~
~
.P-4
"~~
J--'
L1'1
Vol
PERCENT MANDIBLE MARROW FAT
Figure 3.Relationship between percent marrow fat 9£mandibles and longbones for adult moose
in the Nelchina Basin,Alaska.•
moose (n=6)with adults (n=22)and determined that the percent
fat relationship between the two bones was best expressed as a
third degree orthogonal polynomial (Fig.4)where Y=mandible fat
and X=femur fat.His data were subject to an orthogonal regres-
sion.For comparison,we subjected Snider's data to the same
analyses performed on Nelchina Basin moose and determined that
his data also exhibited a significant linear relationship (r=.87,
P<0.05)[Fig.5].In contrast to Nelchina data,however,the
variances between age classes were equal (F=.87,P>0.05)and
there were no significant differences between slopes (F=.87,
P>O.OS)or intercepts (F=.32,P>O.OS).Reasons for the dif-
ferences in homogeneity of variances between the two studies are
unknown but could have been related to a combination of both
studies,or differences in fat deposition and mobilization
between the study moose populations.
Because of small samples sizes in each of the studies and because
samples in the Nelchina study were collected primarily during
winter for predator kills while those in Ontario were collected
primarily in October or June mainly from road-killed moose,we
combined samples in an effort to better describe the relationship
between longbone and mandible fat (Table 1).The analysis as-
sumed that there were no differences in fat mobilization between
the two populations.Variances between calf and adult moose in
these clumped data were not significantly different (P>0.05)sug-
gesting that fat mobilization in the two bones was different for
the two age classes.The relationship between longbone and mand-
ible marrow fat for each age class was best described by linear
regression (Fig.6 and 7).However,because only three adult
moose had longbone fat values of less than 60%,the relationship
between the two bones at lower fat levels warrants further in-
vestigation.
Similar to samples from Nelchina adult moose,bone marrow fat
from mandibles and longbones of adult caribou were also signifi-
cantly correlated (Fig.8,r=.90,P<0.05)suggesting that
154
-
.....
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, I
•.
I..'
,.
0°•
.
.-/a .,"
'0a Q
;
a
CJa ,
C MALES
o
o FEMALES
.2,3
V =-32·13 at 4.4 X +.08X +.0005X
Q
Q
:.,
",
"
:~..
..
,..80
h'~u.;
:;
a
00
~60
:a
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::Jm
o
'~40
:Eo,
...
Z
UJ
U
0:
UJ
a.20
t-'
V1
V1
,•
:PERCENT FEMUR MARROW FAT •
Relationsn1p between percent marrow fat of 'fem~r9 and mandibles for calf and adult moose
in Ontario (from Snider 1980).
.,
~
Figure 4.
:20 40 60 80,100
'I
"...",0.;,/I I Ii.I
0.20.40.60.80.100.
100.I i I I .<.I 7 I
..80.I I I I .~7'I T I
;-
~,.
.1 I
~60.
~
I I I I <.A I I .'
......
IJl
0\
~40.I0 I 7 -,----I I ....j:Q
(!)
~H I I /...1.*H I,Y=4.87 +l:'04Xf;j
u . IIX:1/I I r=O.87 .~
~80.
PERCENT MANDIBLE MARROW FAT
Figure 5.Relationship between percent marrow fat 6f mandibles and longbones for adult and
,I
calf moose in Ontario (data from Snider 1980)~
JI )j t J J J ,~J ~
~.~~~""1 ~1 -----,-·--1
Table 1.Analysis of covariance of percent marrow fat estimated from mandibles
and longbones of calf and adult moose from southcentral Alaska and
Ontar~o.!/
Source
within
Calves
Adults
Pooled Within
Differences Between Slopes
within and Between
Between Adjusted Means
d.f.S.S.M.S.
25 3084.36 117.57
58 6094.55 105.08
83 9178.91 110.59
84 10221.90 121.69
1 1042.99 1042.99
85 30743.212 361.68
1 20521.31 20521.31
Significance Tests
( 1 )aeterscedasticity
F;117.57/105.08 ;1.12 F (25 58).25 ;
1.12 1.23 accept Ho:2 '2c;a
1.23
( 2 )Difference in Slopes
F;1042.99/110.59 ;9.43 F (1 83)·005 ;
9,43 8.30 rejects Ho:BC ;'B a
8.30
(3)Difference Between Intercepts
F;20521.31/121.69 ;168.6 F (1 84).001 =11.8
168.6 11.8 reject Ho:c =A '
!/Ontario data from Snider (1980).
100.I I ,I <I 7 I
80.
I I I
**
"60 _I
1
,~/1*;::
!I *f..:l
Z 40.0
I:Q
t!)
/*I I y=-2.16 +1.15Xz·0
~*E-i I ·1 r=0.91
z
rLl
U~~20.
I /1
I 'I'.b~*'*,. .
...
.,'I
0./80.40 •.80.100.0.60.
PERCENT MANDIBLE MARROW FAT
Figure 6.Relationship between percent marrow fat of mandibles and longbones for calf moose
in Ontario (data from Snider 1980)~nd the ,Nelc:hina Basin ..Alaska.
.J J ]~
I 1 J B I I t 1 I .~J I !I I
·l
,.
0.I I I .I I I
0.20..40.60.80.100.
""PERCENT MANDIBLE MARROW FAT
Figure 7.Relationship between percent marrow fat of mandibles and lo~8bonesfor adult moose
in Ontario (data from Snider 1980)and"the Ne1China Basin,Alaska.·
•
x
X
r=0.90
y=0.95 +1.05X
"../
X ~/.,../..'x·_.-.,...
Xx ./,......-•••••..~
-"...."........................
./.~-a.···~.,.-
..•/.--;:;."~'::'/.......
....••••••••"x ,.'
x.'..'••••••..--."'~.-.....-,..,--.-,........,.~.
,..•'X
100
90
,80
~70~
~~60
~
~50
z
0
~40z
0
t-l
~30
u~20
~
10
10 20 30 40 50 60 70 80 90 100
PERCENT MANDIBLE MARROW FAT
Figure 8.Relationship of percent marrow fat for mandibles and longbonesfor adult caribou in the Ne1china,Basin,
,
Alaska.
.
•
J 1 J J J I I )I .~.~,J J J J .1 I)1
r!
r
I
r
-
mandibles might be useful for estimating longbone fat in caribou.
However,because sample sizes were extremely small and no samples
of caribou in poor condition were collected,this relationship
should be viewed with caution.Also,since no calf caribou were
\
examined it is unknown whether a correlation exists in this age
class as well.
Peterson (In Press)recently compared marrow fat levels between
several longbones of individual moose and determined that fat
mobilization appeared to have proceeded more quickly in proximal
than in distallongbones.If correct,this may partially explain
some of the variabi li ty between longbones (femurs,metatarsals,
and metacarpals)and mandibles found in thi s study.Even with
this variation,however,mandibles appear useful for determining
the percent marrow fat in longbones and consequently appear use-
ful as an indicator of condition.Although results of this study
suggest a positive relationship exists between marrow fat mobili-
zation in mandibles and longbones,we suggest that biologists
collect paired samples from ungulate kills in other populations
to determine if relationships are similar.If this relationship
is confirmed,then biologists should consider using the mandible
in lieu of longbones for marrow fat analyses.Use of mandibles
will allow biologists to greatly increase sample sizes for marrow
fat analysis with minimal effort at relatively small additional
costs.
ACKNOWLEDGEMENTS
Appreciation is expressed to Charles Lucier and Enid Goodwin,
both Alaska Department of Fish and Game (ADF&G)employees,for
processing marrow samples.Donald McKnight,Karl Schneider,and
Karen Wiley (all ADF&G)reviewed the manuscript and made many
helpful suggestions.We also thank Li ta Lewi s for typing the
manuscript on her personal time.The study was funded in part by
Federal Aid in Wi ldlife Restoration Proj ect W-17-R with addi-
tional support provided from the Alaska Power Authori ty.
160
REFERENCES
Baker l F.M.I and F.X.Leuth.1966.Mandibular cavity tissue
as a possible indicator of condition in deer.Proe.Am.
Conf.S.E.Game and Fish Comme 20:69-74
Ballard,W.B.1981.Gray wolf-brown bear
Nelchina Basin of southcentral Alaska.
P.C.Paquet l Co.Eds.Proc.Portland
Oregon:In Press.
relationships in the
F.H.Harrington and
Wolf Symp.Port1and l
BishoPI R.H.,and R.A.Rausch.1974.Moose population fluctu-
ations in Alaska l 1950-1972.Nat.Can.I 101:559-593.
Cheatum,E.L.1949.Bone marrow as an index of malnutrition in
deer.N.Y.state Conser.3:19-22.
Franzmann l A.W"I and P.D.Arneson.1976.Marrow fat in
Alaskan moose femures in relation to mortality factors.J.
Wi1dl.Manage.40(2):336-339.
,~
Greer l K.R.1968.A
of elk (wapiti)
32(4):747-751.
compression method indicates fat content
femur marrows.J.Wildl.Manage.
Nei1and l K.A.1970.Weight of dried marrow as indicator of fat
in caribou femurs.J.Wildl.Manage.34(4):90,4-907.
Peterson l R.O.1977.Wolf ecology and prey relationships on
Isle Royale.Nat1.Park Servo Sci.Monogr.Ser.11.
210pp.
In Press.Aspects of bone marrow fat determination in
moose.J.Wi 1d1.Manage.
161
fI!lIi!P,
Purol,D.A.,J.N.Stuht,and G.E.Burgoyne,Jr.1977.Mand-
ibular cavity tissue fat as an indicator of spring physical
F"'"condi tion for female white-tailed deer in Michigan.
Michigan Dept.of Nat.Resources,Wi'ldl.Div.,Rept.No.
2792.
Rausch,R.A.1969.A summary of wolf studies in southcentral--Alaska,1957-1968.Trans.N.Am.Wildl.and Nat.Resour.
Conf.,34:117-131.
Sergeant,D.E.,and D.H.Pimlott.1959.Age determination in
moose from sectioned incisor teeth.J.Wildl.Manage.
23 (3):315-321.
granti)in Alaska.
Berkeley,California.
Skoog,R.o.1968.Ecology
PhD.
of caribou
Thesis,Univ.
(Rangifer tarandus
of California,
1973.Statistical methods.
.....
Snedecor,G.W.,and W.G.Cochran.
The Iowa St.Univ.Press.
593pp .
Ames,Iowa.Sixth Ed.
_Snider,J.B.1980.An evaluation of mandibular fat as an in-
dicator of condition in moose.Proc.N.Am.Moose Conf.
Workshop 16:37-50.
"...
F'
i
\
162
I""",
,.....
!
.....
-
APPENDIX D.
ALASKA DEPARTMENT OF FISH AND GAME
JUNEAU,ALASKA
STATE OF ALASKA
Jay S.Hammond,Governor
DEPARTMENT OF FISH AND GAME
Ronald o.Skoog,Commissioner
DIVISION ON GAME
Ronald J.Somerville,Director
Donald McKni,ght,Research Chief
NELCHINA YEARLING MOOSE MORTALITY STUDY
by
Warren B.Ballard
Craig L.Gardner
and
Sterling D.Miller
Volume II
Final Report
Federal Aid in Wildlife Restoration
Projects W-17-11 and W-21-1,Job 1.27R
wi th Addi tional Support from the Alaska
Power Authority .
163
I"F"
II
I
State:
Cooperators:
JOB PROGRESS REPORT (RESEARCH)
Alaska
Warren B.Ballard,Craig L.Gardner,
Sterling Miller,John Westlund,and
Dennis McAllister.
Project Title:Big Game Investigations
T
Project No:
Job No.:
W-21-11 &:
W-21-1
1.27R Job Title:Nelchina Yearling Moose
Mortality Study
r
r
Period Covered:March 1,1980 to June 30,1981.
SUMMARY
Causes and rates of calf (0 to 6 months old),short yearling (7
to 12 months old),and yearling (13 to 24 months old)moose mor-
tali ty were studied in Game Management Unit 13 from late March
1979 through June 1981.The status as of 1 July 1981 of 64 moose
captured as short yearlings in 1979 was as follows:22 had lost
their radio-collars due to inadequate collar design;8 had un-
known status probably due to either dispersal or radio failure;
17 had died,primarily from winter-kill (59%)brown bear pre-
dation (18%);and 17 were still alive.Annual mortality rates
for the 1978 cohort were 76-80%the first year and 5%the second
year.
An additional 34 calf moose were captured and radio-collared in
November 1979 to aid in assessing first year survival of moose
calves following a removal of 48 brown bears (Ursus arctos)(see
Ballard et a1.1980).From capture to 1 June 1980 6.1%of the
moose died due to winter kill.No other mortality of radio-col-
lared calves was observed.Second year mortality was 4%,which
was attributable to unknown causes.The low rate of mortality
prior to 1981 was attributed to mild winter conditions and low
predator densities.
164
During fall 1980 both a standard moose sex-age composition survey
----------
and an intensive quadrat sampling technique were used in the bear
transplant area to both compare sex-age data acquired from the
two survey methods and to obtain estimates of moose density.
Resul ts of the comparisons are briefly described.
Census and composition data collected in 1980 were used to adjust
sex-age composition data collected in fall 1979.Based upon this
readjustment it was calculated that the fall 1979 calf:cow
ratio following the transplant of brown bear was 73.Corrected
1979 composition data were used to calculate mortality from birth
to 1 November 1979 following the bear transplant.The total num-
ber of calves that died from all mortality factors in 1979 was
estimated at 9%.In comparison,studies conducted in 1977 and
1978 revealed that mortality during the same time period was 55%;
80%of which was attributable to predation by brown bears.Ap-
proximate rates of first year moose mortality under varying rates
of wolf and bear predation and winter kill are presented and dis-
cussed.This analysis suggested that the largest increases in
first year survival occurred when brown bear densities were tem-
porarily reduced.It was tentatively concluded that the moose
population was not being limited by deteriorating range condi-
tions and that predation,primarily by brown bear,was preventing
the moose population from increasing.
BACKGROUND
Studies of wolf (Canis lupus)food habits in the Nelchina and
Susitna River Basins of southcentral Alaska (Game Management Unit
13)from 1975 through 1980 suggested that wolves were preying
upon calf and yearling moose disproportionately to their presence
in the moose population from January through July of each year
(Ballard et al.1981).Consequently this study was initiated to
determine the importance of wolf predation to yearling moose sur-
vival.First year results of this study were reported by Ballard
and Gardner (1980).
165
-
-
During the second year of study additional calf moose were cap-
tured and radio-collared to assess the causes and extent of year-
ling moose mortality in an area of reduced brown bear (Ursus
T arctos)density.Background for thi s portion of the study was
,I provided by Ballard et al.(1980,1981).
OBJECTIVES
To determine the extent and causes of yearling moose mortality in
the Nelchina and Susi tna River Basins of Southcentral Alaska.
PROCEDURES
r
r
Yearling moose mortality was studied in two areas of GMU-13
(Fig.1).Short yearling moose (7 to 12 months of age)were cap-
tured in Area 1 during March and April 1979,while calf moose (0
to 6 months of age)were captured in Area 2 during late November
1979.An additional 17 short yearling moose were captured in
March 1981 as part of Susitna Dam Studies but this group will be
discussed in the final study report.Topography,weather,
geology and vegetation of this area has been presented elsewhere
(Skoog 1968,Ballard and Taylor 1980,Ballard 1981).
Capture,processing,and radio-monitoring methods were described
by Ballard and Gardner (1980)and Ballard et ale (1979).Proce-
dures used on moose captured in the bear removal area in November
1979 were identical to those used earlier with the following ex-
ceptions:(1)5 cc (1 mgjml)of M-99 was used to immobi lize
calves,(2)radio-collars··were constructed of butyl rubber with
2-3 inches of foam rubber lining the inner circumference to
permi t growth,(3)no blood samples were taken.
Annual mortality rates were calculated by multiplying percent
survival estimates from different sampling periods within a year
by one another and then subtracting from 100%.
166
Fig.1.Location of study area in southcentra1 Alaska where causes of short
yearling moose mortality were studied from March 1979 through June 1981.
-----_._-----------
_Af4r:IJSKA ~vi(
-:::;...::::-:I.-::::-~---..:::.--.:.~--==.==--==.-----=::----:-~-~------:-~
CHUGACH MOUNTAIN RANGE-------
15 10 5 a 25
E3 E3 I
1510 5 0 25
E3 E3 i
'..
167
50km
I
,\\,I
-
TI
I
r
r
I
RESULTS AND DISCUSSION
Capture related statistics including morphometric and blood para-
meters for short yearling moose captured in March-April 1979 were
presented in Ballard and Gardner (1980)and will not be repeated
here.Table 1 summarizes these statistics for Area 2 calves cap-
tured in November 1979.
During this reporting period a paper comparing tooth eruption
patterns between moose populations from I sle Royale,Michigan,
Kenai Peninsula,Alaska and the Nelchina Basin,Alaska was pre-
pared and authored by Rolf Peterson,Charles Schwartz and
Warren Ballard.A draft copy of the manuscript is presented in
Appendix I.Also,a paper comparing bone marrow fat between
longbones and mandibles for moose and caribou (Rangifer tarandus)
was presented at the 17th North American Moose Conference and
Workshop and is presented in Appendix I I.
1978 Cohort
The status on June 1,1981 of 64 short yearling moose captured in
Area 1 during 1979 is given in Table 2.As reported earlier,a
large number (n=22,34%)of the radio-collars feel off due to
problems with collar design (Ballard and Gardner 1980).Of the
remaining radio-collared short yearlings,17 were alive June 1,
1981,and the status of 8 was unknown because of either dispersal
or radio failure.Of the 34 moose of known status,17 (50%)had
died.Causes of death for these were:10 winter-killed (59%),
3 brown bear predation (18%),2 hunter kills (12%),1 uniden-
tified predation (6%)I and 1 tagging mortali ty (6%).
Table 3 summarizes the rates and causes of annual calf and year-
ling moose mortality for Area 1 from March 1979 through June
1981.Moose which either had lost their collars or whose status
168
Table 1.Location,ages.physical measurements.incisor status and other statistics associated with capturing
and collaring of 34 calf moose in Game Management Unit 13 from November 27.1979 through November 30.1979.
---_.._-
Measurements (cm)Drug
Accession Age Total Hind Neck Chest Dosage Induction
U Date Collaring Location Sex (Months)Length Foot Circum.Girth (cc)Time
120375 11/29/79 Between Windy and M 5 215 67 69 156 5
V.a1dez Creek
120376 11/29/79 Windy Creek M 5 210 67 65 148 5
120377 11/29/79 Windy Creek F 5 207 65 65 136 5
120378 11/29/79 Windy Creek F 5 212 66 67 162 5
120379 11/29/79 Between Windy and M 5 ----------5 7
Valdez Creek
120380 11/29/81 Valdez Creek F 5 206 69 60 142 5
"120381 11/29/81 Middle Fork of M 5 ----------5
0 Susitna River
120382 11n9/81 Middle Fork Susitna M 5 ----------5 9
River.
120383 11/29/79 Valdez Creek F 5 213 72 67 173 5 9
120384 11/29/79 Valdez Creek F 5 218 70 62 152 5
120385 11/29/79 Valdez Creek M 5 212 73 64 157 5
120386 11/29/79 Gracious House F 5 202 67 66 152 5
120387 11/29/79 Susitna Lodge F 5 222 68 65 157 5 9
120388 11/27/79 Windy Creek M 5 205 72 70 163 5 9
J 1 '..~1 I I J J J J 1 J ,I J I I ~
-~l ~...1 <--1
Table 1.(cont'd)
Measurements (cm)Drug
Accession Age Total Hind Neck Chest Dosage Induction
II Date Collaring Location Sex (Months)Length Foot Circum.Girth (cc)Time
120389 11/27/79 Between Windy and F 5 205 70 67 150 5 7
Valdez Creek
120390 11/27/79 Between Windy and F 5 202 --68 ---5
Valdez Creek,
120391 11/27/79 Between Windy and M 5 214 72 70 163 5 15
Valdez Creek
120392 11/27/79 Between Valdez and M 5 190 --67 155 5
Windy Creek
......120393 11/27/79 Between Windy and M 5 210 -------5
-...l Valdez Creek
0
120394 11/27/79 Between Windy and F 5 205 -------5
Valdez Creek
120395 11/27/79 Between Windy and M 5 200 --79 ---5
Val!lez Creek
120396 11/30/79 West Fork Susitna River M 5 197 63 66 144 5 3
120397 11/30/79 West Fork Susitna River F 5 199 67 61 140 5 11
120398 11/30/79 Mdl Fork Susitna River M 5 200 68 68 162 5 12
120399 11/30/79 Mdl Fork Susitna River M 5 202 65 70 158 5 12
120400 11/30/79 West Fork Susitna River M 5 211 67 62 148 5 ))24
J J J !B 1 I )J J I )J J ••J ,J
-~--'J ==--1 =--1 -----=1
Table 2.Summary of the status by date of 64 radio-collared short yearling moose from March 1979 to 1 June 1981 in the
Nelchina Basin of Southcentral Alaska.
Surviving Lost Unk.
Radio to Slipped Radio Bear Winter Hunting Tagging I.D.
LD.II Sex II 6/1/81 Collar Contact Predation Kill Mortality Mortality Pred.
120255 F 3600 6/7 /79
120227 M 3617 4/5/79
120229 M 3586 7/2/79
120231 M 3602 5/29/79
120233 F 3589 7/9/79
120235 M 3603 5/31/79
120236 F 3606 4/8/79
120237 F 3619 4/30/79
120239 F 3611 4/11/79
120241 ~1 3582 9/79
120244 F 3581 4/5/79
120249 M 3594 X
120250 M 3616 4/30/79
120252 F 3610 5/22/79
120253 F 3583 5/79
t-'120254 M 3604 4/5/79""-J
N 120255 F 3592 4/8/79
120256 F 3620 4/5/79
120257 M 3607 6/23/79
120258 F 3599 4/5/79
120259 F 3585 4/5/79
120260 F 3561 4/5/79
120261 M 3629 10/7 /80
120262 F 3621 10/7/80
120263 F 3584 4/14/79
120264 F 3588 X
120265 M 3601 1/80
120266 M 3597 4/5/79
120267 F 3618 4/8/79
120268 M 3596 3/08/80
120269 M 3590 X
120270 M 3593 4/6/79
Table 2.(cont'd)
Surviving Lost Unk.
Radio to Slipped Radio Bear Winter Hunting Tagging 1.D.
l.D.II Sex II 6/1/81 Collar Contact Predation Kill Mortality Mortality Pred.
120271 M 3613 8/7/80
120273 F 3612 4/14/79
120275 F 3615 4/10/79
120276 F 3622 4/12/79
120277 M 3623 5/2/79
120278 M 3598 4/20/79
120279 F 3614 4/6/79
120280 M 3608 4/30/79
120281 M 3605 4/27/79
120284 M 3587 4/27/79
120285 M _3595 4/27/79
120287 M 3582 5/2/79
120288 F 3620 4/20/79
120290 F 3599 X
120291 F 3618 X
120292 M 3507 5/18/79
120293 F 3604 X
120294 M 3615 X
120295 M 3591 8/4/79
120296 F 3587 5/18/79
120297 F 3595 X
120298 F 3612 X
120299 l!'3592 X
120300 F 3598 X
120301 M 3622 X
120302 F 3611 X
120303 F 3605 X
120304 M 3617 9/12/79
120305 F 3606 3/18/81
120306 .F 3620 X
120307 F 3593 X
120308 F 3581 X
B .1 J _J J J J J I
"J "1 I ~~!
".I _J
Table 3.Rates and causes of annual mortality (March,1979 to June,1981)of
moose captured as short yearlings in late winter 1979 in GMU-13 of
Southcentral Alaska.
r
I Dates!
3/79 -6/1/79 1/6/2/79 -6/1/80 6/2/80 -6/1/81
No.%No.%No.%
Sample Size 43 100 ~100 U 100
Causes of mortality
Natural
Winter Kill 10 23 0 0 0 0
Bear Predation 2 5 1 4 0 0
Unknown Predation 1 2 0 0 0 0
Subtotal 13 30 1 4 0 0
Hunting 0 0 2 8 0 0
Total 13 30 3 13 0 0.-No.Surviving 30 70 21 87 17 100
I
i 1/Does not include period from birth up to March 1979.
:r
'I
,~
174
as of June 1 was unknown were excluded from calculations for the
preceding year.Natural causes,primarily winter kill (77%of
mortali ty),accounted for 30%mortality of the short yearling
moose from late March to 1 June 1979.Results of calf mortality
studies during 1"977 and 1978 indicated that 45%of the moose
calves survived to November 1 (Ballard et al.1981).Based upon
this survival rate and that of short yearlings (70%)for every
100 calves produced in 1978,a minimum of 69 died.However,this
figure is conservative because it excludes the period between
November 1 to late March during which time some moose doubtless
died during winter 1978-79,the second most severe winter in
terms of total snowfall and due to wolf predation.During winter
1978-79,7 of 17 (41%)short yearlings of radio-collared cows
perished (Ballard and Taylor 1980).Thus,a more accurate es-
timate of first year mortality during a severe winter was 73%;
however,even this estimate may be low--because 16 of 33 of the
radio-collared adults occupied an area of low wolf density
(Ballard and Taylor 1978).Correspondingly,many of the short
yearling losses were probably attributable to winter-kill.
During winters 1978-79 and 1979-80 it was estimated that two wolf
packs had preyed upon 9 to 24%of the short yearlings during late
winter (Ballard et al.1981).During this study no losses of
radio-collared yearlings were attributable to wolf predation even
though wolf studies indicated short yearlings were being
killed.We do not know if the lack of wolf predation on the
radio-collared yearlings during March through June was represen-
tative'or if sample size or other unknown factors precluded its
identification.Thus including losses due to winter wolf pre-
dation we estimate that first-year mortality of moose during this
year wi th a severe winter ranged from 79 to 84%based upon the
following assumptions:55%newborn moose calf mortality attribu-
table primarily to bear predation;30 to 41%winter and spring
morta1i ty of surviving members of this cohort primarily from
starvation and bear preda~ion;and a maximum of 24%mortality due
to wolf predation (total mortality estimated by multiplying in-
dividual survival rates,and subtracting from 100%,therefore,
summing the individual rates will exceed 100%).
175
~,
r-
I
Yearling mortality of the 1978 cohort was 14%,however,hunting-_.-- -"--.-.-.
losses accounted for 66%of this mortality.Therefore,second-
year natural mortality was estimated at 5%which was highly com-
parable to the 6%adult mortality estimated by Ballard and Taylor
(1980)for adult moose.No mortality was observed for 2-year-old
moose.
Rates and causes of mortality for the 1978 bull cohort from late
March through May 1979 are summarized in Table 4.Mortality
during this time period was estimated at 43%,while yearling (2nd
year of life)losses including hunting mortality was 38%.Ex-
cluding hunting,natural mortality from June 2,1979 to June I,
1980 was 17%.No mortality was observed among 2-year olds,which
suggests that new hunting regulations based upon antler restric-
tion (legal bull must have antler spread of 36 inches or more)
are protecting this cohort from hunting mortality.Winter kill
(n =7)and bear predation (n =3)were the causes of natural
mortality.Rate of natural mortality for males of the 1978 co-
hort during the first and second year of life was significantly
(chi-square,P<O.05)greater than that for females.Based upon
blood parameters used by Franzmann and LeResche (1978)to assess
the physical condition of adult moose,male short yearling moose
were in poorer physicalcondi tion at the time of capture than
were females (Ballard and Gardner 1980).
1979 Cohort
The status of 34 calf moose studied in Area 2 from November 1979
through June 1981 is summarized in Table 5.Twenty-seven moose
were known to be alive as of June I,1981.Two died from winter-
kill in March 1980 while two others died of unknown causes and
one slipped its collar during winter 1980-81.Radio contact with
four moose (3 in 1980 and 1 in 1981)was lost due to either radio
failure or dispersal.
176
Table 4.Rates and causes of annual mortality of bull moose captured
as short yearlings in late winter 1979 in GMU-13 of
Southcentral Alaska.
177
-
1 --'-=-=}C---i ----)
Table 5.Summary of the status by date of 64 radio-collared short yearling moose from March 1979 to 1 June 1981 in
the Nelchina Basin by Southcentral Alaska.
Surviving Lost Unk.
Radio to Slipped Radio Bear Winter Hunting Tagging I.D.
LD.1/Sex 1/6/1/81 Collar Contact Predation Kill Mortality Mortality Pred.
120375 M 5712 X
120376 M 5517 12/2/80
120377 F 5199 3/19/81 !I
120378 F 5190 X
120379 M 5185 X
120380 F 5181 X
120381 M 5197 X
120382 M 5182 X
120383 F 5186 7/18/80
120384 F 5175 X
120385 M 5196 3/27/80
120386 F 5171 X
120387 F 5195 X
120388 M 5180 X
120389 F 5176 X
120390 F 5192 X
f-'120391 M 5193 X
"'-l 120392 M 5174 Xco120393M5191X
120394 F 5194 X
120395 M 5200 X
120396 M 5184 X
120397 F 5179 X
120398 M 3992 X
120399 M 3597 12/6/80
120400 M 5183 X
120401 M 5178 3/6/80
120402 M 5198 X
120403 F 3623 X
120404 F 3602 X
120405 F 3614 3/6/80
120406 F 5187 3/19/81
120407 F 5173 X
120408 M 5188 X
!I Radio-failure confirmed-Present with radio-collared twin.
Table 6 summarizes rates and causes of annual mortality to 1 June
1981 for moose captured as calves in November 1979.Excluding
early neonatal losses (pre-November),first-year (11/79-6/1/80)
mortality due to winter kill was 6.1%,which was significantly
less (P 0.05)than that suffered by the 1978 cohort during the
severe winter of 1978-79.This significantly lower mortality
rate can be attributed to mi Ider snow conditions during winter
1979-80 and perhaps to reduced bear predation.High survival was
also confirmed during a spring composition survey flown on May,
23,1980 (Table 7).Because bull moose cannot be accurately
identified from aircraft during.spring,we assumed the spring sex
ratio of adults was identical to the fall 1979 bull:cow ratio
(18M=100E').Subtracting adult bull s (N=14)from the total num-
bers of adults counted (n=93)yielded an estimate of 58 short
yearlings:100 cows (includes cows ~2 yr.olds).This estimate
was comparable with the fall ratio of 58 calves:100 cows
(~2 yr.old)which indicates that survival was quite high.
Second year survival of the 1979 cohort during winter 1980-81 was
similar to that experienced by the 1978 cohort during 1979-80;
96%survival in 1980-81 for the 1979 cohort versus 95%(excluding
hunting)survival in 1979-80 for the 1978 cohort.
Evaluation of Brown Bear Transplant
Routine moose sex and age composition counts were conducted in
Moose Count Area 3 during early November 1980.Immediately fol-
lowing the composition count,the area was censused using quadrat
sampling techniques developed by Gasaway (1978),Gasaway et ale
(1979),and Gasaway and Dubois (unpub.report).Comparison of
the resulting sex and age composition data in addition to the
fall population estimate is summarized in Table 8.Similar to
comparisons made elsewhere in Alaska calf:100 cow ratios provided
by quadrat sampling methods were considerably higher than those
provided from standard composition counts (Gasaway pers.corom.).
179
""'"
~I
""'"
Table 6.Rates and causes of annual mortality of moose captured
as calves in November,1979 in GMU-13 of Southcentra1 Alaska.
Dates
11/2/79 6/1/80 Jj 6/2/80 6/1/81
No.%No.%
Causes of mortality 33 100 28 100
Natural Mortality
Winter Kill 2 6 0 0
Bear Predation 0 0 0 0
Unknown predation 0 0 0 0
Unknown causes 0 0 1 4
Subtotal 2 6 1 4
Hunting 0 0 0 0
Total 2 6 1 4
No.Surviving 31 94 27 96
}j Does not include period from birth up to November 1979.
180
Table 7.Summary of spring moose composition survey conducted in
Moose Count Area 3 on 23 ~my 1980 in the upper Susitna
River Basin of Southcentral Alaska.
II Adults =93
#Short Yearlings =46
%Short Yearlings =36%
Count time:250 minutes
181
-
~i
~,
~--'-i ------'1 ~----i ------1
Table 8.Comparison of moose sex-age composition data collected from standard moose surveys to that obtained from quadrat sampling techniques
used in Moose Count Area 3 during November 1980 in the upper Susitna River Basin.
Date
Tot.bulls
per
100 cows
Sm.Bulls
per
100 cows
Sm.bulls
per 100
19.bulls
Sm.bulls Calves Calves
%in per 100 per
Herd cows >2 yrs.100 cows
Incidence Calf
of twins per %in
100 cows w/calf herd
Animals
per
hour
Total
sample
Count Area
time Samp~ed Hinu~es/
(hr.)(mi)(mi)
Composition 11/1-2 36.7 21.8 146.4 11.9 40.1 31.4 16.3 17 .2 37.0
count
Census 11/2-4 29.9 20.1 203.9 11.6 55.0 43.9 13.9 25.3 27.8
Stratification 11/1 ----------------89.9
x moose/group %of observed moose %of observed moose %of observed moose
comprised of singles comprised of pairs comprised of groups of 3
Composition
count 2.7 9.6
Census 2.6 7.6
I-'Stratification 2.7 7.5
CO
N
29.9
34.4
32.1
8.3
20.9
17.6
344 9.3 273.5 2.0
459 16.5 247.5 4.0
187 2.15 273.5 .5
%of observed moose
comprised of groups of 4 or more
5·2.2
37.0
42.8
Population Estimate =473 -+38 (90%CI -435 -510).
(uncorrected for observability)
This discrepancy is related to survey intensity and a higher
probability of observing large groups of moose which proportion-
ately contain fewer calves.
We used the relationship between the 1980 moose composition
counts and the quadrat sampling to recompute ratios obtained in
1979 following the removal of 48 brown bears by transplant
(Ballard et al.1980;Miller and Ballard in press).Based upon
the 1979 calf:cow ratio of 52.2 estimated from a composition
count,the estimated ratio,had the area been censused,would
have been 72.9 calves/100 cows,while the bull:cow ratio would
have been 14.6.Similarly,the calf percentage of the herd would
have been 45.2%.We estimated calf survival in moose count area
3 following the bear transplant by applying these ratios to a
hypothetical moose population of 1,000.According to our projec-
tions,this hypothetical population in early November would have
been comprised of 450 calves,83 bulls,60 yearlings cows (as-
sumed to be sexually immature)and 407 cows.Assuming a preg-
nancy rate of 90%(Ballard and Taylor 1980)366 cows should have
produced calves in 1979.Twinning rates were calculated based
upon 89 newborn moose calves captured in count area 3 from 1977
through 1979,yielding a twinning rate of 35%or a gross pro-
duction of 135 ca1ves/100 cows (Ballard et al.1980).Therefore,
494 calves should have been produced.Since 450 calves theoret-
ically were alive by 1 November,after the period when most neo-
natal losses occur (Ballard et a1.1981),an estimated 44 calves
died following a reduction in bear density of approximately 60%
(Ballard et al.1980).This provides an estimate of 9%calf mor-
tality from birth to November 1 in 1979.Similarly to the above
analysis we applied the relationship between the 1980 composition
count and census data to the 1977 and 1978 composition counts
which were conducted prior to the reduction in bear density.The
resulting estimated mortality rates were 60 and 55%for 1977 and
1978,respectively.These latter estimates were quite similar to
those observed on radio-collared moose calves.
183
~I
~,
~,
~
I
f'!""
I
I
Calf mortality studies conducted in GMU-13 during 1977 and 1978
prior to bear removal suggested that 55%of the newborn moose
calves died between birth and 1 November of each year (Ballard et
al.1981).During 1979 after bears had been removed the radio-
collared calf data continued to indicate that about half of the
calves were being killed by bears.We discounted·these data for
this analysis because of the following:1)relatively small
sample size (27 in 1979 versus 120 in 1977 and 78);2)the
smaller sample of calves was concentrated in a relatively small
area which made calves vulnerable to a relatively small number of
bears (Ballard et al.1980);and 3)comparisons of fall calf:cow
ratios with other unmanipulated moose count areas suggested that
there had been a significant improvement in calf survival (op.
cit.).Based upon the estimated mortality rate derived from the
1979 composition count the temporary reduction in brown bear den-
si ty may have reduced calf mortality from 55%to 9%,an 84%de-
crease in total mortality.
Mortality of calves and short yearlings in count area 3 during
winter 1979-80 was estimated at 6%.Therefore,.during the year
of the bear reduction program,first year mortality of the 1979
cohort due to early neonatal losses and winter kill totaled an
estimated 14%.No losses were attributable to wolf predation;
however,if wolves had preyed upon 24%of the calves (the maximum
estimate of wolf predation)and short yearlings from 1 November
through early spring 1979,the first year mortality rate would
have been 36%.
Table 9 summarizes approximate rates by cause of first year moose
mortality for the Nelchina and upper Susitna River Basins as de-
termined from several studies conducted from 1975 through 1981.
Based upon these estimates,first year moose mortality varied
from 23 to 84%depending upon the magni tude of bear predation,
wolf predation,and winter severity.These figures do not con-
sider winter-kill and wolf predation as compensatory mortality
factors,nor do they consider variations in moose densi ty.
184
Table 9.Estimates of first year moose mortality by cause and time period for the
Nelchina and Susitna River Basins of Southcentral Alaska.
%Mortality
birth -1 Nov.
(level of bear
predation)
High 1..1
.55
Low 21
.09 -
%Mortality
from winter kill-
1 Nov.-1 June
(winter severity)
Severe 31
.41 -
Mild 41
.06 -
Severe 31
.41 -
Mild 41
.06 -
%Mortality
from wolf predation-
1 Nov.-1 June
(predation level)
High 51
.24 -
Low 61
.09 -
High 51
.24 -
Low 61
.09 -
High 'il
.24
Low 61
.09 -
High i/
.24
Low 61
.09 -
Total calcu-
lated first
year
mortality 21
.84
.77
.68
.62
.68
.54
.36
.23
-
1..1 Mortality rate estimated from calf mortality studies (Ballard et al.1981)
and includes 20%of total mortality which was not attributable to brown
bear predation.
~I Based upon estimated calf production and survival following a 58%
reduction (transplant)in brown bear density (see text,this report).
11 Determined from observations of short yearling losses of radio-collared
adult moose during 1978-79 (Ballard and Taylor 1980).
il Determined from observations of radio-collared calf moose during 1979-80.
'il Extrapolated from predation rates for two wolf packs numbering 7-8 wolves
studies during winter 1979-80 (Ballard et al.1981).Percent of calves
preyed upon was determined by estimating the total pack area moose
population by estimating %of moose observed according to survey
intensity (from Gasaway and Dubois,unpub.rept.).
il Extrapolated from predation rate for one wolf pack numbering 2 wolves
studied during winter 1978-79 (Ballard et al.1981).Percent calves
preyed upon estimated same as that described for tl5.
II Assumes no compensatory mortality.Annual mortality was estimated by
determining percent survival from birth to 1 November and from 1 Nov-
ember to 1 June.The estimates were then multiplied and the sum
subtracted from 100 percent.
185
T
r
I
r
Nevertheless,they provide a general estimate of the extent of
mortality from the three major mortality factors.This analysis
suggests that.the largest increases in survival occurred when
brown bear densities were temporarily reduced to the level at-
tained during the bear transplant.
According to this simple model with high bear predation and
severe winters,a reduction in wolf pack size from 7-8 to 2
wolves only resulted in a 7%decrease in mortality.A simi lar
difference also occurred with the same conditions,but during
mild winters.However,the difference between high and low wolf
predation with high bear predation levels during severe versus
mild winters was 22%.
During years of low bear predation such as that following the
bear transplant (Ballard et al.1980)a difference in high and
low wolf predation during severe winters would result in an es-
timated difference of 14%mortality,while during mild winters
the projected difference was 13%(Table 9).With low predation
by both bears and wolves during a mi ld winter,fi rst-year mor-
tali ty was at its lowest level (23%).
The high survival of calf and yearling moose in Study Area 2
could be attributed to reduced brown bear predation as a result
of the brown bear transplant (Ballard et al.1980)and the mild
winter of 1979-80.The high survival rates documented in this
study suggest that at least on a short term basis,the study
area's.moose population was not being limited by deteriorating
range conditions and that predation,primarily by brown bears,
was preventing the moose population from increasing.
186
1.
2.
RECOMMENDATION
Continue to monitor survival of moose captured in 1979 and
1981.
Ini tiate a long term study to develop a sati sfactory brown
bear harvest strategy which would reduce bear predation and
allow the moose population to increase while maintaining a
viable bear population.
-
ACKNOWLEDGMENTS
Sterling Eide,Albert Franzmann,Dennis McAllister,Christian
Smi th,Robert Tobey,John Westlund,(all Alaska Department of
Fish and Game)and William Martin (U.S.Fish and Wildlife
Service)participated in the tagging operations.
Special appreciation is extended to Mr.Alfred Lee,Lee's Air
Taxi Service,not only for donating flying time,but also for
helping pick up the radio collars which slipped off moose.Many
of the projects successes could be attributed to the interest and
effort given by Mr.Lee.
Appreciation is also extended to Vern Lofstedt,Kenai Air Ser-
vice,for piloting the helicopter,and to Ken Bunch,Sportsman's
Flying Service t for assistance with monitoring radio-collared
moose.
Karl Schneider and G.Bos reviewed this report and made a number
of suggestions for improvement.SuzAnne Miller advised us on
statistical procedure.
187
-
~I
LITERATURE CITED
Ballard,W.B.,and K.P.Taylor.
Moose Population Study.Alaska
Aid Wildl.Rest.Proj.Prog.
Juneau.62pp.
1978.
Dept.
Rept.
Upper Susitna River
Fish and Game,Fed.
W-17-l0,Jub 1.20R.
r
A.W.Franzmann,K.P.Taylor,T.Spraker,c.C.
Schwartz,and R.O.Peterson.1979.Compari son of tech-
niques utilized to determine moose calf mortality in Alaska.
15th N.Am.Moose Conf.Workshop,Kenai,Alaska.
and K.P.Taylor.1980.Upper Susitna Valley moose
population study.Alaska Dept.Fish and Game,Fed.Aid
Wildl.Rest.Proj.Final Rept.W-17-9 and W-17-ll,Job
1.20R.Juneau.102pp.
S.D.Miller,and T.H.Spraker.1980.Moose calf mor-
tali ty study,Game Management Unit 13.Alaska Dept.Fi sh
and Game,Fed.Aid Wildl.Rest.Proj.Final Rept.W-17-9.
l23pp.
and C.L.Gardner.1980.Nelchina Yearling Moose Mor-
tali ty study.Alaska Dept.Fi sh and Game,Fed.Aid Wi ldl.
Rest.Proj.Rept.W-17-ll and W-2l-1.22pp.
1981.Grey wolf-brown bear relationships in the Nelchina
Basin of southcentral Alaska.F.H.Harrington and P.C.
Paquet,Co.eds.Portland Wolf Symposium.Portland,Ore-
gon:in press.r
I
_____,R.O.Stephenson and T.H.Spraker.
Wolf Studies.Alaska Dept.of Fi sh
Rept.,W-17-9 aND W-17-l0.20lpp.
188
1981.Nelchina Basin
and Game.P-R Proj.
Franzmann,A.W.,and R.E.LeResche.1978.Alaskan moose blood
studies with emphasis on condition evaluation.J.Wildl.
Manage.42:344-351.
Gasaway,W.1978.Moose survey procedures development.Alaska
Dept.Fi sh and Game.P-R Pro j.Rept.47pp.
S.J.Harbo,and S.D.DuBois.1979.Moose survey pro-
cedures development.Alaska Dept.Fish and Game.P-R Proj.
Rept.87pp.
Skoog,R.O.1968.Ecology of
granti).in Alaska.Ph.D.
Berkeley.689pp.
PREPARED BY:
Warren B.Ballard
Game Biologist III
SUBMITTED BY:
Karl B.Schneider
Regional Research Coordinator
189
the caribou (Rangifer tarandus
Thesis.Univ.California,
APPROVED BY:
Director,Division of Game
Research Chief,Division of Game -
.....
-
APPENDIX E
190
T
~ppendix E.Daily activity,movements and association of eight members of the Watana
wolf pack as determined from aerial observations from 6 May through
1 July 1981 in the Susitna River Basin of southcentral Alaska.
Wolf -308 -Adult gray femalerr-------------------------------------
f'..ate-Time
•Obs.iF
Distance
traveled
(mi)
Direction
of travel Location
Distance
#Assoc.from den
(ID)(mi)
Direction
from den Misc.
Ti/6-5/8/81 a
am 1600
(11)r
!;/9/81 a
pmfll)
5/10/81 a
fii~
5/11/81 a
n005
;1 :11)
den
den
den
den
1(324)
1(324)
a
a
a
a
a
a
f'i./12 /81
:1 :853,(11)
r:i ;/13/81
!'1.625
(11)
fi/14/81
0820
11)
5/15/81
j:~i~
5/16/81
r~~~
M/17/81
I ~800
;1 (11)
a
a
a
a
a
a
den
den
den
den
den
den
191
2(323)
(325)
3(323)
(324)
(346)
1(324)
2(323)
(346)
2(324)
(325)
1(323)
a
a
a
a
323 returned
346 returned
-
193
194
r, I
Wolf -308 -Adult gray female (cont'd)
Distance Distance
T)ate-Time traveled Direction #I Assoc.from den Direction
Obs.if (mi)of travel Location (ID)(mi)from den Misc.
I'i
I 5/23/81 0 den 5 (11)0
1100rll)
u 7/1/81 .8 N N of den 4(323).8 N Rendez-
1353 (324)vous site
f(24)(345)
(346)
r1,1
Wolf -323 -Yearling gray female
r Distance Distance
Date-Time traveled Direction IF Assoc.from den Direction
rObS.if (mi)of travel Location (ID)(mi)from den Misc.
r/ll/81 2.0 SW W.F.0 39.5 ESE
[I 1225 Gulkana
(59)
r>/12/81 39.2 ENE den 2(308)0
•1853 (325)
(11)
fS/13/81 0 den 3(308)0
1625 (324)rll )(346)
5/14/81 1.0 NE near den 0 1.0 NE
~810
•(65)
5/15/81 1.0 SW den 2(308)0
r1530 (346)
II
5/17/81 0 0 den 1(308)0
'!"""1800
Ii (11)
r
,!
I
195
Wolf -323 -Yearling gray female (cont'd)~~
Distance Distance ~
Date-Time traveled Direction iF Assoc.from den Direction
Obs.iF (mi)of travel Location (ID)(mi)from den Misc.
~
5/18/81 0 0 den 2(308)0
1410 (324)
(11)
5/21/81 7.5 NE .East of 0 7.5 NE
1020 Big Lake ~
(76)
5/22/81·7.5 SW den 2(308)0
1730 (346)~
(11)
5/23/81 4.5 S E.of St.1(325)4.5 S
0950 bear Lake
(81)
5/24/81 4.5 N den 2(308)0
1855 (346)
(11)
5/24/81 5.5 S Is!'on 1(325)5.5 S Ravens
1945 Sue in area.
(85)2 bear -closeby.
5/25/81 5.5 N den 3(308)0 ,~
0955 (324)
(11)(346)
5/26/81 6.3 S S.of 0 6.3 S --
1840 River
(90)
~
5/30/81 6.3 N den 2(308)0
1610
(11)
~,
5/31/81 0 den 3(308)0
1634 (324)
(11)(325)
196
,....
I
Wolf -323 -Yearling gray female (cont'd)
Distance Distance
rate-Time traveled Direction #Assoc.from den Direction
·Obs.#(mi)of travel Location (ID)(mi)from den Misc.
T6/1/81 2.3 E E.of 1(325)2.3 E On 50%
·0925 den consumed
r(l09)calf
:1
moose.
6/2/81 2.3 W den 3(308)0r1930(345)
!(11)(346)r6/3 /81 0 den 3(308)0
i 1255 (324)
·(ll)(346)
f6 /4 /81 0 den 5(308)0
..1945 (324)
(1l)(345)
(346)
6/5/81 4.0 SSE 2 mi.E.4(308)4.0 SSE Suspect
r1930 Std.bear (324)traveling
!i,(l00)Lk (346)from
"I (l )caribou
killed by
bI.bear.
6/6/81 4.0 NNE den 4(308)0r0940(324)
I.(11)(346)
(25)-
.1 6/7/81 0 den 4(308)0
,2050 (324)
(11)(325)r (346)
;1 6/8/81 NO DATAn/9 /81 0 den 4(308)0
:1721 (324)
(11)(345)
(346)
197
Wolf -323 -Yearling gray female (cont'd).EIUm
Distance Distance
Date-Time traveled Direction IF Assoc.from den Direction
Obs.IF (mi)of travel Location (ID)(mi)from den Misc.
6/10/81 0 den 3(308)0
1145 (345)
(11)(346)
6/11/81 0 den 5(308)0
1031 (324)
(11)(325)
(345)
(346)
6/12/81 0 den 5(308)0
2028 (324)
(11)(325)~
(345)
(346)
6/13/81 0 den 3(308)0
1245 (324)
(11)(325)
~
6/15/81 0 den 4(308)0
1557 (324)
(11)(325)~
(345)
6/16/81 0 den 2(308)0
1139 (346)
(11)
6/18/81 0 den 2(308)0 -1600 (346)
(11)
6/19/81 10.3 SW Fog 3(308)10.3 SW on calf
1200 Lake (345)moose
(124)(346)kill.
6/21/81 10.3 NE den 2(308)0
1645 (346)
(11)~
198
Wolf -323 -Yearling gray female (cont'd)
Distance Distance
WDate-Time traveled Direction iF Assoc.from den Direction
.'Obs.iF (mi)of travel Location (ID)(mi)from den Misc.
J6/22/81 0 den 5(308)0
1230 (324)
11).(325)
(345)
(346)
T6/23/81 0 den 5 (11)0
:.100
(11)
f7/1/81 .8 N N.of 4(308).8 N Rendez-
1353 den (325)vous
[(24)(345)site.
(340)
Wolf -324 -Yearling gray female
Distance Distance
rate-Time traveled Direction iF Assoc.from den Direction
IJ Obs.iF (mi)of travel Location (ID)(mi)from den Misc.
0'5/6-5/8/81 3.7 N den 1(308)0
J am 1615
(11)
5/9/81 0 0 den 1(308)0
pm
,(11)
f5/10/8l 5.0 SW Mouth 0 5.0 SW
1600 Watana
r(51)
I,
5/11/81 2.5 NE Delusion ?2.8 SW
~1000
II (55)II
199
200 -
Wolf -324 -Yearling gray female (cont'd)T
I
Distance Distance
rlate-Time traveled Direction iF Assoc.from den Direction
lObs.iF (mi)of travel Location (ID)(mi)from den Misc.
~/27/81 3.0 E E.of den 0 3.0 E
0910
~92)
;IJ/31/81 3.0 W den 3(308)0
1634 (323)r11 )(325)
6/1/81 9.3 SW E.mouth 0 9.3 SWr959Deadmanek.
1110)
~/2/81 10.5 NE N.of den 0 2.8 N on adult
i 945 caribou
kill,
50%con-r sumed.
'I
6/3/81 2.8 S den 3(308)0
f255 (323)
•11)(346)
~':~81 0 den 5(308)0
(323)
,1.11)(325)
(345)
(346)
6/5/81 4.0 SSE 2 mi.E.4(308)4.0 SSE Suspected
f930 Std.Bear (323 traveling
;.100)Lk.from
caribou
kill made
by bI.bear.
6/6/81 4.0 NNE den 4(308)0
f940 (323)
(11)(325)
iF"'(346)
il
II
'l
201
-
Wolf -324 -Yearling gray female (cont'd)
Distance Distance
Date-Time traveled Direction 1ft Assoc.from den Direction
Obs.1ft (mi)of travel Location (ID)(mi)from den Misc.
6/7 /81 0 den 4(308)0
2050 (323)
(ll)(325)-(345)
6/8/81 NO DATA
6/9/81 0 den 4(308)0
1721 (323)
(ll)(345)
(346)
6/10 NO DATA -Absent from den.-
6/ll/81 ??den 5(308)0
1031 (323)
(11)(324)
(345)
(346)
6/12/81 0 den 3(308)a
1245 (323)
(ll)(325)~
6/14/81 5.5 S lsI.on ?5.5 S Suspect
1043 Sue.feeding on
(1l4)on kill of
5/24-6/14.
6/15/81 5.5 N den 4(308)a -1557 (323)
(ll)(325)
(345)
6/18/81 6.8 NW 2.5 mi.a 6.8 NW
1705 .w.Big
(121)Lake
6/19/81 Not at den
ll10
?
~
202
Wolf -324 -Yearling gray female (coned)rII
Distance Distance
T)ate-Time traveled Direction #Assoc.from den Direction
I Obs.#(mi)of travel Location (ID)(mi)from den Misc.
f?/21/81 16.3 SSW Fog 0 13.8 SW On old
1720 Lakes adult mooser126
)
kill.
Wolf -325 -Yearling gray male
r)ate-Time
Distance Distance
traveled Direction iF Assoc.from den Direction
Obs./I (mi)of travel Location (ID)(mi)from den Misc.
[/22/81 13.8 NE den 5(308)0
1230 (323)r:ll )(325)
II (345)
(346)
)/23/81 0 den 5(11)0
1100
(11)
fJ/1/81 .8 N N.of 4(308).8 N Rendez-
1353 den (345)vous site
1:24 )(325)
I (346)
fS/6/-S/8/81 4.0 NW St.Bear 0 4.3 S
I am 1610 Lk
(50)
r.5/10/81 7.5 NE Watana 0 4.7 ENE
1400 Bendr:52 )
'I 5/11/81 2.3 SW Watana 0 2.7 E
,...:lOll Ck.
II I'I :S4)
I""'"
i
203
~
Wolf -325 -Yearling gray male (cont'd)
Distance Distance
11 Assoc.from den Direction -Date-Time traveled Direction
Obs.11 (mi)of travel Location (ID)(mi)from den Misc.
'"""
5/12/81 2.7 ESE den 2(308)0
1853 (323)
(11)..".
5/13/81 4.7 ENE Watana 0 4.7 ENE
1635 Ck.
(52)
5/14/81 3.5 WSW Near den 0 1.2 NE
0815 .
(66)
5/15/81 0 0 den 2(308)0
1020 (324)
(11)-
5/17/81 2.5 SW Delusion 0 2.5 SW
1810 Ck.
(70)-
5/18/81 7.5 NE Upper 5.2 NE
~
5/21/81 4.8 SW den 3 (308)0
0905 (324)
(11)(346)
'-'i
5/22/81 5.8 SSE Susitna 0 5.8 SSE
1725 River
(79)~
5/23/81 1.0 NNW E.St.1(323)4.5 S
0950 bear
(81)
5/24/81 1.0 S lsI.on 1(323)5.5 S Ravens in
1945 Sue 2 -area,
(85)bears
closeby.-
204
r
Wolf -325 -Yearling gray male (cont'd)r
Distance Distance
pate-Time traveled Direction iF Assoc.from den Direction
,Obs.iF (mi)of travel Location (ID)(mi)from den Misc.
r5/25/81 5.8 NNW ESE den 0 1.0 W Possible
,0950 small
(86)mammal
kill.
5/26/81 7.3 SSE S.of 0 7.0 SSE with bears
0 8 :5 5 280 &341
I
I :91)
n/27 /81 7.0 NNW den 2(308)0
!.)900 (346)
..(11)
P/30/81 0 0 den 2(308)0
!1610 (323)
(11)r adult
I 80-90%can.
G/31/81 0 0 den 3(308)0
l634 (323)
(11)(324)
n/1 /81 2.3 E E.of 1(323)2.3 E.on 50%
:J925 den consumed
(l09)calf moose.r
I ,/2/81 3.3 SW 2 mi-0 3.8 SSE
1930 ENE St.r97
)
Bear Lk.
,6/3/81 6.3 NNW N.of 1(345)2.8 N Same kill
1250 den that 324f:98 )pres.on 6/2
6/4/81 2.8 S den 5(308)0r945(323)
I :11)(324).'(345)
iF (346)
jj
r 205
Wolf -325 -Yearling gray male (cont'd)
Distance Distance -Date-Time traveled Direction ifF Assoc.from den Direction ,
Obs.1;(mi)of travel Location (ID)(rni)from den Misc.
6/5/81 0 den 1(345)0 308 travel-
1925 ing on 1st
(11).occasion.~
6/6/81 0 den 4(308)0
0940 (323)-(11)(324)
(346)
6/7 /81 0 1 den 4(308)~
2050 (323
(11)(324),
(346)
6/8/81 NO DATA
NO DATA
,....
6/9 &10 (absent from den)
6/11/81 ??den 5(308)0
1031 (323)
(11)(324)
(345)~
(346)
6/12/81 0 den 5(308)0
2028 (323)
(11)(324)
(345)
(346)
6/13/81 0 den 3(308)0
1245 (323)""'"(11)(324)
6/15/81 0 den 4(308)o .
oClI!mf'I
1557 (323)
(11)(324)
(345)
,""",
~,
206
Wolf -325 -Yearling gray male (cont'd)
fr'
'I
;j
Distance Distance
rate-Time traveled Direction if Assoc.from den Direction
;.1 Obs.#(mi)of travel Location (ID)(mi)from den Misc.
;/10/81 3.3 S 1 mi.N.0 S 308 travel-
1148 ing on 1st
,-'117)occasion.
:1 j/17/81 2.5 SSE lsI.on 0 5.5 S Same loca-
1255 tion as
,114)wolves on
5/24,6/14.
r;/18/81 1.0 S South of 0 6.5 S
II
I'!L300'!.(122)
rl/19/81 1.0 S Isle of 0 5.5 S same lac.
1 1030 River as 5/24,
(114)6/14,6/17.
r'
I
I
I
i
6/21/81 .5 N N.of 0 5.0 Sr630Island
",:125)
I
6/22/81 5.8 N den 5(308)0r
:'1 l245 (323)
J ~11)(324)
(345)
(346)
6/23/81 0 den 3(308)0
,M024 (323)
,'11)
I '(324)
(345)
(346)
7/1/81 .8 N N.of 4(308).8 N Rendezvousr353den(323)site
,I :24)(345)
(346)
207
""'"
Wolf -346 -Yearling gray female
Distance Distance
Date-Time traveled Direction tl Assoc.from den Direction
Obs.tF (mi)of travel Location (ID)(mi)from den Misc.
~
5/11/81 2.5 W Little 0 38.2 ESE
1250 Gap
(60)Lake
5/12/81 NO SIGNAL .....,
5/13/81 38.2 WNW den 3(308)0
1645 (323)
(11)(324)-,
5/14/81 1.7 NE close 1.7 NE
0815 to den ~~
(67)
5/15/81 1.7 SW den 2(308)0 0
1530 323
(11)
5/16/81 8.2 SE opp.8.2 SE Rendezvous
1015 Kosina
(68)Ck.
5/17/81 8.7 NW close to 1.2 SW
1813 den
(69)
5/18/81 8.7 SE opp.8.2 SE suspect
1400 Kosina kill-same
(73)Creek loco as 5/16
5/21/81 8.3 NE den 3(308)0
0905 (324)~j
(11)(325)
5/22/81 0 den 2(308)0 ~
1730 (323)
(11)
5/23/81 8.5 SE On Sus.0 8.5 SE same loco
1000 opp.Kosina as tl346
(83)on 5/18.
jIIIlIII':1I
208
r
r Wolf -346 -Yearling gray female (cont'd)
f>ate-Time
Distance Distance
traveled Direction 11 Assoc.from den Direction
,Obs.II (mi)of travel Location (ID)(mi)from den Misc.
r
d ;/24/81 8.5 NW den 2(308)0
p855 (323)
,:11)
r/25 /81 0 den 3(308)0
:)955 (323)
,{11)(324)
[:/26/81 0 den 3(308)0
(324)
(11)
>/27/81 0 den 3(308)0
0900 (325)r11
)
Jj/30/81 10.5 SE near 0 0.5 SE
1600 den
107)
6/1/81 0.5 NW den 1(308)0 Rendezvousr20
f :11)
n/2 /81 0 den 3(308)0
1 1930 (323)
I ~11)(345)
1'/3/81 0 den 3(308)0
'I l730 (323)
"(11)(324)
f;/4/81 0 den 5(308}0
,1945 (323)
r'11)(324)
(325)
I (345)
ri/5/81 4.0 SSE 2 mi.E 4(308)4.0 SSE Suspect travel-
~I
ii 1930 (323)ling from
(324)caribou kill
(l )made by
bi.bear.
209
-
Wolf -346 -Yearling gray female (cont'd)
Distance Distance -Date-Time traveled Direction If.Assoc.from den Direction
Obs.If.(mi)of travel Location (ID)(mi)from den Misc.
6/6/81 4.0 NNE den 4+(308)0
0940 (323)
(11)(324)
(325 )
(346)
6/7/81 2-5 N N.of den 0 2.5 N.On old
0900 moose kill.
(111)
6/7 /81 2.5 S den 4(308)0
2050 (323)
(11)(324)
(325)
6/8/81 NO DATA 1IlIRIl!',
6/9/81 0 den 4(308)0
1721 (323)
(11)(324)
(345)
6/10/81 0 den 3 (308)0 "'""
1145 (323)
(11)(345)
6/11/81 0 den 5(308)0
1071 (323)
(11)(324)
(325)
(345)
6/12/81 0 den 5(308)0 ~
2028 (323)
(11)(324)
(325)~
(345)
6/13/81 5.5 S 151.on 1(345)5.5 S Suspect
1259 River feeding on
(114)kill same
lac.as
5/24.
210
ii
I
Wolf -346 -Yearling gray female (cont'd)
fJate-Time
Distance Distance
traveled Direction iF Assoc.from den Direction
.,Obs.iF (mi)of travel Location (ID)(mi)from den Misc.
r
6/14/81 5.5 N den 3(308)0r(325)
(345)
rrf/16 /81 0 den (308)0
II i139 (323)
1(11)
r,/18/81 0 den (323)0
I 1600 (308)
(11)
f,/19/81 10.3 SW Fog L.(308)10.3 SW On calf
1200 (323)moose kill.r124 )
li .5/21/81 10.3 NW den (323)0
1645 (308)
11)
6/22/81 0 den (324)0
r 1645 (325)j :41)(346)
(323)
f'/23/81 0 den (324)O·
"1645 (325)
(41)(346)r (323)
7/1/81 0.8 N.of Rendezvous (308)0.8 N Rendezvousr353den(323)
:.'24)(325)(I '(345)
211
212
rr.1 _
f W_O_l_f_-_3_1_0_-_A_d_u_l_t_g_r_ay_m_a_l_e_(_C_on_t_'d_)_
.....
ill'Jate-Time
lObs.11
Distance
traveled
(mi)
Direction
of travel Location
11 Assoc.
(ID)
Distance
from den
(mi)
Direction
from den Misc.
4.5 mi.NW 3(344)
Lone Butte (345)
(1 )
gray)
r>/24/81 \
1820
~84)
I'
115/25/81
~120
'1~87)
5/26/81
f:89)
11.3
2.0
12.8
WSE
S
SW
3.5 E.
Tyone
River
4 mi.ESE
mouth
Tyone River
2(344)
(345)
2(344)
(345)
ESE
36.5
33.5
On adult moose
kill 75%consumed.
ESE
SE
IT""'
[I >/27/81r93)"
I,
U j/28/81
1832
96)
5/29/81r/30 /81
,1025
(105)
L/31/81
1515f105 )
6/1/81
r U38
II :105)
w
12.5 SE
15.0 NE
NO SIGNAL
2.8 NE
o 0
o 0
8 mi.W.
Moore Lk.
Near
Susitna
den
At
Susitna
den
At
Susitna
den
At
Susitna
2(344)
(345)
3(344)
(345)
(1 )
o
o
1(295)
46.0
49.5
53.8
53.8
53.8
SE
SE
SE
SE
SE
On short
yrl.moose
kill.
Wolf 295 of
Susitna Pack
present.
NNW NNW
Tyone Lake
6/2/81
[L817
:96)
'1'/6/81
I L155
.(102)
rI
4.8
2.8 NE
o
At Susitna 0
Rendezvous
Site
213
49.5
53.8
SE
SE
Wolf -310 -Adult gray male (cont'd)""'"
Distance Distance
Date-Time traveled Direction IF Assoc.from den Direction
Obs.IF (mi)of travel Location (ID)(mi)from den Misc.
6/7/81 16.0 NW 1 mi.N.0 39.3 SE
1900 of mouth
(103)Tyone Ck.~
6/8/81 NO DATA
6/9/81 16.3 SE Susitna 2(2 unc.)54.0 SE
1606 den
(105)
"'"
6/10/81 0 Susitna 0 54.0 SE
0928 den
(105)III/IlIl1
6/12/81 12.5 NW W.Tyone 1 (l unc.)43 SE
2202 River ~
(113)
Wolf -344 -Yearling gray male
Distance Distance
Date-Time traveled Direction IF Assoc.from den Direction
Obs.IF (mi)of travel Location (ID)(mi)from den Misc.
5/11/81 3.2 SSW Tyone 0 40.5 ESE
1208 River ~
(57)
5/12/81 10 ..2 SW Tyone 3(310)41.2 SE _.
1800 Ck.(345)
(61)(l unc.)-5/13/81 7.2 N Tyone 2(310)36.5 SE
1715 Ck.(345)
(63)
~~,
5/14/81 5.7 NW Susitna 2(310)31.0 SE On old
0855 Bend (345)adult
(64)moose kill.
214
,-
'I
r Wolf -344 -Yearling gray male (cont'd)
rOate-Time
Distance Distance
traveled Direction if Assoc.from den Direction
,Obs.if (mi)of travel Location (ID)(mi)from den Misc.
....
,15/17/81 9.5 E \Upper 2(310)39.5 ESE Attacking
1915 West (345)cow w/2
f(72).yrls.
:1 5/18/81 2.5 W Little 2(310)37.0 ESE
_1330 Gap (345)
[I (75)
'I
5/21/81 21.0 SE 2 mi.S.1 (310)54.8 SE
'11045 Susitna'I denI(77)
r5/22/81 16.8 N 2 mi.W.2(310)47.3 ESE
11 1530 Vermillion (345)
l (78)
r5/23/81 4.5 N Long Lake 2(310)45.3 E Traveling
"(345)from,I _new-
(88)born calfrmoosekill.
I
5/24/81 11.3 WSW 3-5 E 2(310)35.5 ESE On adultr:82O Tyone (345)moose kill
;1 (84)River 75%con-
sumed.-:1 5/25/81 2.0 S 4 mi.ESE 2(310)36.5 ESE
1I 1120 mouth (345)
(87)Tyone River
U
:i 5/26/81 12.8 SW 4.5 mi.NW 3(344)33.5 SE
Lone Butte (345)
Ir(89)(1 unc.)
j 5/27/81 12.5 SE 8 mi.W.2(310)46.0 SE On short
-Moore Lake (34)yr!.moose
f(93)k kill.
5/28/81 15.0 NE Near 3(310)49.5 SE
!"1832 Susitna (345)I (96)den (l unc.)J
r5/29/81 No Signal
I,
!
215
-----_.-_.
216
T Wolf -345 -Yearling gray male
Distance Distance
rJate-Time traveled Direction II Assoc.from den Direction
II.Obs.II (mi)of travel Location (ID)(mi)from den Misc.
f5/11/81 4.0 SW Tyone 0 38.2 SE
1205 Riverr56)
15/12/81 10.0 S Tyone 3(310)41.2 SE
ri 800 Ck.(344)
l (61)(1 unc.)
I
5/13/81 7.2 N Tyone 2(310)36.5 SE
r17~5 Ck.(344)
l (63)
r5/14/81 5.7 NW Susitna 2(310)31.0 SE On old
:,1 J855 Bend (344)adult moose....(64)kill.
05/17/81 9.5 E Upper 2(310)39.5 ESE Attacking
,1 1915 West (344)cow w/2
(72)yrl.
5/18/81 2.5 W Little 2(310)37.0 ESE
Gap (344)
Lake
5/22/81 10.8 E 2 mi.W.2(310)47.3 ESE
_1530 Vermillion (344)
,I (78)Lk.
5/23/81 4.5 N Long Lake 2(310)45.3 E Traveling
[(88)
(344)from new-
born calf
moose kill.
!r5/24/81 11.3 WSW 3-5 mi.E 2(310)35.5 ESE On adult
,1820 Tyone (344)moose kill
(84)River 75%con-
l sumed.
5/25/81 2.0 S 4 mi.ESE 2(310)36.5 ESE
i P20 mouth (344)
(87)Tyone
River
I~217
Wolf -345 -Yearling gray male (cont'd)
Distance Distance ~
Date-Time traveled Direction if Assoc.from den Direction
Obs.if (mi)of travel Location (In)(mi)from den Misc.
5/26/81 12.8 SW 4.5 mi.W.3(310)33.5 SE
Lone Butte (344)
(89)(1 nne.)
5/27/81 12.5 SE 8 mi.W.2(310)46.0 SE On short
Moore (344)yr!.moose
kill.
(93)kilL
5/28/81 15.0 NE Near 3(310)49.5 SE """',
1832 Susitna (344)
(96)(1 nne.)
!m!'i'1!:,
5/29/81 5.8 W 6.5 mi.0 44.8 SE
W.of Tyone
(94)Lake outlet g,FI,
5/30/81 19.3 NNW Near B-S 0 30.0 E
1436 Lakes
(106)
5/30/81 1.5 S 3 mi.SW 0 30.3 SE
1545 Susitna
(95)den
6/2/81 30.3 NW Watana 3(308)0
1930 den (323)
(11)(346)
6/3/81 2.8 N N.of den 1(325)2.8 N On caribou '"'"
1250 kill which
(98)324 was
obs.6/2.
6/4/81 2.8 S den 5(308)0
1945 (323)""'"(11)(324)
(325)
(340)
~
6/5/81 0 den 1(325)0 308
1925 traveling.
(11)~
218
219
220
-
i
I
"""'Ii
i
,....
j'
~i
I
i