MOOSE MOVEMENT PATTERNS IN THE UPPER KOYUKUK RIVER
DRAINAGE, NORTHCENTRAL ALASKA

Kyle Joly1, Timothy Craig2,4, Mathew S. Sorum1, Jennifer S. McMillan3, and
Michael A. Spindler2

1National Park Service, Gates of the Arctic National Park and Preserve, 4175 Geist Road, Fairbanks,
Alaska, 99709; 2US Fish and Wildlife Service, Kanuti National Wildlife Refuge, 101 12th Avenue,
Fairbanks, Alaska, 99701; 3Bureau of Land Management, Central Yukon Field Office, 1150 University
Avenue, Fairbanks, Alaska, 99709; 4Retired

ABSTRACT: Understanding movement patterns of moose (Alces alces) is critical to understanding
their ecology and sound management. Our study was prompted by concern that the Dalton Highway
Corridor Management Area (DHCMA), where the Dalton Highway facilitates access for non-local
hunting, may be a population sink for moose that also reside in more remote and protected areas
like Gates of the Arctic National Park and Preserve (GAAR) and Kanuti National Wildlife Refuge
(KNWR). We did not detect substantial migrations between DHCMA and GAAR or KNWR. However,
we estimated that 14–60% of moose in our study area were migratory depending on sex, location within
our study area, and methodology utilized to differentiate migratory behavior. A quarter of the animals
displayed mixed-migratory strategies where migration is exhibited by a single individual in some years
but not others. The percentage of moose that were migratory in our study population, and the distances
they migrated, were lower than reported from studies elsewhere in interior Alaska. We hypothesize this
may be related to their very low density (∼ 0.1 moose/km2) and/or higher terrain ruggedness in part of the
study area. Winter severity did not appear to impact migration, but home range sizes were smaller in
severe winters.

ALCES VOL. 51: 87–96 (2015)

Key words: Alaska, Alces alces, conservation, migration, moose, strategy, winter severity

Moose (Alces alces) across their circum-
polar range often exhibit migratory patterns
(e.g., van Ballenberghe 1977, Ball et al.
2001, White et al. 2014), but migration is
not a ubiquitous trait among all populations
(Mueller et al. 2011) and they are not widely
perceived as migratory by the general public.
Migration is thought to increase fitness
because animals gain access to better quality
or quantity of forage, experience a reduction
in predation, or possibly reduced exposure to
parasites or disease (van Ballenberghe 1977,
Avgar et al. 2013). Variability in migration
among individuals may be the norm where
habitat conditions sustain resident popula-
tions but fitness varies spatially (Fryxell
and Holt 2013).

Movements of moose in the upper
Koyukuk River drainage (Fig. 1) are not
well understood. On the lower Koyukuk
River, an area where moose density is com-
paratively high (5/km2), 83% of adult moose
and 58% of cow-calf pairs were migratory,
and movement between summer and winter
ranges averaged 42 and 31 km, respectively
(Osborne and Spindler 1993). In the Arctic
National Wildlife Refuge and neighboring
Canada, northeast of our study area and
where moose density is much lower
(∼0.4/km2; Caikoski 2010) than in the lower
Koyukuk study area, 88% of moose were
classified as migratory (Mauer 1998). There,
the mean maximum migration distance was
123 km (i.e., the mean longest migration

87



distance by individual moose). Moose den-
sity is even lower in the upper Koyukuk sec-
tion of our study area (∼0.1/km2; Lawler
et al. 2006).

Land and wildlife management in our
study area falls within a complicated patch-
work of authorities and includes lands ad-
ministered by the State of Alaska, National
Park Service, US Fish and Wildlife Service,
Bureau of Land Management, and private

entities. Public wildlife advisory groups
interested in hunting opportunities and
management in the upper Koyukuk River
drainage were concerned that harvest was
high within and near the Dalton Highway
Corridor Management Area (DHCMA)
which contains the only road in northern
Interior Alaska (Fig. 1). Harvest opportu-
nity was open to all Alaskan residents in
the DHCMA, whereas only local residents

Fig. 1. The Upper Koyukuk River study area (white polygon) in northcentral
Alaska, encompassing moose locations derived from GPS and VHF
telemetry data from 2008–2013.

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harvest moose within Gates of the Arctic
National Park and Preserve (GAAR) and
much of the Kanuti National Wildlife Refuge
(KNWR). Subsistence hunters in the upper
Koyukuk were concerned that moose were
migrating long distances, as in the eastern
Brooks Range (Mauer 1998), from their
exclusively subsistence hunting areas into
general harvest areas, causing a local decline
in density and hunting opportunity. The pos-
sibility that hunter effort and success in one
portion of the region was negatively affect-
ing another area where distinct user groups
hunted was an important management con-
cern. Therefore, our goal was to better under-
stand the migratory patterns of moose in the
upper Koyukuk River drainage, and we were
specifically interested to identify if moose
moved between lands with different manage-
ment goals and hunting regimes.

METHODS
Study area

Our study area falls within the upper
Koyukuk River drainage in north-central
Alaska (Fig. 1), and encompassed the south-
ern flanks of the central Brooks Range,
including the southeastern portion of
GAAR, all of KNWR, and other state, fed-
eral, private, and native lands. It included
portions of the DHCMA which contains an
all-weather highway and had special hunting
restrictions. The terrain and vegetation com-
munities are diverse. Rugged mountains to
2000 m in elevation with narrowly-confined
glacial river valleys are covered with a mix
of alpine, shrub, and boreal forest habitat
types in the northern portion. Shrub habitats
were dominated by alders (Alnus spp.), wil-
lows (Salix spp.), and dwarf birch (Betula
glandulosa). Black spruce (Picea mariana)
is the most prevalent tree species, with white
spruce (Picea glauca) and poplar (Populus
balsmifera) common in riparian areas, and
birch stands (Betula papyrifera) occur on
south-facing slopes and in areas that burned.

Extensive tracts of tussock (Eriophorum
spp.) tundra occur in wetter, flatter areas.
The landscape becomes progressively flatter
(elevations typically <500 m) to the south
with more muskegs, streams, and lakes inter-
spersed within boreal forest and broad riparian
zones. The regional climate is strongly conti-
nental, with long, extremely cold (dropping
below -45o C) winters, and brief, but hot
(temperatures >30o C) summers. Forest fires
are common during summer months, and
snow depth exceeds 90 cm many winters
with 60 cm during most.

Moose relocation data and GIS analyses
The project ran between March 2008

and April 2013. Adult moose were darted
using a mixture of carfentanil citrate and
xylazine from Robison R-44 helicopters.
Moose were instrumented with either a
GPS collar equipped with a VHF beacon or
a standard VHF collar. Moose captured north
and east of Bettles, Alaska (Fig. 1) were
designated as ‘northern moose’ and those in
and around KNWR as ‘southern moose’.
GPS collars deployed in March 2008 col-
lected 1 GPS location/day, thereafter, all col-
lected 3 locations/day. An attempt to relocate
moose visually from small aircraft using the
VHF beacons occurred ∼monthly from
March 2008 to April 2013. These efforts
were much more consistent for the southern
moose due to budgetary restraints, weather,
and logistics. Movements of northern and
southern moose were compared due to dif-
ferences in terrain, habitats, and sampling
effort.

Migratory status was assessed using
2 different methods using individual moose
as the sample unit for both techniques. First,
for both GPS- and VHF-collared moose,
we calculated distances between summer
(i.e., the closest relocation July 1) and winter
(i.e., the closest relocation to January 15)
locations to assess migratory status and
distance moved. Moose that consistently

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demonstrated separation between summer
and winter locations were categorized as
migratory (i.e., winter locations were not
located sympatrically with summer locations
or vice versa), whereas those without consis-
tent separation between ranges were categor-
ized as non-migratory. Distance between
summer and winter ranges was not a deter-
minate. Moose that had separation in some
years but not others, or did not have a dis-
cernible pattern, were categorized as having
a mixed-migratory strategy. Distances
between winter and summer locations were
averaged for moose that had multiple years
of data. While somewhat subjective, this
methodology mimics other studies (e.g.,
Osborne and Spindler 1993, Mauer 1998)
that were conducted adjacent to ours, thereby
allowing more direct comparison.

Second, net squared displacement
(NSD) of GPS-collared moose was used to
determine migratory status on an annual
basis (see Bunnefeld et al. 2011). The NSD
measures straight line distances between a
starting location and all successive reloca-
tions within the entire annual movement
path. The first position was set to July 1
when moose are in their summer range, and
we excluded all individuals sampled <330
days. Moose displaying “home range”
movement patterns according to Bunnefeld’s
nomenclature were categorized as non-
migratory, “migration” as migratory, and
“mixed migratory” and “dispersal” as mixed
migratory (no moose dispersed from the
study area). Moose that did not consistently
have the same movement status among years
were considered mixed migratory. The shape
of movement patterns, not NSD distance,
was the determinate of migratory status
(Bunnefeld et al. 2011). There were insuffi-
cient VHF data to quantify migration pat-
terns with the NSD technique for any
individual moose.

The 2 methods for determining migra-
tory status are highly disparate and caution

is required when comparing results. GPS
and VHF locations were also used to deter-
mine if individual moose utilized conserva-
tion units (GAAR, KNWR), the DHCMA
exclusively, or multiple units.

Winter severity was classified from the
total number of days with snow and snow
depth as recorded in Bettles, Alaska. The
categories were mild (<135 days with ≥30 cm
snow or <7 days with ≥60 cm snow), moderate
(>170 days with ≥30 cm snow, >50 days with
≥60 cm, or <14 days with ≥90 cm snow),
or severe (>170 days with ≥30 cm snow,
>100 days with ≥60 cm, or >30 days with
≥90 cm snow).

Annual and winter home ranges for GPS-
collared moose were determined by establish-
ing fixed kernels using the 95% utilization
distribution and least square cross validation
method for smoothing using ArcGIS (ESRI,
Redlands, CA; Worton 1989, Seaman and
Powell 1996). There were insufficient data
to develop kernels for VHF collars.

RESULTS
Moose Relocation Data

In total we captured 120 adult moose
(27 bulls and 93 cows); 58 in March 2008,
10 in October 2008, 15 in November 2009,
and 37 in April 2011. One presumed capture
myopathy (adult cow in 2008) was censured.
There were 52 southern moose and 67 northern
moose providing ∼265 moose-years of data to
analyze allopatric migration and movements
among conservation units. Aerial telemetry
flights yielded 2119 high-quality relocations
(positive visual identification). GPS collars
were deployed on 14 northern cow moose
in 2008 (25 moose-years of data), 2 in 2009
(2 moose-years), and 2 in 2011 (2 moose-
years); 8 GPS collars were deployed on
southern cows (8 moose-years of data) and
11 on northern bulls in 2011 (18 moose-
years). In total, the 37 GPS units collected
71,675 locations.

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Migration
Allopatry of winter and summer ranges

and the migratory status of 86 cows and
21 bulls were ascertained through the use
of both VHF and GPS data. Of the 10 moose
with only a single year of data, 2 displayed
non-migratory behavior and 8 displayed
mixed-migratory behavior. Using only GPS
data for NSD-based models, we determined

the migratory status of 20 cows and 11
bulls; 25 had 2–4 years of data (5 non-
migratory, 8 migratory, and 12 mixed)
and 6 had only a single year of data
(4 non-migratory, 1 migratory, and 1 mixed).
Non-migratory, migratory and mixed-
migratory strategies were exhibited in varying
proportions by sex, location, and methodology
(Fig. 2).

0

10

20

30

40

50

60

Non-migratory Migratory Mixed-Migratory

All Moose - Allopatry
All Moose - NSD
All Cows - Allopatry
All Cows - NSD
Southern Cows - Allopatry
Southern Cows - NSD
Northern Cows - Allopatry
Northern Cows - NSD
All Bulls - Allopatry
All Bulls - NSD
Southern Bulls - Allopatry
Southern Bulls - NSD
Northern Bulls - Allopatry
Northern Bulls - NSD

Pe
rc
en
ta
ge

Fig. 2. Percentages of non-migratory, migratory, and mixed-migratory moose in the upper Koyukuk
River drainage, northcentral Alaska, 2008–2013. Light bars represent cows and darker bars
represent bulls. Rectangular shaped bars represent percentages determined by winter and summer
range allopatry, and cylindrical bars represent those calculated by net squared displacement (NSD)
methods outlined by Bunnefeld et al. (2011). Percentages associated with southern moose are
represented with diagonal striping, and northern moose are represented by horizontal striping;
stippled bars represent all moose combined. Note: missing bars occur where n = 0.

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Mean distances moved between succes-
sive summer and winter locations were 2.5
times greater for migratory cows (n = 86)
than non-migrators (Table 1). Migratory
southern cows (n = 46) tended to move less
between summer and winter locations than
migratory northern cows (n = 40; Table 1).
Migratory distances (mean per individual)
for all cows ranged from 557-88,402 m.
Bulls (n = 21) had a similar relative pattern
with individual mean distance between sum-
mer and winter locations ranging from
2,886–52,624 m (Table 1). Summer ranges
of both sexes were located in all cardinal
directions relative to winter ranges; however,
59% of winter ranges of migratory moose
were located north of summer ranges. Net
displacement (ND) of GPS-collared moose
(mean per individual) ranged from 10,970–
94,972 m for cows and 13,060–52,660 m
for bulls (means in Table 1). Only 45% of
moose were categorized with the same
migratory pattern using the NSD techniques
when compared to the winter-summer range
allopatry analysis. Although we did not
detect any movements that led to dispersal
or forays outside the study area, we also

did not collar younger moose that would be
more likely to undertake measurable disper-
sal movements.

Home Range
The mean annual home ranges were

243.5 ± 96.9 km2 for GPS-collared cows
(n = 21; range = 106.5 – 498.3 km2) and
262.0 ± 67.7 km2 for GPS-collared bulls
(n = 11; range = 185.7 – 400.9 km2). As
expected (Joly 2005), home ranges of cows
(n = 4; 305.0 ± 47.2 km2) calculated from
1 GPS location/day were larger than cows
(n = 17; 228.9 ± 22.9 km2) with 3 locations/
day. The mean home range size of non-
migratory (NSD technique) cows was
∼10–15% smaller (n = 5; 228.1 ± 45.2 km2)
than that of migratory cows (n = 7; 247.9 ±
38.2 km2) or cows with mixed-migration
strategies (n = 8; 261.7 ± 35.7 km2). Non-
migratory (n = 4; 260.3 ± 33.4 km2) and
migratory (n = 4; 295.9 ± 33.4 km2) bulls con-
formed to this same pattern; bulls with mixed-
migration strategies had home ranges about
25% smaller than other bulls (n = 3; 219.0 ±
38.5 km2). The mean annual home range
was smallest for cows found primarily within
the KNWR (n = 5; 199.6 ± 63.0 km2),

Table 1. Distance (mean ± SE) between summer and winter locations (allopatry) and net displacement (ND;
sensu Bunnefeld et al. 2011) of cow and bull moose for the entire and portions of the upper Koyukuk
River drainage study area, northcentral Alaska, 2008–2013.

Methodology Location/Sex Non-migratory (m) Migratory (m) Mixed Migratory (m)

Allopatry All Cows 8621 ± 2241 21598 ± 2462 17598 ± 2827

Allopatry Southern Cows 7697 ± 2233 16128 ± 2233 18619 ± 2658

Allopatry Northern Cows 9494 ± 3809 29347 ± 4665 16373 ± 5110

Allopatry All Bulls 5923 ± 5923 15709 ± 3419 18820 ± 5297

Allopatry Southern Bulls 5015* 906 ± 3261 2886*

Allopatry Northern Bulls 6226 ± 7176 17644 ± 4143 22804 ±6215

ND All Cows 23584 ± 8410 49046 ± 8410 29084 ± 5947

ND Southern Cows 16526 ± 7663 15604* 40430 ± 5419

ND Northern Cows 28290 ± 10197 57406 ± 8831 21520 ± 7210

ND Northern Bulls 22267 ± 6473 37757 ± 6473 22808 ± 7474

*N = 1, no SE reported.

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increased in size in the DHCMA (n = 11; 227.9
± 88.5 km2), and was largest for cows (n = 3;
279.1 ± 77.4 km2) in the GAAR. The 2 cows
that utilized both KNWR and the DHCMA
had the largest annual home ranges (384.9 ±
160.0 km2). All 11 GPS bulls were collared
in GAAR and had larger average annual
home ranges than cows in GAAR. East of
GAAR, home ranges often overlapped the
Dalton Highway and Trans-Alaska Pipeline
System (TAPS).

For all GPS moose, winter home
ranges were slightly larger (n = 32; 24.2 ±
13.0 km2) in mild winters than moderate
(n = 36; 20.2 ± 8.6 km2) and severe winters
(n = 11; 21.0 ± 9.0 km2). Using only
3 locations/day data, we similarly found
larger winter home ranges during mild than
moderate winters, and smallest ranges during
severe winters (23.3 ± 12.5 km2, 19.4 ±
7.0 km2, and 16.4 ± 8.1 km2, respectively).
Cows appeared to be more sensitive than
bulls to severe weather; their winter home
ranges were 24.2 ± 13.7 km2, 19.0 ±
7.7 km2, and 16.4 ± 8.0 km2 in mild, moder-
ate and severe winters, respectively. No GPS
bulls were collared during a severe winter,
but their winter home range sizes were
nearly identical in mild and moderate winter
(20.9 ± 9.3 km2, 20.2 ± 8.6 km2,
respectively).

Movement Relative to Conservation Units
VHF and GPS data (n = 116 moose)

were combined to examine movements in
and around GAAR, KNWR, and the
DHCMA. Thirty-two moose stayed exclu-
sively within the confines of KNWR, includ-
ing all 5 bulls originally collared there.
An additional 21 used KNWR and areas
outside the refuge, and of these, only 1 bull
and 2 cows (14.3%) were also found
within the DHCMA; 1 cow primarily used
the DHCMA (1 location within KNWR)
and 1 bull primarily used the GAAR. Two
(1 bull and 1 cow) of these 21 moose used

GAAR extensively and did not enter the
DHCMA, and the cow consistently migrated
from GAAR in the winter to calve within
KNWR. The GAAR had 16 moose (10 bulls
and 6 cows) that stayed within its borders
and 24 that moved in and out of the area;
of the 24, 9 bulls and 11 cows (83.3%)
moved into the adjacent DHCMA (Fig. 1).
Nonetheless, most (∼70%) moose using the
GAAR and DHCMA resided primarily in
one or the other unit, with a single or few
relocations falling in the other. Moose
located commonly in both units had home
ranges with measurable overlap of both.
Robust migratory movements between these
2 units were not detected for bulls or cows.

DISCUSSION
Moose in the upper Koyukuk River

drainage exhibited partial migration (25–34%
of cows and 36–57% of bulls), similar to
other areas in Alaska (van Ballenberghe
1977, Osborne and Spindler 1993, Mauer
1998, White et al. 2014). However, the pro-
portion of moose that were migratory in the
upper Koyukuk River (35–38%) was less
than in the lower drainage (Osborne and
Spindler 1993) or the eastern Brooks Range
(Mauer 1998). Similarly, the distances
traveled from winter to summer range by
our migrators (22 km for cows and 16 km
for bulls) were less than those measured in
adjacent studies (31–123 km; Osborne and
Spindler 1993, Mauer 1998) or elsewhere
in Alaska (see review in Mauer 1998). Our
NSD calculations revealed longer migration
distances than obtained by simply comparing
mid-summer to mid-winter locations (range
allopatry). Although not directly comparable
to earlier studies because migration was
calculated differently, moose in the upper
Koyukuk River drainage exhibited migratory
behavior less frequently than moose else-
where in the region, and migrated shorter
distances. This may be due to lower moose
density, differences in habitat quality and

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distribution, and/or physiographic differ-
ences between study areas.

However, moose migration patterns are
more complicated than parsing individuals
into migratory or non-migratory categories.
In addition to partial migration, we docu-
mented that a large segment (26–50% of
cows and 24–27% of bulls) exhibited a
mixed-migration strategy (i.e., migratory
behavior only in certain years), and that the
annual strategy varied individually. Our
estimates are conservative as a portion of
our population was classified during a single
year only. This phenomenon was documen-
ted in southcentral Alaska and believed
related to snow depth (van Ballenberghe
1977). Alternatively, adaptive migration
may convey fitness benefits especially in
highly variable climates (e.g., our study
area) and in rapidly changing environments
such as the Arctic. However, migration strat-
egy does not appear to be unilaterally linked
to environmental conditions because not all
our moose, or entire populations in other
areas (van Ballenberghe 1977, Ball et al.
2001), respond similarly in a given year. As
our analyses reveal and as expected (Singh
et al. 2012, Fryxell and Holt 2013), a mix
of migratory strategies are displayed by sym-
patric moose under varied environmental
circumstances.

The mixed-migration strategies we
detected highlight the importance of long-
term studies and procuring an adequate sam-
ple size. The migratory characteristics of
a study population would probably not be
categorized accurately by short duration
studies because mixed-migratory behavior
could be erroneously categorized as either
migratory or non-migratory behavior (Dettki
and Ericsson 2008). Long-term, more
focused studies are required to determine if
migratory strategy is a behavioral trait
acquired from maternal experience as docu-
mented elsewhere (e.g., Sweanor and
Sandegren 1988). If migratory strategy is a

learned behavior, it would explain, in part,
why multiple strategies exist.

Our results also reveal that the metho-
dology used to determine migratory status
is important because:

“Migratory behavior is persistent and straight-
ened out movement effected by the animal’s
own locomotory exertions or by its active
embarkation upon a vehicle. It depends on
some temporary inhibition of station keeping
responses but promotes their eventual disin-
hibition and recurrence” (Kennedy 1985,
Dingle and Drake 2007).

The NSD technique appears to more
robustly and objectively parse moose migra-
tory status relative to this definition. How-
ever, the NSD requires data beyond
identifying if summer and winter locations
are allopatric. Because the allopatric techni-
que is simpler, requires less data, and has
been used in past studies, we employed it
to make more direct comparisons to regional
studies. Because both techniques have posi-
tive and negative aspects, identifying objec-
tives and definitions are key in determining
which is more appropriate for a specific study.

We found home ranges to be larger in the
northern part of our study, which was con-
sistent with past studies (see review by
Hundertmark 1997). Highly variable season-
ality and climatic conditions in the north
where forage productivity is lower, and
habitat is patchier due to more rugged and
higher terrain, may foster migratory behavior
and larger home ranges (Ballard et al. 1991,
Hundertmark 1997). The size of home
ranges appeared to be influenced by winter
severity as smaller home ranges occurred in
harsher winters.

Our study was promoted by public con-
cern about moose moving between conserva-
tion units (GAAR and KNWR) and areas
with different hunting regulations and acces-
sibility, such as the DHCMA. Peak move-
ment rates of bulls occurred from mid-
September through early October (Joly et al.

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MIGRATION IN NORTHCENTRAL ALASKA – JOLY ET AL. ALCES VOL. 51, 2015



2015) coinciding with much of the hunting
season; presumably this timing of bull move-
ment stimulated public concern. Exposure
of KNWR moose to the DHCMA was
negligible, likely a result of the separation
(>10 km apart) of the units in most areas
and smaller home ranges of moose in that
area. In contrast, GAAR and the DHCMA
shared a common boundary and a large pro-
portion (50%) of the moose collared in
GAAR used both units. Nonetheless, a pre-
ponderance of these moose resided primarily
in either unit, with a single or a few reloca-
tions at the edge of the adjacent unit. Moose
that spent large portions of time in both units
had home ranges that measurably overlapped
both units. With a single exception, we did
not detect regular or substantial migratory
movements between the DHCMA and
GAAR or the more distant KNWR.

This study was the first of its kind in this
region of Alaska and these data provide a
basis for evaluating moose movements rela-
tive to harvest concerns and distribution of
the moose resource, specifically in our study
area. Understanding the spatial ecology of
moose improves the understanding of moose
ecology, behavior, and demographic and
genetic processes. It is also critical to devel-
oping a comprehensive management pro-
gram (Hundertmark 1997), especially where
harvest allocation, resource damage, and
local management issues are influenced by
seasonal migration and distribution of
moose.

ACKNOWLEDGEMENTS
This project was promoted by public

subsistence and wildlife advisory groups.
Funding was provided by the National Park
Service, US Fish and Wildlife Service,
Alaska Department of Fish and Game, and
the Bureau of Land Management. We thank
pilots T. Cambier, M. Spindler, M. Webb,
P. Zaczkowski, C. Cebulski, L. Dillard,
P. Christian, A. Greenblatt, H. Bartlett,

N. Guldager, D. Sowards, and S. Hamilton
for making this project possible and safe.
S. Backensto, J. Burch, J. Caikoski, K.
DeGroot, J. Dillard, M. Flamme, A. Morris,
C. Harwood, T. Hollis, E. Julianus, H.
Kristenson, J. Lawler, N. Pamperin, T. Paragi,
K. Rattenbury, C. Roberts, L. Saperstein, G.
Stout, and many others provided critical
assistance with project implementation.
We thank G. Stout for his contributions
to project management and insights into
the ecology of moose within the region.
N. Bywater, C. Harwood, S. Miller, R. Sarwas,
and A. Quist provided database and GIS
expertise. We appreciate the constructive
reviews by E. Addison, R. Churchwell, and
anonymous reviewers. All moose captures
adhered to State of Alaska Animal Care
and Use Committee (ACUC) guidelines
(#07–11).

LITERATURE CITED

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MIGRATION IN NORTHCENTRAL ALASKA – JOLY ET AL. ALCES VOL. 51, 2015


	MOOSE MOVEMENT PATTERNS IN THE UPPER KOYUKUK RIVER DRAINAGE, NORTHCENTRAL ALASKA
	Methods
	Study area
	Moose relocation data and GIS analyses

	Results
	Moose Relocation Data
	Migration
	Home Range
	Movement Relative to Conservation Units

	Discussion
	ACKNOWLEDGEMENTS
	LITERATURE CITED