HomeMy WebLinkAboutAPA4119Alaska Department of Fish and Game
Division of Gamer Federal Aid in Wildlife Restoration
Research Progress Report
LOWER SUSITNA VALLEY MOOSE POPULATION
IDENTITY AND MOVEMENT STUDY
by
Ronald D. Modafferi
Project W-22-5 and W-22-6
Job 1.38
June 1988
S79' v' ;W t
- I
STATE OF ALASKA
Steve Cowper, Governor P
5DEPARTMENT OF FISH AND GAME
Don W. Collinsworth, Commissioner
DIVISION OF GAME
W. Lewis Pamplin, Jr., Director
Steven R. Peterson, Research Chief
Persons intending to cite this material should obtain prior permis-
sion from the author(s) and/or the Alaska Department of Fish and
Game. Because most reports deal with preliminary results of conti-
nuing studies, conclusions are tentative and should be identified as
such. Due credit will be appreciated.
Additional copies of this report, or reports on other species covered
in this series may be obtained from:
Publications Technician
ADF&G, Game Division
P.O. Box 3-2000
Juneau, AK 99802
(907) 465-4190
PROGRESS REPORT (RESEARCH)
State: Alaska
Project No.: W-22-5 Project
W-22-6 Title: Big Game Investigations
Job No.: IB-1.38 Job Title: Lower Susitna Valley Moose
(2nd year) Population Identity and
Movement Study
Period Covered: 1 July 1985 -30 June 1987
SUMMARY
This report includes aerial-survey, marking, and radio-
relocating data collected from moose (Alces alces gigas
Miller) observed and/or cap iured between October 1985 and
May 1987 in alpine areas of the western foothills of the
Talkeetna Mountains and lowland forests of the lower Susitna
River Valley in Southcentral Alaska. This report also con-
tains pertinent survey and radio-relocation data gathered
between April 1980 and May 1986 during previous moose studies
in lowland riparian areas of the Susitna River Valley.
Periodic aerial surveys conducted from October through April
1985-86 and from November through March 1986-87 documented
increase, peak, and decrease phases in moose utilization of 7
subareas of alpine habitat in the western foothills of the
Talkeetna Mountains. Maxima of 919 and 1,405 moose were
observed in alpine habitats on 18 November 1985 and 26 Novem-
ber 1986, respectively. Moose numbers decreased earlier and
more dramatically in 1986-87. During both winters, moose
numbers in the Sunshine area continued to increase after the
areawide peak. Differences in peak numbers (53%) and dynamics
were attributed to variation in pcpulation size and/or clima-
tic conditions. Antlerless, calf, and antlered moose
accounted for 68%, 28%, and 4% of the increase, respectively.
Relative to antlerless moose, antlered moose decreased by 30%
(40:100 to 28:100) and calf moose increased by 44% (18:100 to
26:100) from 1985-86 to 1986-87. Changes within subpcpula-
tions on Bald Mountain and Willow Mountain, respectively, were
primarily responsible for a decrease in antlered and an
increase in calf moose.
After moose numbers declined in alpine habitats, about 150
moose were observed in lowland mature forests west of Willow
Mountain. About one-half of the moose observed occurred in
less than 23% of the area surveyed. Moose densities were
i
greatest between Little Willow Creek and the Kashwitna River.
Commercial timber harvest is proposed for this area. Timber
harvest will alter plant communities and may impact moose.
Herd size, composition, and mortality of moose were assessed
for 3 lowland riparian areas (Alexander, Moose, and Kroto
Creeks) during 3 winters. Alexander and Moose Creeks func-
tioned as typical moose winter ranges; as expected, these
winter ranges supported relatively large numbers of moose
though March and April, respectively. Moose use of Kroto
Creek, however, was different; it more closely paralleled
moose use of alpine postrut areas. Moose numbers on Kroto
Creek decreased greatly between February and March; herd
composition and winter mortality there also differed from
other areas, and more male and fewer calf moose were observed.
Significant winter-kill was also observed during all winters
on Kroto Creek. Hypotheses based on nutritive condition of
dams were posed to explain area differences in production
(calf numbers) , herd composition, and winter-kill. Data on
fetus sex ratios skewed towards males (2.4:l) were used to
implicate postrut range quality as an ultimate factor affec-
ting dam nutritive condition.
Moose mortality and movements were assessed from remains of
moose found dead and relocations of 44, 7, and 15 moose
radio-marked in alpine areas of the Talkeetna Mountains in
1985-86, in adjacent lowland forests in 1987, and along the
Susitna River floodplain in 1980-85, respectively.
Hunters killed 48% of the 25 marked moose that died. Death of
the remaining 13 moose was allocated between winter-kill
(16%), predation (12%), poaching (12%), injury/wounds (8%),
and collisions with trains (4%).
Mortality rate for some factors appeared to be sex dependent.
Forty-one percent of 22 marked males were killed by hunters.
Only 7% of 42 marked females were killed by hunters. Hunting
regulations favor the killing of male moose.
Death of 3 females and 1 male was attributed to winter-kill.
It was hypothesized that winters are more likely to weaken and
kill females (vs. males) because most are diverting part of
their energy and nutrient resources to grow feti.
Three females (and no males) were presumed killed by bears.
It was hypothesized that female moose (vs. males) are more
susceptible to bear predation because of their association
with neonates.
Data from radio-marked moose indicated that a major portion of
moose subpopulations that occur in alpine areas of Bald, Moss,
and Willow Mountains during the postrut period moved south to
lowland disclimax habitats near human settlements in winter
1986-87. In winter 1985-86, most marked moose remained at
higher elevations. Differences in movement patterns were
related to temperature and snowpack depth. Browse is abundant
in disclimax plant communities among human settlements.
During both winters, small numbers of moose moved west among
human settlements along railroad and highway rights-of-way and
near the Susitna River floodplain. Westerly, rather than
southerly, movements to winter range appeared more charac-
teristic for moose subpopulations north of Little Willow
Creek. Winter range near Pittmann and Wasilla, which was
commonly utilized by more southern subpopulations, may be too
distant.
Marked female moose throughout the Talkeetna Mountains
occurred west of the Susitna River during parturition. This
movement was more prevalent for moose north of Little Willow
Creek. It was hypothesized that lack of preferred calving
habitat and higher predator levels in the Talkeetna Mountains
foothills were responsible for this seasonal movement.
Most moose marked in the Talkeetna Mountains began aggregating
near timberline in June and July. Some moose moved higher
into alpine habitats in August. By early September, most
moose moved back into forested habitats at lower elevations.
The latter movement was associated with rutting activities.
However, influx of hunters into alpine habitat at about this
same time may also have prompted this movement. Marked moose
remained in midelevation forested habitat until late October
when movements into alpine postrut areas near initial capture
sites commenced. While in postrut areas, males commonly
occurred in sex-segregated groups at slightly higher eleva-
tions than females.
Potential impacts on moose of land development, timber har-
vest, and other alternative land uses proposed for the lower
Susitna River Valley were discussed. Concern was expressed
about potential impact of access that would be created inci-
dentally by the development.
Growth characteristics of browse in alpine and lowland habi-
tats were related to moose fall-winter movements and foraging
strategies. Hypotheses proposed suggested that (1) moose
should not be discouraged from overbrowsing lowland riparian
forage and (2) moose should not be encouraged to overbrowse
alpine forage.
Contemporary hypotheses on plant antiherbivory evolutionary
strategies suggest that fast, tall-growing plants on fertile
iii
lowland floodplain substrates probably provide more palatable
moose forage than do slower, lower-growing plants that occur
in less productive environments at higher elevations. These
strategies may partly explain why large numbers of moose move
to lowland floodplains in winter, rather than remain at higher
elevations. Future research plans are outlined.
Key Words: Moose, Alces alces gigas, Susitna Valley,
radiotelemetry, habitat, movements, aerial survey, population
identity, Southcentral Alaska.
iv
CONTENTS
SUMMARY. . .. . . .. . . .. . . .. . . .. . . .. . .
BACKGROUND .. .. . . . .. . . . . .. . . . .. . . . .
OBJECTIVES .. . . . .. . . . . .. . . . . .. . . . . .
Primary .. . . . . . . . .. . . . . . . .. . . . .
Peripheral. .. . . . . . . . . .. . . . . . . . . .
STUDY AREA . . .. . . .. . . . . .. . . . . . .. . . .
METHODS. . .. . ... . .. . . . . .. . . .. . .. ..
RESULTS AND DISCUSSION ... . . .. . . .. . . .. . . .
Dynamics of Moose Distribution, Abundances, and Herd
Composition in Alpine Habitats .. . .. . . ..
Moose Distribution and Herd Composition in Kashwitna
Forest . . .. . .. .. . . . .. .. . .. . .
Herd Size, Composition, and Mortality of Moose in
Lowland Riparian Wintering Areas .. . . .. . .
Dynamics in Use of Ranges. .. .. . .. . . ..
Herd Composition ... . .. . . .. . .. . ..
Winter Mortality . .. . .. . . .. . .. . ..
Fate of Radio-marked Moose. . . .. . . .. . . . ..
Moose Marked in This Study .. . .. . .. . ..
Moose Marked in Other Studies. . .. . . .. . .
Mortality Factors for Radio-marked Moose . .. . ..
Moose Movements . .... . . . .. . . .. . ... .
Bald Mountain ... . . .. .... . .. .
Moss Mountain ... . . . . . . .. . . . . . .
Willow Mountain. ... . . . .. . . ... ..
Witna Mountain . . ... . . .. . . .... ..
Brownie Mountain .. . . . .. . . . .. . . . .
Wolverine Mountain . . . . .. . . . . .. . . .
Sunshine Mountain. .. .. . . . .. . . . .. .
Kashwitna Forest .. .. . . . . . . . . .. . .
Study Area Summary .. ... . . .. . .. . ..
Habitat Considerations. .. . .. . . .. . . .. . .
Impact of Access on Moose Subpopulations. . .. .. .
Access Related to Hunting. ... .. .. . .. .
Access Unrelated to Hunting. .. .... . .. ..
Winter Forage and Foraging Strategies in Alpine
and Lowland Habitats . . .. .... .. . .. .
Management Implications. .. . .. . .. . .. .
i
2
4
4
4
5
5
8
8
10
12
13
14
15
17
17
18
18
20
20
21
21
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24
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25
27
FUTURE RESEARCH PLANS. . .. . .. . .. . .. . .. .. .30
ACKNOWLEDGEMENTS. .. . .. .. .. .. . .. .... .. .31
LITERATURE CITED .. . .. . .. . .. .. . .. . ...32
Figures. .. ... .. ..... . .. . .. . .. ...36
Tables .. .. ... ... . .. ... ...48
Appendix. Fate and capture data for radio-marked moose in
subareas of the lower Susitna River Valley in southcentral
Alaska, 1980-87. .. ... .. ... . .. . ......59
1
BACKGROUND
Prior to statehood, the Susitna River Valley was ranked as the
most productive moose (Alces alces gigas Miller) habitat in
the territory (Chatelain 1951). Today, the innate potential
of this area as habitat for moose is probably unsurpassed
throughout the state.
Presently, the lower Susitna Valley is the focal point of more
development than any other region in the state. Proposed and
progressing projects involving grain and crop agriculture,
dairy and grazing livestock, commercial forestry and logging,
personal-use cutting of firewood, mineral and coal mining,
land disposals, hydroelectric development, capitol site selec-
tions, wildlife ranges and refuges, human recreation, human
settlement, urban expansion, further development of the
highway system, and increased railroad traffic in the region
may greatly detract from the potential of the area to support
moose.
Though development and associated activities may tend to
decrease overall moose abundance in the Susitna Valley, there
is pressure from resource user groups to increase moose
populations so that their demand for greater direct alloca-
tions to commercial, consumptive, and nonconsumptive uses can
be satisfied.
The development activities and conflicting demands of resource
users have created a tremendous need for timely and accurate
general and site-specific knowledge about moose populations in
the lower Susitna River Valley (Game Management Units [GMU]
14A, 14B, 16A, 16B, and 13E). The demand for this information
originates from an array of local, state, and federal land and
resource management agencies, and it will likely intensify in
the future in response to (1) increased pressures to develop
additional lands, (2) increased numbers of users and types of
resource use, and (3) a more complex system for allocating the
resource to potential users.
Game Division presently lacks appropriate and/or sufficient
information about moose populations in the lower Susitna
Valley to accurately, consistently, and satisfactorily assess
ultimate impacts of contemporary demands on the moose
resource. The Division is therefore unable to knowledgeably
dispute or condone specific demands or provide recommendations
that would effectively regulate and minimize negative impacts
on moose populations or habitat. Additionally, the Division
must be knowledgeable about moose subpopulation behavior in
order to propose, design, and implement mitigation plans to
offset unavoidable negative impacts to moose subpopulations or
habitat.
2
Since major decisions on land use and resource allocation in
the lower Susitna River Valley are presently being made and
will continue to be made in the future, it is imperative that
the Game Division (1) proceed to review, unify, and summarize
the present state of knowledge on lower Susitna Valley moose
populations; and (2) proceed with new studies to augment this
data base so that future actions having an impact on moose
populations or their habitat may be promptly recognized,
evaluated, and minimized and/or mitigated. The lower Susitna
Valley is extensive in size and its habitats and environmental
conditions vary greatly. Because many resource use conflicts
require site-specific knowledge, numerous interrelated sub-
studies will be required to adequately understand movement
patterns and identities of major moose subpopulations.
Initial substudies should be conducted in areas where imme-
diate problems or conflicts in moose management exist.
When I evaluated conflicts in resource use for the entire
lower Susitna Valley, it was apparent that initial research
efforts should begin in the western foothills of the Talkeetna
Mountains (GMU's 14A and 14B) for the following reasons: (1)
this area possesses the largest, densest postrutting aggre-
gation of moose in the region and, perhaps, the state; (2) it
is the nucleus of development activities and resource use;
(3) it provides recreation and resources accessible to over
half of Alaska's human population; and (4) it has unique
problems involving railroad and highway systems. Also, recent
information obtained from Susitna River hydroelectric environ-
mental studies and a habitat suitability assessment project
has pointed out a lack of basic knowledge about moose in the
area.
Historical information available on moose populations in the
Susitna Valley is limited to (1) harvest statistics (ADF&G
files), (2) annual but inconsistently conducted sex-age
composition surveys (ADF&G files), (3) inconsistently collec-
ted data for train- and vehicle-killed moose (ADF&G files),
(4) an outdated population movement study based on resightings
of "visually collared" moose (ADF&G files), (5) studies on
railroad mortality and productivity of the railbelt subpopula-
tion (Rausch 1958, 1959), (6) a sporadically monitored radio-
telemetry population identity study in the Dutch and Peters
Hills (Didrickson and Taylor 1978), (7) a past, uncompleted
study of moose-snowfall relationships in the Susitna Valley,
and (8) a study of extensive moose mortality in a severe
winter (1970-71) for which there is no final report.
Recent studies designed to assess the impact of a proposed
hydroelectric project on moose have provided substantial
amounts of contemporary data on populations in areas adjacent
to the Susitna River and downstream from Devil Canyon
3
(Arneson 1981; Modafferi 1982, 1983, 1984). These data have
not been summarized to provide general or specific information
about those moose subpopulations when they are occupying areas
removed from the Susitna River floodplain. Circumstantial
evidence and cursory examination of these data suggest that
traditional sex-age composition counts conducted in widely
spaced alpine areas of GMU's 14A and 14B give biased results
and do not include samples from large segments of hunted moose
subpopulations. These data also suggest that moose killed
during late-winter hunting seasons in Subunit 14B originate in
Subunit 16A and that moose killed during hunting seasons in
Subunit 16A are included in composition surveys for Sub-
units 14A and 14B. I believe that moose subpopulations in
most of Subunit 16A (subpopulations that remain largely
unsurveyed because they occur in forested habitats) could be
surveyed during winter when they occur in riparian habitats
common to both Subunits 14B and 16A. The aforementioned data
and the fact that traditional composition surveys have
remained relatively insensitive to large annual changes in
moose mortality rates indicate that contemporary assumptions
about movements and identities of moose subpopulations in the
western foothills of the Talkeetna Mountains (Subunits 14A and
14B) are incorrect or, at least, too simplistic.
A recent joint study conducted by Divisions of Game and
Habitat (Modafferi and Albert, unpubl. data) and designed to
evaluate methods for assessing moose population status and
habitat suitability has begun to identify important moose
wintering areas and to document moose-snowfall relationships
in a large portion of the lower Susitna River Valley (GMU's
14A 14B, 16A, 16B, and 13E).
OBJECTIVES
Primary
To identify and delineate major moose subpopulations in the
lower Susitna River Valley.
To more precisely delineate annual movement patterns and
location, timing, and duration of seasonal habitat use.
To identify habitats and land areas that are important for
maintaining the integrity of moose subpopulations in the
lower Susitna River Valley.
Peripheral
To identify location of winter range and calving areas used by
lower Susitna River Valley moose subpopulations.
4
To determine natality rate and timing of calf and adult
mortality.
To assess effects of seasonal timing on results of sex-age
composition surveys on results obtained.
To identify moose subpopulations that sustain "accidental"
mortality on highway and railroad rights-of-way and mortality
from open hunting seasons.
STUDY AREA
The overall study area encompasses the lower Susitna River
Valley in Southcentral Alaska. This area includes all water-
sheds of the Susitna River south of Talkeetna (Fig. 1) and all
or portions of Subunits 14A, 14B, 16A, and 16B (Fig. 2).
Initial field studies were centered in alpine habitats along
the western foothills of the Talkeetna Mountains between the
Little Susitna River and the Talkeetna River (Subunits 14A and
14B). Moose were marked (Fig. 3) and aerial surveys (Fig. 4)
were conducted in these areas.
In late winter 1987, parallel field studies were initiated in
2 lowland moose wintering areas (see Figs. 3 and 4): (1)
forested habitats between the Kashwitna River and Willow Creek
(GMU 14B) where the Alaska Division of Forestry has proposed
to let leases for timber harvest and hunter access has greatly
increased and (2) riparian habitats along Alexander Creek (GMU
16B) where human settlement has escalated and late-winter
subsistence moose hunting seasons occur.
In subsequent years, similar field studies involving radio-
marked moose will be initiated to study subpopulations in
other geographical areas within the lower Susitna River
Valley. Data on moose herd composition were gathered on
aerial surveys of lowland riparian wintering areas in the
lower Susitna River Valley (Fig. 3).
METHODS
To identify and delineate moose subpopulations, to determine
annual movement patterns and timing, duration, and magnitude
of use of seasonal habitats, and to identify lands (habitats)
that are important to specific moose subpopulations, it was
necessary to periodically locate individually identifiable
moose. To provide individually identifiable moose that could
be periodically relocated, individuals were captured and
marked with ear tags as well as visual and radio-transmitting
neck collars. Each ear tag featured a discrete numeral, and
each neck collar featured a discrete, highly visible numeral
and radio-transmitted frequency.
5
For marking and capturing purposes, moose were located in
(1) an alpine substudy area along the western foothills of the
Talkeetna Mountains between the Little Susitna River and the
South Fork of Montana Creek (GMU 14A and B) during winter
1985-86, (2) a lowland forested substudy area between Willow
Creek and the Kashwitna River during late winter 1986-87, and
(3) a lowland riparian substudy area along Alexander Creek
during early spring 1987.
Moose were typically immobilized with 4-6 mg carfentanil
(Wildlife Laboratories, Ft. Collins, Co.), dissolved in 2-3 cc
H 0 and administered with Palmer Cap-Chur equipment by person-
nil aboard a hovering Bell-206B or Hughes-500D helicopter.
While immobilized, moose were marked with ear tags and neck
collars and aged by visual inspection of wear on incisor
teeth. Antler conformation was considered when assessing ages
of males. Though specific birth years were assigned to
captured moose, age categories of calf, yearling, 2-5, 6-12,
and 12+ years are more realistic because of imprecision in the
aging technique. Sex of marked moose and their association
with young of the year were noted. Immobilized moose were
revived with an intramuscular injection of 90 mg naloxone
hydrochloride (Wildlife Laboratories, Ft. Collins, Co.) per mg
of carfentanil administered.
Forty-four moose were captured and marked between 23 December
1986 and 4 February 1987 in the alpine substudy area
(Appendix A). Marking procedures were initiated after 18
November 1985, when peak numbers of moose were observed on
prior aerial surveys (Modafferi 1987). Distribution of
sampling effort between subareas within the alpine area
roughly paralleled moose distribution observed on aerial
surveys. Proportion of male moose marked was higher than that
observed on sex composition surveys, because male moose
usually dominate the open-hunting season harvest and more
complete information about their behavior (vs. females) was
desired.
Seven moose were captured and marked on 28 January 1987 in the
Kashwitna Forest substudy area. Sampling effort roughly
paralleled moose distribution observed in the area proposed
for timber harvest.
Fourteen and 6 moose were captured and marked in the Alexander
Creek area on 10 and 27 March 1987, respectively. Sampling
procedures roughly paralleled moose distribution previously
observed along the creekbed. Further details of research
activities in this substudy will be reported under a separate
job number.
6
Fifteen moose captured and radio-marked during previous
studies (Arneson 1980; Modafferi 1982, 1983, 1984) in the
lower Susitna River valley typically and frequently ranged
throughout GMU's 14A and 14B. Information gathered from these
individuals supplemented that obtained from moose that had
been specifically radio-marked for this study.
Survey flights in Cessna 180 or 185 and Piper PA-18 aircraft
equipped with a 2-element yagi antenna (Telonics, Mesa, Az.)
fixed on each wing were conducted periodically to relocate
marked moose. Moose relocations (audio-visual or audio) were
noted on USGS topographic maps (1/63,360 scale) during aerial
surveys. Relocation points were later transferred to translu-
cent overlays of those maps in preparation for computer
digitization and geoprocessing. Relocation surveys were
conducted at about 2-3 weeks intervals and provided about 22,
7, and 165 relocations through 27 May 1987 for moose marked in
the Talkeetna Mountains, Kashwitna Forest, and Susitna River
areas, respectively.
To determine moose distribution, abundance, and herd composi-
tion and to help delineate timing, magnitude, and duration of
use of habitats during winter, periodic surveys were conducted
in areas where moose were marked. Four, 2, and 2 aerial moose
surveys were conducted in the Talkeetna Mountains, Kashwitna
Forest, and Alexander Creek areas, respectively. Results of
these surveys were, in part, employed to determine when to
initiate moose marking procedures and how to distribute
sampling effort between subareas.
Information on moose herd composition and mortality were
collected during previous studies of lowland riparian winter-
ing ranges in the lower Susitna River Valley (Modafferi,
unpubl. data). Data gathered on surveys of Moose Creek, Kroto
Creek, Kroto Creek Islands, Yentna River, and Alexander Creek
are presented and analyzed to provide supplemental information
for assessing moose use of lowland riparian winter ranges.
Information gathered on occurrence and sex composition of
multiparous litters of moose killed during late-winter hunting
seasons (ADF&G files) was analyzed to help explain observed
differences in moose herd composition. Data on fate of marked
moose were analyzed to describe sources of mortality for moose
subpopulations in the lower Susitna River Valley.
For ease of reference and to denote hypothetical moose subpopu-
lations, 7 subareas were identified within the Talkeetna
Mountains alpine substudy area: Bald Mountain Ridge, Moss
Mountain, Willow Mountain, Witna Mountain, Brownie Mountain,
Wolverine Mountain, and Sunshine Mountain (Fig. 3). Subarea
names are those associated with Vertical Datum Bench Mark
(VDBM) notations on USGS topographic maps (1:250,000 scale).
7
RESULTS AND DISCUSSION
This report presents field data collected from April 1980
through May 1987. Data collected prior to October 1985 were
accumulated during previous moose studies in the Susitna River
Valley. Data collected from October 1985 through May 1987
were gathered specifically for this study.
Dynamics of Moose Distribution, Abundance, and Herd Composi-
tion in Alpine Habitats
Eight and 4 surveys were conducted to document distribution,
abundance, and herd composition changes of moose in 7 alpine
subareas in the western foothills of the Talkeetna Mountains
during the winters of 1985-86 and 1986-87, respectively (Table
1). During both winters, numbers of moose observed in alpine
areas fluctuated greatly and appeared to peak toward the
latter part of November. Peak number of moose observed in the
winter of 1986-87 was considerably greater than that observeed
in the previous winter. After the mid- to late-November peak,
the number of moose observed declined. Moose numbers
decreased more abruptly in the winter of 1986-87 than in the
winter of 1985-86. In both winters, numbers of moose observed
in the Sunshine Mountain subarea continued to increase past
the areawide late-November peak and significant decreases were
not recorded until until March.
In the winter of 1985-86, numbers of moose observed in alpine
areas peaked at 919 on 18 November and decreased to 202 by
April. In the winter of 1986-87, a peak of 1,405 and a low of
133 moose were observed on 26 November and 2 March, respec-
tively. The peak number of moose observed in the winter of
1986-87 was 53% greater than that recorded in the previous
winter. Moose numbers declined more abruptly in the winter of
1986-87, compared with that in the winter of 1985-86. In
1986-87, moose numbers declined by 77% from late November to
mid-January, compared with a 24% decrease from mid-November to
late February in 1985-86.
Temperature (Edwards and Ritcey 1956) and snow (Coady 1974)
may affect moose movements and behavior. Dynamics in numbers
of moose observed on aerial surveys in alpine habitats may be
related to climatic conditions. Moose numbers in alpine areas
may be negatively related to ambient temperatures and snowpack
depth.
October and November air temperatures were normal in 1985-86
(Clagett 1986:19) and above normal in 1986-87 (Clagett
1987:22). Relatively cold ambient temperatures in the winter
of 1985-86 were associated with relatively low numbers of
moose in alpine habitats, whereas above normal air
8
temperatures in the winter of 1986-87 were associated with
relatively large numbers of moose in alpine habitats. Ambient
temperatures could have directly affected moose behavior, or
more likely, ambient temperatures may have affected forage
quality that, in turn, influenced moose movement patterns.
Snowpack depth for February through April was below the 22-yr
recorded minimum in 1986 (Clagett 1986:29) and above the
historical average in 1987 (Clagett 1987:22). Shallow snow-
pack in the winter of 1985-86 probably did not discourage
moose from remaining in alpine habitats, whereas above-average
snowpack early in the winter of 1986-87 probably encouraged
moose to vacate those areas after November.
Another possibility is that moose behavior did not vary
between years, but that the size of moose populations in the
study area was larger in the winter of 1986-87 and, therefore,
led to observations of more moose. However, this contention
seems unlikely because of the relatively low proportion of
calves (potential recruits) observed in the population in
November 1985 (11 calves:100 moose or 18 calves:100 antlerless
[female] moose) (Table 2). If the increase in moose numbers
was attributable to recruitment, I would expect to observe a
similar significant increase in moose numbers in November 1987
(vs. 1986).
Antlerless moose accounted for 68% of the increase in moose
observed in November 1986 (Table 3). One hundred and thirty
percent (i.e., 135) more calves were observed. Nine percent
of the increase was attributable to antlered moose. The
proportion of antlered:100 antlerless moose observed decreased
(i.e., 40 vs. 28) between 1985 and 1986, whereas the propor-
tion of calves:100 antlerless moose increased (i.e., 18 vs.
26). Occurrence of antlerless adult moose and, to a lesser
extent, calves largely accounted for the increased numbers of
moose observed in November 1986; these herd characteristics
could have resulted from increased production and/or survival
of moose calves during 1986 or from drastic annual differences
in moose distribution.
Except for the Sunshine Mountain subarea, relationships of
moose numbers between areas remained relatively constant
during both winters. Contrary to this general data pattern,
during both winters the number of moose observed in the
Sunshine Mountain subarea continued to increase through
November and did not exhibit a significant decrease until
March. Moose use of this subarea differed from that of other
subareas.
When I combined data for subareas, the percentage of
antlerless moose observed differed little between years (63%
9
vs. 65%), whereas the percentage of antlerless and calf moose
decreased (25% vs. 18%) and increased (11% vs. 17%), respec-
tively, during 1985 and 1986 (Tables 2 and 3). Relative to
the antlerless moose observed, males decreased from 40:100 to
28:100 and calves increased from 18:100 to 26:100. The
greatest decrease in antlered moose (29%) occurred in the Bald
Mountain subarea. Large contributions to the overall increase
in antlerless moose occurred in Bald, Willow, Witna, and
Brownie Mountain subareas.
The most dramatic increase in the percentage of calf moose
occurred in the Willow Mountain subarea; in spite of a 66%
increase in number of antlerless moose, there was a 4-fold
increase in the proportion of calves:antlerless moose. Nearly
7 times as many calf moose were observed on Willow Mountain in
November 1986 than in November 1985. Causes of a parallel 70%
annual increase in calf moose observed on Bald Mountain may be
different because, in contrast, the calf:100 antlerless moose
ratio there changed only slightly between years (25:100
vs. 28:100).
The relative decrease in males and increase in calves are most
easily explained by an increase in the proportion of females
with calves in alpine habitats in November 1986. It is
generally assumed that in fall and early winter females with
calves more commonly occur in forested rather than alpine
habitats (ADF&G files). Possibly, annual variation in clima-
tic conditions (as previously described) was responsible for
the altering distribution of females with calves from forested
to alpine habitats in November 1986.
The 29% decrease in antlered moose (51% for the ratio of
antlered:100 antlerless moose) observed on Bald Mountain may
be attributed to the harvest during open hunting season. The
recent tremendous increase in ownership and use of ATV's by
hunters and the relative ease of access to Bald Mountain by
hunters lead me to believe that hunting contributed to the
decrease in antlered moose observed in November 1986.
Moose Distribution and Herd Composition in Kashwitna Forest
Because the Division of Forestry was planning to promote
timber harvests in the mixed forests adjacent to Willow
Mountain, I initiated a study to more fully understand how
moose utilize these habitats. As moose numbers observed in
alpine habitats on Willow Mountain decreased, substantial
numbers of moose were observed in adjacent midelevation
forested habitats. Aerial surveys conducted on 7 January and
6 February 1987 provided baseline information on distribution
and relative abundance of moose in this area (Table 4).
Because these surveys were primarily designed to ascertain
10
moose distribution, numerical totals are merely indices of
absolute numbers of moose. They are not meant to represent
the absolute number of moose present. I estimate that 30% of
the moose present were actually observed. This approximation
is realistic because about 500 moose were observed above
timberline on Willow Mountain in November and in January;
these observations occurred after moose numbers in this
habitat had decreased significantly and 151 moose (roughly
30%) had been observed on surveys in the adjacent forest
habitats.
By 6 February the number of moose observed in forested
habitats had decreased to 98. Results from this survey may
not be directly comparable to results from the 7 January
survey because of different observers and because snowcover
had deteriorated and proportionately fewer moose could be
observed. Regardless of possible shortcomings with surveys, I
suspect moose numbers in these forested habitats had
decreased. Other data also suggest that when snowpacks become
deep (e.g., in February), moose from the western foothills of
the Talkeetna Mountains move farther west (out of forested
habitats), presumably to forage near human settlements and
along the Susitna River floodplain (ADF&G files).
Data from two of 7 moose radio-marked in the Kashwitna Forest
area on 28 January 1987 indicated movements to other areas and
habitats by 24 February. One individual moved west to lower
elevations near the railroad right-of-way, and the other one
moved east to higher elevations. Both moose moved to loca-
tions where snowpack depth was less.
Substantial numbers of moose utilized the mixed-forest
habitats west of Willow Mountain and between Willow Creek and
the Kashwitna River. Timing and magnitude of moose use of the
area may be closely related to snowpack depth in surrounding
areas. Moose were not evenly distributed within the lowland
mixed-forested areas surveyed (Table 4); subsection f (Fig. 4)
encompassed roughly 23% of the survey area but contained 49%
and 50% of the moose observed on 7 January and 6 February
1987, respectively. The ecological basis for nonrandom moose
distribution within forested habitats remains unknown. If
moose movement out of alpine areas is primarily westerly
(i.e., little north-south movement), observed differences in
density may reflect local differences in subpopulation size.
More moose may move from areas nearer Little Willow Creek and
the Kashwitna River than from areas near Willow Creek and
Peters Creek. Local differences in moose subpopulation size
may result from differential hunter harvests. Southern
portions of Willow Mountain are more accessible to hunters and
have been more heavily hunted.
11
These data raise important questions regarding timber harvest
and moose management in mixed-forest habitats of the Kashwitna
forest. Moose in this area prefer to occupy either forest, or
the more typical shrub-dominated habitats during the winter.
If the former preference is correct, it would be wise moose
management to minimize timber harvests, at least, in subsec-
tion f of the Kashwitna Forest. If the latter preference is
correct, it would be wise moose management to encourage timber
harvests throughout all of the Kashwitna forest. Continued
studies will provide additional information so that the best
moose-related forest management practice can be determined.
The percentage of calves observed on 7 January 1987 in lowland
forest habitats adjacent to Willow Mountain was sightly less
than that observed on 26 November 1986 in alpine habitat on
Willow Mountain (20% vs. 17%, respectively) and similar to the
total observed for all subareas. Previous data (ADF&G files)
suggested that the percentage of calves in lowland forest
habitats would be greater than that in alpine habitats;
however, either the proportion of calf moose present in
forested habitats was not greater than that in alpine habitats
or a disproportionate calf mortality between areas or an
influx of adult moose occurred prior to 7 January. Data from
radio-marked individuals along with documented decreases in
numbers of moose observed in alpine habitats suggest that
there was a movement of adult moose into lowland forested
habitats. The continued decrease in the percentage of calves
observed in lowland forest habitats between the 7 January and
6 February surveys may be attributable to either calf mor-
tality or differential movement patterns between other sex or
age classes. Based on the aforementioned data, I suspect calf
ratios observed in forested habitats will continue to be
diluted with the influx of adult moose from alpine habitats.
Herd Size, Composition, and Mortality in Lower Susitna Valley
Riparian Wintering Areas
Status of moose populations is typically evaluated by aerial
composition surveys. Herd composition surveys are commonly
conducted in late November and early December when moose are
in postrut aggregations in alpine habitats.
Evaluating status of lowland moose subpopulations is difficult
because they typically occur in timbered habitats during the
postrut period when herd composition surveys are normally
conducted. Snow cover is frequently inadequate in lowland
areas, and mature mixed-forest canopies obstruct observability
of moose. However, later in winter, moose from lowland
subpopulations commonly depart forest habitat and gather in
relatively open-shrub habitats on riparian floodplains where
the status of these lowland herds .can be more determined. To
12
evaluate this possibility, one must understand the factors
affecting dynamics of moose herd composition on riparian
floodplains.
Moose population growth is largely determined by recruitment
that is assessed by enumerating calf and, to a lesser extent,
yearling proportions in the postrut aggregations (November or
December). Unfortunately, these assessments are conducted
before the winter period when calf and yearling moose cohorts
sustain considerable mortality. Late-winter herd composition
surveys would more accurately assess recruitment. Likewise,
winter mortality assessments should weigh heavily when formu-
lating management strategies for subsequent fall hunting
seasons.
Population simulation models help biologists understand the
population dynamics of moose. Year-round information on moose
mortality and herd composition provide the necessary data to
formulate realistic population simulation models. Information
on moose herd composition and mortality in winter is lacking
for the lower Susitna River Valley.
I collected data on dynamics, herd composition, and mortality
of moose on lower Susitna River valley riparian ranges in
winter 1983-83, 1984-85 (Modafferi, unpubl. data). Parallel
demographic data were gathered for the same areas in winter
1986-87 (Table 5).
Dynamics in Use of Ranges:
Data obtained from winter moose herd composition surveys
indicate that different riparian areas may serve different
ecological functions. Moose use (i.e., moose numbers) of the
Kroto Creek floodplain peaked in early December and appeared
to decline thereafter. Timing for this sequence of events was
similar to that observed for alpine areas in the western
foothills of the Talkeetna Mountains.
In contrast, moose use of the Moose and Alexander Creek
floodplains appeared to peak during January and February,
respectively. Moose use of the former areas remained high
until late March. Timing of moose use of these riparian
floodplains closely paralleled that observed for the Susitna
River floodplain (Modafferi 1984).
Moose and Alexander Creeks functioned as typical moose winter
ranges. Moose use of Kroto Creek more closely paralleled that
of the alpine postrut range.
Experimental studies indicate that while in alpine postrut
range moose consume large quantities of high-quality forage,
13
maintain a positive energy balance, and improve nutritive
condition (Schwartz et al. 1984); whereas, while on lowland
winter range moose consume substantially less forage, experi-
ence a negative energy balance, and lose nutritive condition
(Schwartz et al. 1984). The basic underlying difference in
moose ecology is that the energy balance is positive on the
early winter, postrut range and negative on the winter range.
Snowpack depth could be responsible for area differences in
moose use of lowland riparian winter ranges. The early
occurrence of deep snowpacks may have encouraged moose to
leave the Kroto Creek floodplain before moose had vacated
other floodplain winter ranges. Excessive snowpack depths in
midwinter may have prevented moose from leaving the Moose and
Alexander Creek floodplains for more desirable winter ranges.
Presently, I cannot explain the observed differences in moose
use of "apparent" lowland riparian winter ranges.
Herd Composition:
Composition of moose herd counts varied over winter and
between lowland riparian winter areas (Table 5). Variation in
moose herd composition may be attributed to antler drop in
males and age or sex differences in movement or mortality.
Male moose with "half-racks" (antlers on one-side only) have
been observed in early November. "Half-racks" were observed
on less than 2% of 281 males in mid-November. Most moose have
shed antlers by mid-January. Ratios of male and calf
moose:adult females were calculated from the 12 December 1985
survey (Table 6.).
Male moose were observed in all areas. Percentage of male
composition varied between areas. Kroto Creek exhibited the
highest percentage of males (22%) as well as the highest ratio
of males:100 adult females (37). The disparate sex ratios may
have resulted from differential movement patterns or produc-
tion and/or the greater survival of males using the Kroto
Creek drainage.
Fetus samples from pregnant moose killed by hunters in late
winter along the Parks Highway in the early 1980's indicate
that in utero sex ratios are skewed towards males (ADF&G
files). Nine of 13 (69%), 10 of 15 (67%), and 8 of 12 (67%)
feti examined in 1981, 1982, and 1983, respectively, were
males (Table 7). Seventy-two percent of the feti in 12
multiparous litters were males. Previous studies indicated
that in most years moose from the Kroto Creek area (GMU 16A)
move to areas along the Parks Highway by January and are
available to hunters during late winter. It is likely that
females sampled during late-winter hunts represented moose
14
from the Kroto Creek area. In contrast, samples from dams
killed by collisions with trains and vehicles in the
"Railbelt" of GMU 14B (between Matanuska and Curry) indicated
that fetal sex ratios were about equal in the mid 1950's
(Tables 7 and 8).
Differences in fetal sex ratios between time periods may be
accounted for by differences in subpopulation behavior,
nutritive condition, .or stage of population growth. Possibly,
behavior of subpopulations has changed between the mid-1950's
and early 1980's. Moose sampled in the same areas during
different time periods may have represented different subpopu-
lations with contrasting fetal sex ratios.
Fetal sex ratios for a given subpopulation can vary with
nutritive condition of dams that, in turn, may be related to
population status. During the earlier time period
(mid-1950's), moose populations in the Matanuska Valley were
at high levels, range conditions probably were of high
quality, and moose were generally well nourished. By the
1980's, moose population levels had declined, range quality
had deteriorated, and moose were not as well nourished.
Proportions of calves observed on moose herd composition
counts were lower on Kroto Creek (19%) than on Moose (25%) and
Alexander (23%) Creeks; even after accounting for the rela-
tively high proportion of males in the sample, the discrepancy
was still apparent. Disparate proportions of calves may have
been caused by (1) differential movement patterns or (2) lower
calf production and/or survival in the Kroto Creek drainage.
Calf composition on all 3 riparian floodplains decreased
through winter.
Winter Mortality:
The numbers of moose carcasses observed on floodplain areas
increased throughout the winter (Table 5). These losses,
primarily calves, became apparent in February and were attri-
buted to "winter kill." Although "winter kill" may result
from many causes, the ultimate common factors are low body fat
reserves and inadequate nutrition; it is also more likely to
occur when snowpacks are deep. The snowpack in the winter of
1984-85 was rated as the deepest in 10 years (Clagett et al.
1985) , and the "winter kill" was common to all areas during
that period.
For all areas, proportions of calves observed on surveys
generally decreased from approximately 20% in early December
to less than 10% in March. Moose carcasses were first
observed in February as calf composition approached 10%.
Carcasses were observed on Kroto Creek during the winters of
15
1984-85, 1985-86, and 1986-87. No carcasses were observed on
Moose Creek in March 1984 or on Alexander Creek in 1987.
Nineteen of 24 carcasses examined in these areas in the winter
of 1984-85 were calves (Modafferi unpubl. data).
Observations of moose carcasses imply that range quality on
all riparian areas was inadequate, relative to moose popula-
tion size and winter weather conditions in two of 3 years.
Based on observations of moose carcasses, range quality on
Kroto Creek was probably inadequate in all years. Alterna-
tively, lowland winter ranges were of adequate quality, but
the postrut ranges, where most moose improve nutritive condi-
tion prior to winter (Schwartz et al. 1984), were of inade-
quate quality, relative to moose population size.
The simplest explanation for the herd composition and winter
mortality data is a lightly hunted moose subpopulation. Light
hunting mortality levels led to excessive subpopulation levels
that caused overutilization and degradation of range quality
that, in turn, affected moose nutrition and resulted in high
winter mortality (Table 5) and low productivity (Table 6).
A less direct, alternative explanation that considers bull
ratios, winter mortality, and productivity but does not
require low male mortality is based on the nutritive condition
of dams. Studies of moose (Reuterwall 1981) and White-tailed
deer (Verme 1965, 1969; Verme and Ozoga 1981) provide evidence
that occurrence of unequal sex ratios in mammals is not
uncommon. These studies indicate that differential mortality
can lead to skewed adult sex ratios. Their data also indicate
that poor nutrition in dams, delayed mating, or delayed
fertilization may lead to male-dominated sex ratios at concep-
tion.
Even if a winter range is determined to be of high quality,
nutritional stress cannot be ruled out as the ultimate factor
causing "winter kill." "Winter kill" and high-quality winter
range are not mutually exclusive. Julander et al. (1961)
provided evidence that nutritional stress may originate long
before animals arrive on winter range. They linked low
productivity in mule deer to poor quality in the summer range.
Boisonnas (1935, in Verme 1981) provided circumstantial
evidence that delayed mating in red deer (Cervus elaphus)
resulted in hinds carrying a preponderance or male fetuses.
Delayed mating resulted either from (1) hunter harassment of
rutting animals and interference with rutting activities or
(2) relatively low proportions of breeding males to estrous
females. In the former case, moose were constantly moving to
avoid hunters instead of rutting, and in the latter case, too
16
few males were present to service all females during their
first estrus.
Accessible parts of the Kroto Creek area (along river and lake
shores) are heavily hunted; however, because large portions of
the area are relatively inaccessible, overall hunting activity
there has been light. Hunter harassment of moose and inter-
ference of rutting activities may occur locally but are
probably not prevalent throughout the area.
Relatively high proportions of antlered moose in the Kroto
Creek area during the winter suggest that males were lightly
hunted. Observations of "winter kill" indicate that moose
wintering on Kroto Creek were nutritionally stressed. The
fetal sex ratios (i.e., skewed towards males) and the low
proportions of calves are additional factors supporting the
nutritional stress hypothesis; these factors also indicate
that nutritional stress may occur before moose are on lowland
winter range. The high proportions of adult males observed on
Kroto Creek may have resulted from nutritional stress of dams
on the postrut range.
Previous studies in the area have provided data supporting the
correlation between fetal sex ratios, population status, and
range quality (Rausch 1959). In the early 1950's, when moose
populations in the area were increasing and range conditions
were likely better, fetal sex ratios were essentially equal.
Fate of Radio-marked Moose
Moose Marked In This Study:
Forty-four moose were radio-marked between 23 December 1985
and 28 January 1987 in the western foothills of the Talkeetna
Mountains. Seven additional moose were radio-marked on
28 January 1987 in the Kashwitna Forest area (Appendix).
Thirty-two radio-marked individuals were under surveillance on
27 May 1987. Seven of 19 males (37%) and one of 30 females
(3%) were killed during open hunting seasons. Two other
females (7%) whose transmitting collars have not been
recovered were suspected killed by hunters (perhaps ille-
gally). Three females were suspected killed by brown bears.
Death of 1 male and 1 female was attributed to "winter kill."
One male that died shortly after hunting season was suspected
to have died from a bullet wound.
One female (visual collar No. 481) died shortly after she had
been captured and handled. Although this mortality was
classed as captured related, the individual (estimated to be
18 years old) was extremely emaciated at capture. I suspect
17
capture trauma only precipitated an inevitable "winter kill"
mortality.
I killed 1 male that had been incapacitated by an infection in
the femur-tibia joint. Bone fragments in the infected area
indicated the infection had resulted from an injury. The
injury was probably the result of fighting during the rut, a
bullet wound, or a fall.
Moose Marked in Other Studies:
Fifteen moose that had been radio-marked between April 1980
and January 1984 for other studies in the lower Susitna River
Valley (Modafferi, unpubl. data) were found to utilize habi-
tats in the present study area (Appendix). Data obtained from
these individuals were included in the present study. Seven
of these moose were under surveillance through 27 May 1987.
Two of 3 males and one of 12 females were known to have been
killed by hunters during open hunting season. Another female
whose remains had been consumed by brown bears was suspected
to have been legally killed by hunters during open hunting
season. Death of 2 moose was attributed to "winter kill."
One female was killed by a collision with a train and 1 male
whose collar was found in the Talkeetna landfill shortly after
being after being located along the highway right-of-way was
suspected to have been killed illegally.
Mortality Factors for Radio-marked Moose
Twenty-five (39%) of the 64 radio-marked moose (Table 10) that
provided data for this study are dead. Mortality of radio-
marked moose was attributed to 6 factors (Table 9). Twelve
(48%) of the deaths were attributed to hunting, four (16%) to
"winter kill," three (12%) to predation, three (12%) to
poaching, two (8%) to injury or bullet wounds, and one (4%)
to a collision with a train.
Mortality rates for some factors varied between sex cate-
gories. Nine of 22 (41%) and three of (7%) 42 radio-marked
moose, respectively, were killed by hunters (Table 10).
Deaths of 3 (7%) females and 1 (5%) male was attributed to
"winter kill." Mortality of 3 (7%) radio-marked female moose
was attributed to bear predation.
One male and 2 female moose, representing 5% of the marked
moose, were thought to have been illegally killed (poached) by
hunters. Both female moose were poached during an open
hunting season (male only). The male was poached after the
season had closed. Mortality of 2 male and 1 female radio-
18
marked moose was attributed to injuries from wounds and a
collision with a train, respectively.
Hunting was a major source of mortality for male moose in the
study area (Table 10). Restrictive hunting regulations
regarding female moose concentrates the harvest on antlered
males. Regulations specifying harvest for only bull moose
essentially preclude harvest of calf moose. Subarea specific
data indicate that harvest of male moose was greatest in
subpopulations south of Willow Mountain (Bald and Moss Moun-
tain) and north of Brownie Mountain (Wolverine and Sunshine
Mountain) where easy access is afforded by all-terrain vehi-
cles (ATV's). Only one of 10 radio-marked moose on Willow
Mountain was killed by hunters. Hunter harvest rates in these
areas appear largely dependent on hunter access. I expect the
moose harvest in these areas to increase in relation to the
increasing ease of access caused by the use of ATV's. The
road proposed by Alaska Division of Forestry for timber
harvest will also provide easy access into forested portions
of Willow and Witna Mountain areas, and ATV's will provide
hunters with additional access to move freely throughout most
of the areas. Under present regulations, the increased access
will increase the moose harvest from these subpopulations.
"Winter kill" was the next most important mortality factor,
and its significance was greatly underestimated because it
primarily affects calf and yearling moose, age categories that
were not represented in the radio-marked samples. Addition-
ally, "winter kill" is not an important mortality factor when
average winter conditions prevail. The significance and
impact of "winter kill" can only be evaluated during severe
winter conditions (e.g., winters of 1970-71 and 1984-85).
Entire cohorts of calves can be lost during a severe winter
(ADF&G files). I suspect that "winter kill" would rate
significantly higher as a mortality factor if I were to
collect data during a severe winter. However, I doubt that
"winter kill" rates for Talkeetna Mountain moose subpopula-
tions would approach those for lowland moose subpopulations
west of the Susitna River where deeper snowpacks more
typically occur (Modafferi, unpubl. data).
Data available from other studies indicate that brown (Ballard
et al. 1980) and black bears (Ursus americanus) (Franzmann et
al. 1980) can be significant predators on moose calves.
Circumstantial data (Modaferri, unpubl. data) suggest that
bear predation on adult moose is both common and sex
dependent; bears may prey on females more commonly because of
their association with calves. These predators actively
pursue calf moose. Dams may have been killed while trying to
protect their calves; e.g., the death of 2 females attributed
to bears occurred during July, a time when neonates are
vulnerable.
19
I have obtained no evidence of predation on male moose.
Though brown bears on the Alaska Peninsula have been known to
attack and kill large bull moose during the rut (ADF&G files)
when they are probably less wary, I suspect the larger size of
male moose affords added protection from bears. Because male
moose are not accompanied calves they may be able to escape
pursuing bears. Data collected also imply that predation
rates on adult moose may be less in lowland than in alpine
habitats where brown bear occur more commonly.
Documentation of "injury-induced" moose mortality during the
winter period (ADF&G files) prompts me to speculate that those
mortalities occurring shortly after hunting season (and before
midwinter) may ultimately be attributed to infections from
injuries sustained from fighting during the rut, bullet
wounds, or falls, rather than "winter kill."
Moose poaching in the study area should not be regarded
lightly. Depressed economies, increases in rural inhabitants,
timing and duration of moose movements into settled areas, and
deep snowpacks that may affect moose temperament and daily
movements can affect rate of moose poaching; however, the
greatest factor influencing poaching rate is the tremendous
increase in human settlements in remote areas. Human settle-
ments typically occur in lowlands along watercourses, and
floodplains are typical wintering areas for moose.
Loss of moose to collisions with trains (and highway vehicles)
can be a significant mortality factor to specific subpopula-
tions (Modafferi unpubl. data). Rights-of-way for railroads
and highways are typically constructed in lowland areas along
major drainages. Common use of lowland areas creates a high
potential for train- and vehicle-induced mortality. Moose
mortality from this source increases with numbers of moose on
winter ranges (population size and movement patterns) and
length of time they utilize these areas. Mortality is grea-
test when deep snowpacks persist for extended periods and
moose occupation of winter ranges is lengthened.
Moose Movements
Summaries of moose movements by subarea of capture
(Figs. 5-12) illustrate annual, ranges for moose subpopulations
observed during typical ADF&G sex-age composition counts in
Subunits 14A and 14B.
Bald Mountain:
Movements for the Bald Mountain moose subpopulation were
basically bounded on the north by Willow Creek (Fig. 5).
Movements south occurred during midwinter (December) and
20
terminated near Pittman and Wasilla; by February 1987, all
radio-marked moose in this subarea had also moved south.
These data imply that roughly all 400 moose observed on Bald
Mountain (see Table 1) made similar movements to the win-
tering areas near Pittman and Wasilla. Shallow snowpacks and
availability of preferred browse apparently attracted moose to
this area. One 2-year-old male moved west over 100 km from
its capture site and was killed by a hunter near Hiline Lake
in September.
Moss Mountain:
In 1986-87 the only remaining radio-marked moose from the Moss
Mountain subarea moved south about 25 km to winter near
Wasilla (Fig. 3). This movement paralleled that for moose
from Bald Mountain. This behavior pattern probably charac-
terized movements for other moose in the Moss Mountain subpopu-
lation. The other radio-marked moose in this subarea was
killed by a hunter near Houston in September 1986.
Willow Mountain:
Five of 10 radio-marked moose on Willow Mountain that were
relocated during winter 1986-87 moved south about 35 km to
winter near Houston and Wasilla. These findings imply that
approximately 250 of the 500 moose composing the Willow
Mountain subpopulation made similar movements to this winter
range. At parturition time, two of 6 radio-marked females
were located west of the Susitna River near Kroto Creek.
These data imply that about 30% of the females observed on
willow Mountain moved to lowlands west of the Susitna River
for parturition. One radio-marked male moved west to winter
along highway and railroad rights-of-way near the Susitna
River at Montana. The later data suggest that only a small
percentage (10%) of the moose observed on Willow Mountain in
November moved west to winter along railroad and highway
rights-of-way near the Susitna River.
Witna Mountain:
The only radio-marked moose in this area wintered near highway
and railroad rights-of-way. This individual moved west of the
Susitna River during parturition. In August it returned to
alpine habitats of the Talkeetna Mountains. These data
suggest that females from this subarea winter near the Susitna
River and calve west of the Susitna River.
Brownie Mountain:
Radio-marked moose in this subarea generally remained near
higher elevations not far from their alpine capture sites.
21
For a short period of time after marking, several individuals
moved south across the Kashwitna River into the Witna subarea.
The following November, 1 female moved north into the
Wolverine subarea. One female ranged from the upper
Kashwitna River in late winter to the Susitna floodplain
during parturition, a linear distance of 60 km. Another
radio-marked female ranged south into the Willow Mountain
subarea.
Wolverine Mountain:
Two males accounted for major movements recorded for moose
marked in this subarea. One individual ranged from upper
Sheep Creek, Sheep River, and the North Fork of Kashwitna
River in summer to the Susitna River floodplain in winter.
Another male ranged from Sheep River in spring and summer to
near Caswell Lakes during the rut. During parturition, one
radio-marked female moved far up Sheep Creek in summer and
fall and another moved west of the Susitna River near the
Sunshine Bridge.
Sunshine Mountain:
Three radio-marked females that provided data for this area
moved across the Susitna River during parturition. One
individual traveled southwest about 70 km to a calving area.
These data suggest that preferred calving environments (i.e.,
those that may have fewer predators) are of limited availa-
bility to this subpopulation. Predators such as brown and
black bears and wolves (Canis lupas) become increasingly more
common in northern portions of the Talkeetna Mountains study
area. Moose from this subpopulation commonly moved west to
lowland wintering areas near human settlements along highway
and railroad rights-of-way.
Kashwitna Forest:
Shortly after capture, 2 marked females moved about 20 km
easterly to higher elevations in the Purches and Sheep Creek
drainages; they remained there until spring. Another female
moved west to winter in lowland areas along the highway and
railroad rights-of-way. In April a female moved west about 45
km across the Susitna River. The latter movement was presum-
ably for parturition.
Study Area Summary:
Large numbers of moose from Little Willow Creek winter in
lowland areas between Houston and Wasilla. Prevailing winds
in these areas keep snowpacks shallow, and human disturbance
to lands has resulted in revegetation by preferred moose
22
browse. Significant numbers of these moose are killed by
collisions with highway vehicles. Smaller numbers of moose
from this area move west to winter among human settlements and
along railroad and highway rights-of-way where human distur-
bance to lands has resulted in revegetation by preferred moose
browse. Moose from subpopulations near Kroto Creek and the
Susitna River also utilize this same winter range (Modafferi
1984). Moose begin to leave lowland wintering areas in March
or April and return to forested habitats near timberline in
the Talkeetna Mountain foothills. Females are in calving
areas by late May. Some travel to wet muskeg habitats across
the Susitna River, others remain in forested habitats slightly
below timberline. By late June or early July, many moose are
in timberline and alpine habitats. Moose remain there until
late August when they begin to move into more timbered habi-
tats for rutting activities. Moose begin gathering in alpine
areas in November after the rut. Moose numbers in postrut
alpine areas peak by about the last week of November and
decline thereafter as moose begin moving to lowland wintering
areas. Snowpack depth can influence timing of this movement.
In some aspects, moose behavior in subareas between Little
Willow Creek and the South Fork of Montana Creek differs from
that in more southern subareas. In inclement winters, migra-
tory moose from these subareas move west to lowland areas near
human settlements along railroad and highway rights-of-way and
the Susitna River floodplain. Significant mortality can
result from collisions with vehicles and trains. Greater
proportions of these more northern subpopulations appear to
remain in alpine areas and near timberline through winter.
Some moose from these same subpopulations winter in birch
forest habitats between timberline and the lowlands. In
inclement and/or long winters, many of the later moose
probably continue moving westward to destinations in lowland
areas. A greater proportion of female moose from these more
northern subareas calve in muskeg habitats west of the Susitna
River. This may be because of lack of preferred calving
habitat and/or high predator levels.
Habitat Considerations
Changing patterns of land use threaten to alter moose habitat
in the lower Susitna River Valley. Habitat alterations
associated with a ski resort in the Hatcher Pass area, land
management of the Willow Capitol site, agricultural and
homesite land disposals, development of industries based on
timber harvest, development of a coal mine and subsequent
plant succession, and development of lands previously cleared
for homesteads can impact local moose populations.
23
Timber harvest may not always be beneficial to moose; however,
it can have positive effects on moose when it causes regrowth
of preferred moose browse. Alternatively, timber harvest may
eliminate mature forest habitat components preferred by moose
for other reasons or during other seasonal periods. In these
instances and when winter range is not limiting, timber
harvest can be detrimental to maintenance and growth of moose
populations.
In many instances, mitigation for loss of moose or habitat is
appropriate. However, in cases where conflicts in land use
pertain to lands utilized by large numbers of moose or where
new or additional winter range will be of little benefit to
impacted subpopulations, alternative land uses should be
strongly opposed. Habitats important to large numbers of
moose (critical habitats) should be preserved.
Large portions of moose subpopulations in the lower Susitna
River Valley utilize wintering areas in disclimax habitats
among human settlements. Alternate uses of these private
lands would reduce available winter range and significantly
impact those subpopulations.
Impact of Access on Moose Subpopulations
Access Related to Hunting:
Use of ATV's increases the efficiency and success of hunters
over large land areas and will likely lead to shortened
hunting seasons. In areas where hunters utilize ATV's,
success of hunters without ATV's (foot-hunters) is probably
reduced. To maintain or increase length of hunting seasons
and more equally allocate the moose harvest among all hunters,
I suggest that some subareas be closed to the use of ATV's in
the taking of game.
The use of ATV's negatively impacts landscape. Presently, the
proportion of habitat destroyed by ATV's is relatively small;
however, small areas of affected habitats can negatively
impact the aesthetics of large landscapes. Because ATV's are
primarily used in remote areas by hunters in pursuit of game,
hunting regulations indirectly affect the area and timing of
that use. Wildlife managers must become concerned with impact
of ATV's on habitat quality as well as on moose population
levels.
Access Not Related to Hunting:
Human disturbance of moose during the postrut (when moose are
preparing for winter by regaining nutritive condition lost
during rut activities) and winter periods (when moose are
24
attempting to conserve energy) may negatively affect moose
energy bugets. Easy access into the postrut and winter ranges
of moose may encourage human use of these areas. Disturbances
from human activities (e.g., recreational snowmachining,
skiing, and photography, etc.) may alter normal activity
patterns of moose or displace them from their preferred
ranges. This unnecessary harassment during critical periods
may ultimately affect the survival of moose. Newly created
access into the Willow Mountain subarea, a popular snow-
machining area, may lead to human disturbance of that moose
subpopulation.
Winter Forage and Foraging Strategies in Alpine and Lowland
Habitats:
The winter forage of moose in lowland areas of the lower
Susitna River Valley primarily includes willows, birches, and
poplars in riparian or disclimax seral plant communities.
These plant species typically exhibit rapid growth rates that
in 5-10 years place the most palatable components out of reach
to moose. At this time, the plant may remain productive, but
moose are unable to utilize them. Regardless of the influence
of moose, these plant species can only be considered as
temporarily available food sources. Intensive browsing
pressure by moose may protract the period of availability of
plants browsed. Unbrowsed plants will continue to grow
rapidly. Degeneration (senescence) of browse plants resulting
from "overbrowsing" would be of little consequence in the long
term. In this situation, the strategy of moose would be to
utilize available browse heavily because it would only be
available for a short period of time.
Moose winter forage in alpine areas of the lower Susitna River
Valley consists primarily of willows, birches, and nonbrowse
species in climax communities. Growth rates of plants at
these higher elevations are probably slower than those at
lower elevations, and mature plants would be seldom out of
reach to moose. Plants in alpine communities are always
available to browsing by moose (i.e., they never grow out of
reach to moose and are not a temporary food source if not
covered by snow). Overbrowsing of plants in alpine commu-
nities could result in decreased long-term production and
encourage premature degenerative senescence. An evolutionary
strategy for moose exploiting browse in alpine plant commu-
nities would be to not heavily browse or "overbrowse" avail-
able plants. Overutilization of food sources available in
alpine communities would result in premature senescence or
degeneration of plants and lead to deterioration and permanent
loss of the range in the long term. It appears that over-
browsing on alpine ranges may have more profound and longer-
lasting impacts.
25
In terms of evolutionary strategies, moose may be encouraged
to heavily utilize available forage on lowland riparian winter
range and be discouraged from overutilizing available forage
on alpine postrut and winter range. It appears that over-
browsing on alpine range may have more profound long-term
affects on quantity of available browse.
Perhaps, frequent and periodic weather-related winter moose
die-offs and range overutilization observed on lowland ranges
function as a control of a subpopulation's size that, in turn,
deter overutilization of the more vulnerable alpine postrut
and winter range. Likewise, movements of the majority of
moose out of alpine postrut and winter ranges to lowland
winter ranges may be an evolutionary adaptation to guard
against overutilization of browse in alpine plant communities.
In contrast, plants have apparently evolved counter mechanisms
to discourage overutilization by herbivores. Recent studies
suggest that plant life form (MacLean and Jensen 1984), plant
growth rate and leaf life time (Coley 1988), and secondary
chemical compounds that inhibit forage digestion (Bryant and
Kuropat (1980) or intake (Robbins et al. 1987) are important
components of the antiherbivory defense mechanisms of plants.
Concepts basic to these hypotheses, may be summarized in the
following manner: The preferred evolutionary strategy of a
plant is to grow rapidly enough to escape browsing. This
antiherbivory strategy is probably common for plants growing
in fertile substrates (probably more so in lowland floodplains
than at higher alpine elevations). In less fertile substrates
where plants may grow more slowly and are accessible to
browsers for relatively longer periods, they have evolved
chemical or physical mechanisms (e.g., thorns) to discourage
herbivory. Recent hypotheses suggest that chemical components
that act by discouraging intake are more appropriate than
those compounds that inhibit digestion because the latter
substances may only encourage consumers to increase forage
intake (browse more!) to obtain required amounts of minerals
and nutrients.
Obviously, plants that never grow out of reach of herbivores
(i.e., shrubs, low growing subspecies, etc.) and/or remain
excessively attractive in winter (i.e., evergreen conifers and
ericaceous shrubs) have to rely on chemical and physical
mechanisms to discourage herbivory. Considering the antiher-
bivory strategies of plants, I would hypothesize that low,
slow-growing forage plants (which rely primarily on chemical
and physical deterrents to discourage herbivory) available to
moose at higher elevations in relatively unfertile substrates
in alpine habitats are not as "palatable" as the faster
growing, early successional forage plants (which rely pri-
marily on rapid grow rates to avoid herbivory) available in
26
relatively fertile lowland substrates along river floodplains.
This may explain why large numbers of moose seek winter range
in lowland floodplain areas along the Susitna and other
rivers.
However, these same scenarios lead one to raise the following
question: if plants in alpine habitats employ antiherbivory
"tactics" to discourage moose browsing, why do they attract
such large numbers of moose during the important foraging
period succeeding the rut and preceding winter? It may be
that plants only synthesize and/or translocate antiherbivory
chemical compounds into "browseable" shoots immediately
preceding winter senescence or, at least, after leaf
abscission. If defensive chemicals were incorporated into
"browseable" plant parts prior to leaf abscission or "down"
translocation of other substances into storage depots, plants
would have to synthesize much larger amounts of the compounds
to maintain relatively high (and functional) concentrations
of deterrents in "browseable" plant parts. Because production
of additional deterrent compounds would be "energetically"
more costly to plants, it is avoided by waiting until
"browseable" plant biomass is minimal.
Management Implications:
Traditional sex-age composition trend surveys are conducted in
early winter in alpine habitats where moose aggregate during
the postrut period. Data (Modaferri, unpubl. data) from
surveys in alpine habitats during the postrut period indicate
that (1) weather and timing of surveys can drastically affect
numbers and composition of moose observed, (2) annual varia-
tion in number of moose observed may be attributed to weather
conditions, rather than changes in subpopulation size, and
(3) annual variation (changes) in subpopulation demography may
be "masked" when data from several subpopulations are
combined.
Decreased numbers and percent of antlered moose on Bald
Mountain, an accessible and popular hunting area, most likely
resulted from loss of males killed during open hunting sea-
sons. Twenty-seven percent of the radio-marked adult male
moose in the Talkeetna Mountains were killed by hunters after
2 open hunting seasons. Three of 5 radio-marked adult males
on Bald Mountain were killed during the same time interval.
These data indicate relatively high hunter harvest rates for
males in GMU's 14A and 14B. Impact of these harvest rates on
moose management goals should be evaluated.
Higher moose calf ratios were observed on Willow Mountain in
winter 1986-87 than in winter 1985-86. Calf ratios on Bald
Mountain remained relatively unchanged between those years.
27
These data could be the result of open antlerless moose
hunting seasons in GMU 14B or annual differences in environ-
mental conditions. Elevated calf ratios could occur if
(1) hunters selected and killed lone antlerless moose rather
than antlerless moose accompanied by calves, (2) hunters did
not select for lone antlerless moose, and (3) orphaned calf
moose were not misclassified as adults during composition
surveys. An open antlerless hunting season occurred on Willow
Mountain (GMU 14B), whereas a limited number of permits were
issued for antlerless moose for Bald Mountain (GMU 14A).
Annual and subpopulation differences in calf ratios observed
on Willow and Bald Mountains may also be attributed to effects
of the relatively severe 1984-85 winter (excessive snow pack)
on nutritive condition of pregnant females, resulting in
lowered calf production and survival. While winter environ-
mental conditions could have depressed calf ratios in the
Willow Mountain subpopulation, they may not have adversely
affected calf ratios in subpopulations on Bald Mountain. The
Willow Mountain subpopulation primarily winters between
Kashwitna River and Willow Creek, and most of the Bald Moun-
tain subpopulation winters between Pittman and Palmer; exces-
sive snowpack depth seldom occurs there. If this scenario is
accurate and the productivity of the moose subpopulation on
Willow Mountain was depressed by winter conditions in 1984-85,
then improving winter range quality (carrying capacity) for
the Willow Mountain subpopulation would likely increase the
size of that subpopulation.
Annual differences in environmental conditions from fall
through winter could have altered distribution of calves in
the Willow Mountain moose subpopulation. If this was the
case, then the elevated calf composition observed in November
1986 (vs. 1985) was an artifact of nonrepresentative sampling.
In January 1987 substantial numbers of moose were observed in
forested habitats west of Willow Mountain. The Division of
Forestry is proposing timber harvests in this area. Previous
studies have indicated that moose also use nearby forested
habitats during spring and summer. In order to knowledgeably
comment on proposed timber harvests, the Game Division must
gather additional information to identify the reasons moose
occur in forested habitats.
Bull:cow ratios are frequently utilized to assess the impact
of hunter harvests on moose populations. Data from several
different sources indicated that elevated bull:cow ratios
observed in GMU 16A riparian wintering areas may be attributed
to effects of poor quality range on dams or skewed sex ratios.
Dams that have mated late or ones that have been in poor
nutritive condition during mating tend to produce more males
28
than females. In the absence of population estimates, ratios
should never be solely used to assess status of moose popula-
tions.
Because proportions of yearling males in fall moose popula-
tions are utilized to estimate the annual increment of
breeding females, it is extremely important to know fetus sex
ratios. If fetus sex ratios are skewed 2:1 toward males, then
observations of 6% yearling males would perhaps suggest an
equal number of yearling females rather than a higher number.
In male-only hunting areas, it is typically assumed that
survival rates for yearling females is considerably higher
than for males. Current data on fetus sex ratios should be
obtained where hunting of females occurs.
Moose mortality on lowland riparian winter ranges in GMU 16A
must be quantified annually; this factor must be considered
when calculating allowable hunting harvest. Additional
studies should be conducted to determine cause of mortality in
lowland riparian wintering areas.
Movements of radio-marked moose indicate that postrut aggre-
gations in GMU 14B represent numerous, relatively discrete
subpopulations. Losses (mortality) and gains (productivity)
to a specific moose subpopulation only affect that subpopula-
tion. These findings indicate that moose in GMU 14B should
be managed at the subpopulation level.
Moose from several subpopulations may utilize the same winter
range. Areas among human settlements south of the Little
Susitna River and north of the Parks Highway between Houston
and Palmer are an important moose winter range. Negative
impacts to this winter range through development or advances
in plant succession will affect large numbers of moose from
numerous subpopulations. I estimate that about 1,000 moose
presently utilize this winter range. Loss of moose winter
range in this area should be avoided. If range loss is
unavoidable, alternative winter ranges should be established.
In winter, moose frequently become a nuisance to humans when
they occur among human settlements and interfere with human
activities. Additionally, because highway and railroad
rights-of-way occur within moose winter ranges, large numbers
of moose are killed by collisions with trains and highway
vehicles. Information on movement patterns of moose in the
lower Susitna River Valley may be utilized to designate areas
where moose winter range could be established to help preclude
mortality and conflicts with human activities.
Use of ATV's and snowmachines have become popular recreational
endeavors in unsettled areas of the lower Susitna River
29
Valley. Snowmachine and ATV use may alter habitats, displace
moose from seasonal ranges, and/or cause them to alter usual
behavior patterns. Disturbances from human recreational
activities may directly or indirectly result in mortality of
moose. Disturbance of moose by snowmachines in winter on
winter range and by ATV's on fall range (during rut) are areas
of concern. Wildlife managers must become cognizant of the
potential impact of these human activities on moose.
Use of ATV's by hunters in GMU 14B has increased substantially
in the last several years. In many areas, ATV trails are
obvious features of the landscape. ATV use in wet marshy
areas alters those habitats. Wildlife managers must recognize
that hunting regulations may indirectly affect habitat
quality.
Data on fate of radio-marked moose in the lower Susitna River
Valley indicate that the following component categories should
be considered when calculating annual mortality (losses) for a
moose subpopulation: (1) hunter harvest, (2) crippling kill,
(3) illegal harvest, (4) predator kill, (5) collisions with
trains, (6) collisions with highway vehicles, (7) winter kill
(range carrying capacity, population size, and winter severity
all relevant factors), (8) old age, (9) injuries from fighting
in rut (males), (10) injuries from accidents (falling or
slipping on ice), (11) drowning (after falling through ice or
open water leads in rivers, while crossing rivers, or getting
caught in ice jams during breakup).
Population demography and movements of radio-marked moose
indicate that a distinct subpopulation occurs on Bald Moun-
tain. This subpopulation sustains substantial mortality from
collisions with vehicles and trains and poaching while on
winter range in lowland areas between Houston and Palmer.
Mortality from these sources is less common to other GMU 14A
moose subpopulations east of Government Peak and more similar
to GMU 14B subpopulations north of Bald Mountain. The Bald
Mountain moose subpopulation exhibits levels of productivity
higher than subpopulations on Willow Mountain, presumably
because of higher quality winter range. These data indicate
that the Bald Mountain moose subpopulation may require more
site-specific management than presently exists under the
GMU 14A classification.
FUTURE RESEARCH PLANS
1. Periodically Continue radio-relocating marked moose.
2. Conduct herd distribution, abundance, and composition
surveys as snow cover permits through the 1987-88 winter.
30
3. Conduct surveys in alpine areas of Talkeetna Mountains,
on Alexander Creek, and on the Yentna River near the new
proposed Skwentna and McDougall study areas.
Skwentna and McDougall, on the Yentna River, were selec-
ted as component areas for extension of moose research in
the lower Susitna River Valley. These riparian habitats
are commonly known to be important moose wintering areas.
Extensive use of these habitats by moose was documented
during previous winter surveys (Modafferi, unpubl. data).
Late winter subsistence hunting seasons occur in these
areas. Similar to the Alexander Creek substudy area,
this area represents a wintering habitat (lowland
riparian) grossly different from alpine areas in the
western foothills of the Talkeetna Mountains.
4. Conduct field excursions into Kashwitna forest subarea to
assess moose winter use of that habitat.
5. Initiate a cooperative study with U. S. Fish and Wildlife
Service involving marking moose with satellite-tracking
collars.
Satellite-tracking collars provide information on acti-
vity and movement of marked moose. Potentially discern-
ible activities include feeding, running, and bedding.
These data may provide baseline information to quantita-
tively evaluate the influence of human disturbances on
moose winter behavior. Satellite-tracking collars
provide daily movement data regardless of weather condi-
tions. Satellite-tracking collars have a 1-year useful
life expectancy. Because satellite collars must be
attached "snugly" to obtain activity data, they are only
installed on females. Two females in the Willow Mountain
subarea will be marked with satellite-tracked collars.
6. Radio-mark additional moose in the Willow Mountain
subarea near where timber harvests are proposed.
Additional moose may also be radio-marked in the Bald
Mountain subarea because of the reduced sample size in
that area and to provide additional information on moose
use of the Willow Capitol Site Land Management Area.
ACKNOWLEDGEMENTS
I am especially grateful to Dennis C. McAllister, Alaska Dept.
Fish and Game, who made useful suggestions, pertinent criti-
cisms, and willfully provided his assistance throughout this
study. Dennis always managed to be available when needed;
sometimes at the expense of his own time. Dennis was
31
invaluable in field aspects of this study. I am thankful to
Dennis for his able assistance is all facets of this study.
I am grateful to my immediate supervisor Karl Schneider,
Alaska Dept. Fish and Game, who has always diligently suppor-
ted and lobbied for the study of moose in the lower Susitna
Valley. Karl's sincere interest in the study has been a
welcomed stimulus to my efforts. Karl has also labored to
keep my administrative duties to a minimum so I might concen-
trate on the biological problems. I greatfully appreciate all
his efforts.
The following persons also deserve special thanks:
C. Soloy, Soloy Heli-Ops, Wasilla, for expertly piloting
helicopters during moose capture procedures; B. Wiederkehr,
Wiederkehr Air Inc. Palmer, for piloting Piper Super Cub
aircraft during herd distribution, abundance and composition
surveys, and aerial surveys to locate radio-marked moose; and
Larry Rogers, Southcentral Air, Kenai, for piloting Cessna
180/185 on moose radio-relocation surveys. These individuals
are commended for piloting aircraft safely and efficiently
during low-level flight operations. B. Taylor and J.
Didrickson are recognized for assistance in field aspects of
this study. J. Didrickson and N. Steen, area game management
biologists responsible for GMU's 14A and 14B (the study area),
are acknowledged for freely providing local knowledge, logis-
tic assistance, a congenial working atmosphere, and "area
support" from conception through the present stage of this
study. L. Pank, U. S. Fish and Wildlife Service, is acknow-
ledged for stimulating my interest in the application of
satellite telemetry on moose and willingly cooperating in a
joint-agency study.
LITERATURE CITED
Arneson, P. 1981. Big game studies. Vol. II. Moose. Ann.
Prog. Rep. Susitna Hydroelectric Proj. Alaska Dept.
Fish and Game. Juneau. 64pp.
Ballard, W. B., T. H. Spraker, and K. P. Taylor. 1980.
Causes of neonatal moose calf mortality in southcentral
Alaska. J. Wildl. Manage. 45:335-342.
Boisonnas, J. 1935. In Verme L. J. and J. J. Ozaga. 1981.
L'appel des cerfs. Review de Paris. 21:130-147.
Bryant, J. P. and P. J. Kuropat. 1980. Selection of winter
forage by subarctic browsing vertebrates: the role of
plant chemistry. Ann. Rev. Ecol. and Systemat.
11:261-285.
32
Chatelain, E. F. 1951. Winter range problems of moose in the
moose in the Susitna Valley. Proc. Alaska Sci. Conf.
2:343-347.
Clagett, G. P. 1986. Alaska snow surveys. U.S. Dept. of
Agriculture, Soil Conservation Service. Anchorage.
30pp.
Clagett, G. P. 1987. Alaska snow surveys. U.S. Dept. of
Agriculture, Soil Conservation Service. Anchorage.
3 1 pp.
Clagett, G. P., R. McClure, and T. Robles. 1985. Snow
surveys and water supply outlook for Alaska. U.S. Dept.
of Agriculture, Soil Conservation Service. Anchorage.
Coady, J. W. 1974. Influence of snow on behavior of moose.
Naturaliste can. 417-436.
Coley, P. D. 1988. Effects of plant growth rate and leaf
lifetime on the amount and type of antiherbivore defense.
Oecol. 74:531-536.
Des Meules, P. 1964. The influence of snow on the behavior
of moose. Trans. NE. Wildl. Conf., 21. 11 Figs., 17pp.
Didrickson, J. C. and K. P. Taylor. 1978. Lower Susitna
Valley moose population identity study. Alaska Dept. of
Fish and Game. Fed. Aid Wildl. Rest. Proj. Final Rept.,
W-17-8 and 9. Jobl.16R. Juneau. 20pp.
Edwards, R. Y. and R. W. Ritcey. 1956. The migrations of a
moose herd. J. Mammal. 37:486-494.
Franzmann, A. W., C. C. Schwartz, and R. O. Peterson. 1980.
Moose calf mortality in summer on the Kenai Peninsula,
Alaska. J. Wildl. Manage. 44:764-768.
Julander, 0., W. L. Robinette, and D. A. Jones. 1961.
Relation of summer range condition to mule deer herd
productivity. J. Wildl. Manage. 25:54-60.
MacLean, S. F. and T. S. Jensen. 1985. Food plant selection
by insects in Alaska arctic tundra the role of plant life
form. Oikos 44:211-221.
Modafferi, R. D. 1982. Big game studies. Vol II. Moose-
Downstream. Final Phase I Rep. Susitna Hydroelectric
Proj. Alaska Dep. Fish and Game. Juneau. 114pp.
33
.1983. Big game studies. Vol. II.
Moose-Downstream. Prog. Rep. Phase II. Susitna Hydrol-
electric Proj. Alaska Dep. Fish and Game. Juneau.
114pp.
.1984. Big game studies. Vol. II.
Moose-Downstream. Prog. Rep. Phase II. Susitna Hydro-
electric Proj. Alaska Dep. Fish and Game. 116pp.
1987. Lower Susitna Valley moose population
identity and movement study. Alaska Dept. Fish and Game.
Fed. Aid Wildl. Rest. Final Rep. Proj. W-22-5. Job
1.38R. Juneau. 17pp.
Rausch, R. A. 1958. The problem of railroad-moose conflicts
in the Susitna Valley. Alaska Dept. of Fish and Game.
Fed. Aid Wildl. Rest. Final Rep. Proj. W-3-R. Job 1-4.
Juneau. 116pp.
Rausch, R. A. 1959. Some aspects of population dynamics of
the railbelt moose populations, Alaska. M.S. Theses.
Univ. of Alaska, Fairbanks. 81pp.
Robbins, C. T.. 1959. Some aspects of population dynamics of
the railbelt moose populations, Alaska. M.S. Thesis.
Univ. Alaska, Fairbanks. 81pp.
, T. A. Hanley, A. E. Hagerman, O. Hjeljord, D. L.
Baker, C. C. Schwartz, and W. W. Mautz. 1987. Role of
tannins in defending plants against ruminants: reduction
in protein availability.
Reuterwall, C. 1981. Temporal and spatial variability of the
calf sex ratio in Scandinavian moose Alces alces. Oikos
37:39-45.
Schwartz, C. C., W. L. Regelin, and A. W. Franzmann. 1984.
Seasonal dynamics of food intake in moose. Alces
20:223-242.
Verme, L. J. 1965. Reproduction studies on penned
white-tailed deer. J. Wildl. Manage. 29:74-79.
____. 1969. Reproductive patterns of white-tailed deer
related to nutritional plane. J. Wildl. Manage.
33:881-887.
and J. J. Ozaga. 1981. Sex ratio of white-tailed
deer and the estrous cycle. J. Wildl. Manage.
45:710-715.
34
PREPARED BY:
Ronald D. Modaferri
Game Biologist III
APPROVE BY:
W. L p in, ir., Director
Si en R. Peterson
Chief of Research
35
j
Fogure 1. Map showing location of the study area In Alaska with names listed for
rivers, lakes and other prominent landscape features.
36
Fig. 2. Location of game Mansgement Subunits (13E, 14A, 148,
I6A and 161) and state and national parks In the study area.
37
NOTH
NORTH
llia
Fig. 3. Location of Talkeetna Mountains subareas (A-G), Kashwltna Forest (H)
and Alexander Creek (I) where moose were radio-marked in Winter 1986-86,
January 1987 and March 1988, respectively and the Yentna River (J) where
moose will be radio-marked In March 1988.
38
r
---- -- ---
goOk In et
• IN I IIII Il l ln N in
Fig. 4. Locationsi for TalkeetflS Mountains subareas (A-0). Kashwltns Forest (1H)
subareas (a-b). Moose Creek (1). Kroto, Croek (A) Yentne River (K) and Alexander
Creek (L) where moos* surveys were conducted (Asulaid Mtn..1, SMosg Mtn.,
C-Willow Mtn.. DaWitna Mtn.. E.=Brownie Mtn., F*Woiverine. Mtn., and G=Sunshine Mtn.).
39
Fig. 6. Polygon encompassing relocations for moose-radio marked In the Bald Mtn.
subarea (G).
40
i
Fig. 6. Polygon encompassing relocations for moose radio-marked In the Moss Mtn.
subarea (4).
41
Fig. 7. Polygon encompassing relocatlons for moose radio-marked In the Willow Mtn.
subarea (C).
42
Fig. 8. Polygon encompassing relocations or moose radio-marked In the Witna Mtn.
subarea (C).
43
_··
I
Fig. 9. Polygon encompassing relocations for moose radio-marked In the Brownie Mtn.
subarea (@).
44
Fig. 10. Polygon encompassing relocatlons for moose radio-marked In the Wolverine Mtn.
subarea (4).
45
I _
C
Fig. 11. Polygon encompassing relocations for moose radio-marked In the Sunshine Mtns.
subarea (0).
46
t
NORTH
Scale 1:915000
0
Km.
50
Fig. 12. Polygon encompassing relocations for moose radio-marked In the Kashwitna
Forest subarea (4).
47
--·
Table 1. Number of moose observed on periodic surveys in alpine areas of
the Talkeetna Mountains foothills in southcentral Alaska during 2
winters, 1985-87.
Winter
1985-86 1986-87
Areaa 4 17 8 18 3 23 31 17 26 24 15 2
Oct Oct Nov Nov Dec Feb Mar Apr Nov Dec Jan Mar
A 37 109 264 302 260 275 191 40 408 120 47 20
B 0 19 37 50 54 33 26 15 38 45 11 9
C 5 148 262 268 313 164 121 59 492 43 15 15
D 0 9 24 19 20 42 13 11 101 6 9 26
E 0 25 110 125 112 104 96 49 197 61 41 30
F 0 41 54 129 93 32 22 14 148 18 8 14
G 0 2 21 26 39 50 21 14 21 56 51 19
Total 42 353 775 919 890 703 487 202 1,405 349 181 133
a Areas A, B, C, D, E, F, and G -Bald Mountain, Moss Mountain,
Willow Mountain, Witna Mountain, Brownie Mountain, Wolverine Mountain,
and Sunshine Mountain, respectively.
48
F
Table 2. Herd composition for moose observed in alpine subareas of the
Talkeetna Mountains foothills in southcentral Alaska, 18 November 1985.
Moose Ata Al C At: C:
Subarea No. % N No. No. % No. % 100 Al 100 Al
Bald Mountain 302 33 66 22 189 63 47 16 35 25
Moss Mountain 50 5 7 14 35 70 8 16 20 23
Willow Mountain 268 29 81 30 173 65 14 5 47 8
Witna Mountain 19 2 6 32 8 42 5 26 75 63
Brownie Mountain 125 14 41 33 71 57 13 10 58 18
Wolverine Mountain 129 14 23 18 90 70 16 12 26 18
Sunshine Mountain 26 3 8 31 17 65 1 4 47 6
Total or Avg. 919 100 232 25 583 63 104 11 40 18
a At -antlered moose, Al -antlerless adult moose and C -calf moose.
49
Table 3. Herd composition for moose observed in alpine subareas of the
Talkeetna Mountains foothills in southcentral Alaska, 26 November 1986.
Moose Ata Al C At: C:
Subarea No. % No. 2 No. % No. % 100 Al 100 Al
Bald Mountain 408 29 47 12 281 69 80 20 17 28
Moss Mountain 38 3 10 26 25 66 3 8 40 12
Willow Mountain 492 35 108 22 287 58 97 20 38 34
Witna Mountain 101 7 25 25 63 62 13 13 40 21
Brownie Mountain 197 14 40 20 138 70 19 10 29 14
Wolverine Mountain 148 11 22 15 104 70 22 15 21 21
Sunshine Mountain 21 1 2 10 14 67 5 24 14 36
Total or Avg. 1,405 100 254 18 912 65 239 17 28 26
a At -antlered moose, Al -antlerless adult moose and C -calf moose.
50
4
*L
Table 4. Herd composition for moose observed in subsections of the
Kashwitna forest survey area in southcentral Alaska, 7 January and 6
February 1987.
Survey Number of Mooseb
date Subsectiona At Al C Total %C
7 Jan a 0 8 0 8 0
b 0 18 4 22 18
c 1 15 4 20 20
d 0 16 4 20 20
e 0 6 1 7 14
f 6 55 13 74 18
Total 7 118 26 151 17
6 Feb a 0 7 4 11 36
b 0 5 0 5 0
c 0 14 1 15 7
d 0 8 0 8 0
e 0 8 2 10 20
f 0 44 5 49 10
Total 0 86 12 98 12
a Subsections a,
10, 13, 10, 10, 11,
b, c, d, e, and f were estimated to encompass
and 16 mi2, respectively.
b At -antlered moose, Al -antlerless adult moose and
C * calf moose.
51
Table 5. Moose herd composition (At -antlered adults, Al -antlerless
adults, Cf -calves, and carcasses -Cs) observed on aerial surveys of
Moose Creek, Kroto Creek, Kroto Creek Islands, and Yentna River areas,
1984-1987.
Moose
At Al Cf Cs
Location Date No. % No. % No. % No.
0
5
11
13
8
0
0
0
0
0
0
0
0
0
0
46
56
21
5
0
0
01
0
0
0
0
0
0
5
29
12
28
11
7
20
5
9
20
28
4
16
18
5
29
12
27
11
7
20
5
9
20
28
4
16
18
20-
28
4
0
16
14
11
6
0
0
0
0
0
0
0
0
0
0
32
22
12
3
0
0
1
0
0
0
0
0
0
Mar
Nov
Dec
Dec
Jan
Feb
Feb
Mar
Mar
Mar
Mar
Apr
Apr
Mar
Mar
Nov
Dec
Dec
Jan
Feb
Feb
Mar
Mar
Mar
Mar
Apr
Apr
Mar
Mar
Mar
Apr
58
18
50
70
114
123
162
155
143
106
69
62
41
47
37
71
150
124
141
127
134
81
61
36
29
18
11
34
84
84
84
84
85
85
85
85
85
85
85
85
85
87
84
84
84
84
85
85
85
85
85
85
85
85
85
87
85
85
85
0 0 133
87
56
62
61
83
84
90
92
91
91
99
93
93
89
93
50
59
70
80
88
89
90
95
97
100
95
92
89
89
94
87
9
9
20
32
16
24
19
14
15
11
1
5
3
6
3
25
48
32
30
17
17
8
3
1
0
1
1
4
5
2
4
13
28
25
28
12
16
10
8
9
9
1
7
7
11
7
18
19
18
17
12
11
9
5
3
0
5
8
11
11
6
13
92 11 8
52
Moose
Kroto Ck.
Kroto Is.
Yentna
0
0
0
0
0
0
39
29
27
0
0
0
0
0
0
1
0
2
3
13
10
18
12
13
0
0
0
0
0
4
4
2
1
2
9
6
9
0
3
1
922 Feb 85
1 11
Table 5. (cont'd)
Moose
At Al Cf Cs
Location Date No. % No. % No. % No.
Alexander 29 Nov 84 12 23 27 51 14 26 0
12 Dec 84 17 15 68 62 25 23 0
27 Dec 84 11 9 94 79 14 12 0
11 Jan 85 3 1 191 78 52 21 0
7 Feb 85 0 0 172 86 28 14 2
20 Feb 85 0 0 149 90 16 10 3
5 Mar 85 0 0 186 88 26 12 0
9 Mar 85 0 0 170 90 18 10 1
20 Mar 85 1 1 142 91 13 8 3
28 Mar 85 0 0 134 94 8 6 7
4 Apr 85 0 0 151 94 9 6 6
16 Apr 85 0 0 124 92 11 8 6
18 Feb 87 0 0 125 85 22 15 0
12 Mar 87 0 0 85 85 15 15 0
53
Table 6. Herd composition (At -antlered, Al -antlerless adults, and C -
calves) for moose observed on aerial surveys of Moose, Kroto, and Alexander
Creeks, 12 December 1985.
Moose Ata Al C At: C:
Subarea No. % No. % No. % No. % 100 Al 100 Al
Moose 81 11 14 50 62 20 25 22 40
Kroto 254 56 22 150 59 48 19 37 32
Alexander 110 17 15 68 62 25 23 25 37
a At -Antlered moose, Al -antlerless adult moose and C -calf moose.
54
i0
Table 7. Sex ratio of feti from dams killed by hunters and by collisions with
trains or vehicles in Game Management Subunit 14B during the winters of 1955-58
and 1980-83.
Collision killb Hunter killc
1955 1956 1957 Total or 1980 1981 1982 Total or
Sexa -56 -57 -58 Average -81 -82 -83 Average
M 6 20 4 30 9 10 12 31
F 7 19 3 29 4 5 4 13
M:1 F 0.9 1.1 1.3 1.0 2.3 2.0 3.0 2.4
a SM -malesbData from
Data fromcData from
and F -females.
Rausch 1959, Table 5 p.20-22.
ADF&G files.
55
Table 8. Sex of feti in multiparous litters of moose killed by collisions with
trains and vehicles and hunters in Game Management Subunit 14B in the winters
1956-58 and 1980-83.
Collision killb Hunter killc
1955 1956 1957 Total or 1980 1981 1982 Total or
Sex -56 -57 -58 Average -81 -82 -83 Average
FF 0 3 0 3 0 0 1 1
FM 1 1 1 3 2 1 1 4
MM 0 2 0 2 2 2 2 6
MMF 0 0 0 0 0 1 Q 1
TM 1 5 1 7 6 7 5 18
TF 1 7 1 9 2 2 3 7
M:1 F 1.0 0.7 1.0 0.8 3.0 3.5 1.7 2.6
a aFF
female,
* 2 females, FM -1 female and 1 male, MM
TM -total males, TF -total females, and
S2 males, MMF - 2 males and 1
M:1F -males per 1 female.
b Data from Rausch 1959, Table 5 p.20-22.
SData from ADF&G files.Data from ADF&G files.
56
Table 9. Mortality factors for radio-marked moose in the Talkeetna Mountains
(N-49) and along the Susitna River floodplain (N-15).
Study
Mortality Talkeetna to Susitna River Total
Factors No. % No. X No. %
Hunter kill 8 47 4 50 12 48
Winter kill 2 12 2 25 4 16
Predation 3 18 0 0 3 12
Injury/wounding 2 12 0 0 2 8
Poaching 2 12 1 13 3 12
Collision with train 0 0 1 13 1 4
Total 17 101 8 101 25 100
aIn cases where actual cause of death was uncertain, individuals were
assigned to the most probable cause after considering circumstances.
57
Table 10. Capture location, sex composition, and fate of radio-marked moose in
subareas of the lower Susitna River Valley in southcentral Alaska, 1980-87.
Malesa Females Total
Subarea Mk Hk OK Mk Hk OK MK Hk Off OK
Bald Mountain 5 3 2 5(1) 0 3 10 3 5 5
Moss Mountain 1 1 0 1 0 0 2 1 2 0
Willow Mountain 6 1 4 8 0 6 14 1 4 10
Witna Mountain 0 0 0 1 0 1 1 0 0 1
Brownie Mountain 3 0 2 4 0 4 7 0 1 6
Wolverine Mountain 2 1 0 3(1) 1 2 5 2 3 2
Sunshine Mountain 2 1 1 1 0 1 3 1 1 2
Kashwitna Forest 0 0 0 7 0 6 7 0 1 6
All subareas 19 7 9 30(2) 1 23 49 8 17 32
Susitna Riverb 3 2 0 12 2 7 15 4 8 7
Total 22 9 9 42(2) 3 30 64 12 25 39
a Mk, Hk, Ok, and Off -No. moose marked, killed by hunters, presently under
surveillance, and no longer under surveillance, respectively. Numbers in
parentheses indicate moose that were captured and marked but subsequently died
as a result of capture procedures. These moose are excluded from other totals.
b Radio-marked moose in parallel studies and which provided supplemental
information for this study.
58
(BL,
APPENDIX
Table A. Fate and capture data for radio-marked moose in subareas of the lower
Susitna River Valley in southcentral Alaska, 1980-87.
Number
Capture Capture Left Right Visual Trans-
date location Sex Agea ear tag ear tag collar mitter Status
12/23/85 Bald Mt. M 3 2354 1546 31 18135 HK
12/23/85 Bald Mt. F 8 2360 1506 262 18136 CM
12/23/85 Bald Mt. F 8 2389 2388 33 18130 OK
12/23/85 Bald Mt. F 5 2400 2395 27 10591 OK
12/23/85 Bald Mt. F 5 1510 2485 29 10598 OK
12/23/85 Bald Mt. F 2 2392 2399 371 6359 WK
12/23/85 Bald Mt. M 1 2397 2396 35 18131 HK
12/23/85 Willow Mt. F 10 1551 1524 34 18137 OK
12/23/85 Willow Mt. F 6 1575 1570 36 18138 OK
12/23/85 Willow Mt. F 4 2482 2440 3 6397 PK
12/23/85 Willow Mt. F 4 2433 2368 9 6396 PK
12/23/85 Willow Mt. M 8 1568 1569 8 6383 WM
12/23/85 Willow Mt. M 4 2394 2391 32 18134 OK
12/23/85 Willow Mt. M 5 2398 2393 5 6374 OK
12/26/85 Willow Mt. M 6 1571 2497 28 6425 OK
12/26/85 Bald Mt. F 3 2357 1512 2 12807 IK
12/26/85 Bald Mt. M 12 1573 1574 1 10498 OK
12/26/85 Bald Mt. M 9 1501 1554 4 6372 OK
12/26/85 Bald Mt. M 4 1504 2390 7 6356 HK
12/26/85 Moss Mt. F 15 1538 1513 38 10498 IK
12/26/85 Moss Mt. M 4 1517 1532 25 6438 HK
01/02/86 Brownie Mt. F 7 2387 2382 49 6460 OK
01/02/86 Brownie Mt. F 4 2380 2386 50 6495 OK
01/02/86 Brownie Mt. M 3 2378 2385 54 6499 OK
01/02/86 Brownie Mt. M 4 2381 2383 52 6454 WK
01/02/86 Brownie Mt. M 3 2379 2384 53 6504 OK
01/02/86 Witna Mt. F 5 1508 1503 55 6402 OK
01/02/86 Wolverine Mt. F 9 2376 2377 51 6496 OK
01/07/86 Brownie Mt. F 3 1562 2414 43 10496 OK
01/07/86 Brownie Mt. M 8 1528 2409 30 6411 OK
01/07/86 Sunshine Mt. F 3 1511 1555 11 6410 OK
01/07/86 Sunshine Mt. M 5 1560 1561 47 6500 OK
01/07/86 Sunshine Mt. M 3 2411 2479 10 6494 HK
01/07/86 Wolverine Mt. F 18 1586 2436 481 6501 CM
01/07/86 Wolverine Mt. F 12 2423 2370 46 18133 HK
01/07/86 Wolverine Mt. M 5 1505 1509 441 10594 HK
02/04/86 Wolverine Mt. M 7 1698 2158 48 6501 IM
02/04/86 Wolverine Mt. F 13 2073 2150 581 23933 OK
02/04/86 Willow Mt. F 8 2071 2106 721 6458 OK
02/04/86 Willow Mt. F 4 2161 2116 60 6457 OK
59
Table A. Cont.
Number
Capture Capture Left Right Visual Trans-
date location Sex Age ear tag ear tag collar mitter Status
02/04/86 Willow Mt. M 7 2162 2190 16 6365 HK
02/04/86 Willow Mt. M 11 2200 2059 17 6380 OK
02/04/86 Willow Mt. F 3 2156 1652 61 6517 OK
02/04/86 Willow Mt. F 3 2101 2142 261 18136 OK
01/28/87 Kashwitna For. F 8 42 50 21 26100 PK
01/28/87 Kashwitna For. F 3 39 43 731 26101 OK
01/28/87 Kashwitna For. F 1 62 44 24 26104 OK
01/28/87 Kashwitna For. F 6 69 67 25 26106 OK
01/28/87 Kashwitna For. F 18 80 45 15 26107 OK
01/28/87 Kashwitna For. F 7 --- 23 26109 OK
01/28/87 Kashwitna For. F 4 66 38 14 26110 OK
04/17/80 Susitna River F 2 15753 15752 26 10603 OK
04/17/80 Susitna River F 5 15754 15755 22 10592 OK
04/17/80 Susitna River F 2 15737 15378 23 10594 RK
04/17/80 Susitna River M 2 15371 15370 27 6388 HK
03/10/81 Susitna River F 11 8442 -- 79 6502 HK
03/10/81 Susitna River M 5 -- 8453 84 6460 HK
02/24/82 Susitna River F 6 16984 -- 94 10597 OK
01/31/84 Susitna River F 4 16 -- 812 6424 OK
02/24/82 Susitna River F 3 16704 -- 100 10602 OK
02/24/82 Susitna River F 2 16998 -- 41 6494 HK
01/03/85 Susitna River F 9 2149 2076 2 13128 OK
02/24/82 Susitna River F 7 -- 16937 87 10593 WK
03/10/81 Susitna River M 10 8477 -- 65 6413 IK
01/31/84 Susitna River F 7 6 17 61 6459 OK
02/26/82 Susitna River F 11 -- -39 10596 WK
a Age determined from incisor wear; assigned age probably encompassed within
intervals of: 1, 2-3, 4-6, 7-12, and 12+ years.
b OK -alive and functional; HK -hunter kill; WK -winter kill; CM -capture/drug
related mortality; PK -predator kill, these are not documented but presumed to be
most likely cause of death; IK -illegal kill; WM = hunting/wound mortality and IM =
mortality from injury/wounding. Date for all OK -05/27/87.
60
4
S