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Title
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Migratory and Non-Migratory Movements and Habitat Use by Female Elk on the Cascade Range of Washington
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Date
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2010
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Creator
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Moeller, Barbara J
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Subject
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Environmental Studies
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Migratory and Non-migratory Movements and Habitat Use by Female Elk in the
Cascade Range of Washington
by
Barbara J. Moeller
A THESIS
Submitted in partial fulfillment of the requirements for the degree of
Master of Environmental Study
The Evergreen State College
Completed December 2010
© 2010 by Barbara J. Moeller. All rights reserved.
ii
This Thesis for the Master of Environmental Study Degree
by
Barbara J. Moeller
Has been approved for
The Evergreen State College
by
______________________________
Dylan Fischer, PHD
_______________________________
Brian A. Gilbert, PHD
________________________________
Steven G. Herman, PHD
_______________________________
Date
iii
ABSTRACT
Migratory and Non-migratory Movements and Habitat Use by Female Elk in the
Cascade Range of Washington
Barbara J. Moeller
Most research studies have evaluated non-migratory or migratory elk (Cervus elaphus)
behavior and their use of habitat as unique separate groups. Few studies have evaluated
migratory and non-migratory elk habitat use within the same elk herd. This thesis focuses on the
habitat use of female elk (Cervus elaphus) with a specific emphasis on the similarities and
differences between migratory and non-migratory groups of elk in the southwestern Cascade
Range of south-central Washington during 2004 – 2008. I used relocations on 53 radio-collared
female elk and >80,000 locations on 15 GPS (Global Positioning System) collared female elk.
Habitat use and movement, home range, survival, and herd fidelity were studied and measured on
the collared elk to assess elk habitat use across the 1,028 km2 study area.
Differences were detected between the migratory and non-migratory groups of elk where
migratory elk were found to use steeper terrain, were in habitat areas in closer proximity to cover
(safety), and tended to be found in habitat areas that were greater distances from paved roads than
non-migratory elk. Annual survival rates ranged between 71% and 90% over the course of the
study. Home range size ranged from 5.4-102 km2. Differences in home range size were detected
between the migratory and non-migratory groups of elk for each biological year (BYR) of the
study (2004-2008) and for the overall life home range. The elk in our study also demonstrated
strong site fidelity (99%).
ACKNOWLEDGEMENTS
While driving my truck the other day, I was contemplating acknowledging
friends, family and colleagues that have contributed to my thesis process. During the
drive Janis Joplin came on the radio. Her raspy soulful voice rang out embedded within
the rich accompaniment of her band. It made me think about how talented she truly was,
but what also became very clear, were the players involved in contributing to her success.
The possible range of family, friends, artists that inspired her, producers, and band mates
that influenced her final products were probably countless. Although my research and
the resulting thesis takes place in a vastly different discipline, venue and at a much
smaller scale, I am as grateful as Janis could have been to the many people that have
contributed to my research project and my thesis.
I am eternally grateful to the Puyallup Tribe of Indians for providing me the
opportunity to work for them, conduct research, and use the data I have collected for the
thesis. Within the Tribal organization, I am especially grateful for the support of the
Tribal Council, the Hunting Commission, staff and the general membership.
Specifically, I would like to thank Bill Sullivan, Natural Resources Director for his
continual support, flexibility, and encouragement. Very special thanks to Paul Arnold for
his superb GIS expertise and very considerable mapping contributions on the project.
Additional thanks to tribal project participants include: Mary Basballe, Phillip Dillon,
Don Coats, and Paul Herrera. Special thanks to my predecessor Mike McDonald for
initiating research on the South Rainier elk herd.
The project would not have been possible without the support of the United
States Fish and Wildlife Service Tribal Wildlife Grant and the Bureau of Indian Affairs
Unresolved Hunting and Fishing Rights grant. Additional thanks to cooperating
agencies, tribes and organizations include: Northwest Indian Fish Commission, especially
Chris Madsen, Muckleshoot Indian Tribe, especially Dave Hamilton and David Vales,
Squaxin Island Tribe of Indians, especially Pete Kruger, Nisqually Tribe of Indians,
Georgiana Kautz and Dave Trout, Quinault Indian Tribe, especially Grover Oakerman,
Washington State Department of Fish and Wildlife, especially Briggs Hall, DVM, Robin
Woodin, and Richard Stone, Mount Rainier National Park, especially Jim Schaberl,
Gifford Pinchot National Forest, especially Tom Kogut and Joe Kulig.
iv
Very special thanks to my thesis committee chairman, Dylan Fischer, for his
exceptional attention to detail, guidance, and effort on the manuscript. As well, special
thanks to Brian Gilbert and Steve Herman for participation on my thesis committee. Also
special thanks to Ted Whitesell of The Evergreen State College.
Capturing and collaring of the elk for the study would not have been possible
without expert helicopter pilot Jess Hagerman. Honorable mention goes out to Jerry and
Bridgette White for sharing their home, coffee, and wood spirits and allowing us to stage
elk capture from their property. Additional thanks to N. Carroll, Janie Unbehagen, DVM
and staff at the Animal Care Clinic, B. Anderson, J. Foster, and the residents of
Packwood and Randle for their patience and support during elk projects.
I am extremely grateful to Lesley Richardson for the inspiration to enter into
graduate school and sustained support on all aspects of this project, including master edits
and formatting of the manuscript. Also thanks to Charmane Ashbrook, Shon Kayli, Dr.
Jennifer Holt, and sister Julie Jeronimus for extended support and encouragement.
Special thanks to my partner Kathryn McMahon for loving support, thoughtful
sweetness, and encouragement to persevere.
This thesis is dedicated to the memory of my father Donald D. Moeller who
passed on in February of 2010. Without my father’s encouragement to play hockey as a
girl and pursue my dreams, as non-traditional as they were, I would not be the woman I
am today! Thanks Dad!
v
TABLE OF CONTENTS
Item
Page
________________________________________________________________________
HOME RANGE, SURVIVABILITY, AND HABITAT ANALYSIS OF ELK IN
SOUTHWEST WASHINGTON………………………………………………………….1
INTRODUCTION/HISTORY……………………………………….......1
STUDY AREA……………………………………………………….…8
METHODS………………………………………………………….…11
Elk Captures…………………………………………..………11
Radio Telemetry………………………………………………13
Home Range and Core Use Areas……………………………..13
Survival of Cow Elk…………………………………………...15
Habitat Variables……………………………………………....15
Habitat Use, Seasonal Habitat Use and Time of Day……….....19
RESULTS……………………………………………………................20
Elk Captures………………………………………………..….20
Home Range …………………….……………………….........20
Migration ……..…………………………………………….…24
Distribution………………………………………………….....24
Survival Rates………….………………………………………28
vi
Item
Page
________________________________________________________________________
Habitat Use: Activity………………………………………....28
Aspect…………………….…………………....30
Proximity to Cover…………………..……...…..32
Proximity to Shrub/Scrub……………….…...….37
Slope………………………………..……..…..…40
Proximity to Water………………..…………….44
Proximity to Roads and Trails……………….….47
Habitat Use with Season and/or Time as Predictors……..……54
DISCUSSION……………………………………………..…………...55
Home Range………………………………..……………….…55
Migration and Distribution…………………………………….57
Survival Rates………………………………………...………..57
Habitat Use: Aspect…………………………………...……….58
Slope………………………………………...…...60
Proximity to Cover…………………...…………..60
Proximity to Shrub/Scrub………..……..………..61
Proximity to Water….………………………..…..62
Proximity to Roads and Trails………….…..……..63
MANAGEMENT IMPLICATIONS……………………………..…....66
vii
LITERATURE CITED……………………………………………..….70
APPENDICES….…………………………………………………….75
viii
LIST OF APPENDICES
Appendix
1
Winter range plant list for Western Hemlock zones………………………….....76
2
Summer range plant lists for Pacific Silver and Mt. Hemlock zones……….......77
3
Batch fixed kernel density for ten GPS collared elk at the 50, 90 &95% range
area………………………………………………………………………………78
4
Habitat attributes and distances classes for habitat use analysis of female elk in
southwest Washington…………………………………………………………..84
5
All elk. Radio frequencies, capture dates and total number of locations of radiocollared elk in southwest Washington 2004-2008………..……………………..86
6
Elk name, life home range area, and migratory status of radio-collared elk in
southwest Washington, 2004- 2008………………………………………..……87
7
Habitat use of aspects by elk in the combined Randle and Packwood study areas
at the 50% and 95% range scales in the Southwest Cascades of Washington.
2007-2008……………………………………………………………………….89
8
Habitat use by elk relative to distance to cover in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008……............................................................91
9
Habitat use by elk relative to distance to shrub/scrub in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008………………...………………………….92
10
Habitat use by elk of slopes in the combined Randle and Packwood study areas at
the 50% and 95% range scale in the Southwestern Cascades of Washington,
2007-2008…………………………………………………………………..…...93
11
Habitat use by elk relative to distance to water in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008………………………………………...…94
12
Habitat use by elk relative to distance to paved roads in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008………………………………………….95
13
Habitat use by elk relative to distance to non-paved roads in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the
Southwestern Cascades of Washington, 2007-2008…………………………….96
ix
14
Habitat use by elk relative to distance to trails in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008………………………………………….97
15
R2 values for Individual elk by habitat attribute at both the 50% and 95%
scales……………………………………………………………………..……98
16
GPS collared elk data points per migratory status in the Randle subunit study
area…………………………………………………………………………..101
LIST OF FIGURES
Figures
1
Study Area for elk in the Southwestern Cascades of
Washington……………..9
2
Batch fixed kernel density for Elk #30 at the 50, 90 &95% range area………...14
3
Packwood elk habitat analysis area……………………………………………..16
4
Randle elk habitat analysis area…………………………………………..……..18
5
GPS collared elk data points per migratory status in the Packwood subunit study
area........................................................................................................................23
6
Elk distribution in the Packwood study area……………………………………25
7
Elk distribution in the Randle study area……………………………..………...27
8
Activity of migratory and non-migratory elk in the combined Randle and
Packwood
study areas in the Southwest Cascades of Washington. 20072008………………………………………………………………………….….29
9
Distance to cover analysis for migratory and non-migratory elk at the 50% scale
in the Southwestern Cascades of Washington……………….………………….33
10
Distance to cover analysis for migratory and non-migratory elk at the 95% scale
in the Southwestern Cascades of Washington………………………….……….36
11
Distance to shrub/scrub analysis for migratory and non-migratory elk in the
Southwestern Cascades of Washington…………………………...……………39
12
Migratory elk habitat selection with slope analysis………………………..…...43
13
Non-migratory elk habitat selection with slope analysis………………...……...43
14
Migratory and Non-migratory elk habitat use and distance to roads in the
Southwestern Cascades of Washington…………………………………………48
x
LIST OF TABLES
Tables
1
Minimum Convex Polygon (MCP) home range estimates for migratory and nonmigratory elk for years 2004 – 2008 and life home range comparisons for
Western Washington elk in the upper Cowlitz River valley……………………21
2
Overall annual habitat use of aspects by elk in the combined Randle and
Packwood study areas at the 50% and 95% range scales in the Southwest
Cascades of Washington. 2007-2008……………………………………………30
3
Overall annual habitat use by elk relative to distance to cover in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the
Southwestern Cascades of Washington, 2007-2008……………………………34
4
Overall annual habitat use by elk relative to distance to shrub/scrub in the
combined Randle and Packwood study areas at the 50% and 95% range scale in
the Southwestern Cascades of Washington, 2007-2008……………………….37
5
Annual overall habitat use by elk of slopes in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008……………………………………….…..41
6
Overall annual habitat use by elk relative to distance to water in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the
Southwestern Cascades of Washington, 2007-2008………………...……….….45
7
Overall annual habitat use by elk relative to distance to paved roads in the
combined Randle and Packwood study areas at the 50% and 95% range scale in
the Southwestern Cascades of Washington, 2007-2008………………….….…47
8
Overall annual habitat use by elk relative to distance to non-paved roads in the
combined Randle and Packwood study areas at the 50% and 95% range scale in
the SW Cascades of Washington, 2007-2008……………………………….….49
9
Overall annual habitat use by elk relative to distance to trails in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the SW
Cascades of Washington, 2007-2008…………………………………………...52
xi
xii
INTRODUCTION
Habitat loss is one of the biggest challenges wildlife species face worldwide.
Conservation of elk (Cervus elaphus) winter range in particular continues to be a
challenge to wildlife managers nationwide. Since the 1960s there has been a dramatic
increase in human occupation of elk range in Western Washington State (U.S.A.), and
most notably those geographic locations supporting elk winter ranges at lower elevations
(Lyon and Christensen 2002). Conservation of winter range is critical for the
sustainability of elk herds globally. Examining the way migratory and non-migratory elk
use habitat may provide valuable insight into planning conservation efforts for the entire
herd.
Differences in habitat use between migratory and non-migratory elk within the
same herd are poorly understood both globally and locally. The literature includes a vast
volume of studies on elk behavior while very few, if any, describe migratory and nonmigratory elk behavior within the same herd.
Prior research has identified a need for herd-specific studies to be conducted on
habitat selection of elk (Holthausen et al. 1994). Research gaps include information on
the movements and wintering areas of the South Rainier elk herd (WDFW 2002).
Movement patterns inside Mt. Rainier National Park have been described (WDFW 2002).
However, movement patterns of elk living outside the park are poorly understood
(WDFW 2002). A better understanding of winter range habitat use is important in order
to: 1) identify areas to pursue for elk conservation in the most conflict prone area of the
winter range via land acquisition and conservation easements, 2) assess potential impacts
of increased development (e.g., along the upper Cowlitz River) and modified Forest
practices in the Gifford Pinchot National Forest (WDFW 2002), and 3) identify whether
impacts are likely to affect migratory, non-migratory, or all segments of the herd.
This thesis will provide habitat use analysis for both summer and winter range
areas in the Packwood and Randle areas of Lewis County in Washington State, and will
also examine the habitats and behavior of migratory vs. non-migratory elk. In this study,
I used four population metrics to analyze and compare the South Rainier elk herd. I will
address migration, fidelity, home-range, and survival, as indicators of stability,
demography, and movement. The intention of this thesis is to provide herd-specific
habitat use data to fill an existing research gap for the South Rainier elk herd.
1
I will explore differences in resource selection between and among migratory vs.
non-migratory female elk by analyzing habitat use areas of South Rainier female elk at
the core (50% contour level using a fixed kernel approach )) and home range (95%
contour level fixed kernel). The major goals of this study are to: 1) compare and analyze
home range areas of migratory and non-migratory elk in the study group; 2) compare and
analyze winter and summer range use areas of migratory and non-migratory elk in the
study group; 3) compare and analyze rates of pregnancy and mortality (where sample size
is large enough) of migratory and non-migratory elk in the study group; and 4) describe
herd dynamics and habitat use. Based on previous studies in other systems, I hypothesize
that habitat use and home range size between migratory and non-migratory elk will be
significantly different. The intent of this study is to identify and describe any differences
in habitat usage between, and among, migratory and non-migratory elk at both homerange, and the 50% and 95% MCP use areas within the greater home range on both
summer and winter range. Based on my findings, I will recommend management
strategies, and help to identify future research needs.
HISTORY
In the state of Washington, 10 distinct elk herds (Cervus elaphus) have been
identified as inhabiting various regions of the State. The native elk that were historically
found in greater Washington State have been identified as the subspecies Roosevelt elk
(Cervus elaphus roosevelti). The historic range of the Roosevelt elk subspecies extended
from just North of San Francisco, CA to Vancouver Island, BC, and inland from the
Pacific Ocean to the Cascade Mountains (O’Gara 2002). The current range includes the
coastal areas of Humboldt and Del Norte counties, CA, northward through western
Oregon and Washington to Vancouver Island (O’Gara 2002). Historic, archeological,
and anthropologic records support evidence of substantial numbers of elk in Washington
State prior to European exploration and settlement at the start of the 19th century
(Schullery 1982). By 1900, much of western Washington (with the exception of the
Olympic Peninsula) was devoid of elk. In many instances, native elk were thought to
have never existed in areas such as the Mt. Rainier National Park area (Bradley 1982).
Elk in the Cascade Range of Washington were eliminated to the extent that many early
settlers, explorers, and their descendents held the belief that elk were not a native species
of the area.
2
Historic records not only affirm the presence of elk in Washington State prior to
European settlement, but also include documentation of Native Americans relationship
with elk. Some examples of the subsistence and cultural importance of elk include uses
of elk horn by the Puyallup and Nisqually Tribes to make clubs, elk horn wedges to split
cedar planks, elk and deer hides to wrap bodies for burial, the use of untanned elk skin to
make “parfleches” that were used as containers, and consumption of elk for sustenance
(Haeberlin and Gunther, 1930). Members of the Puyallup Tribe were known to hunt and
to use fire to manage for deer and elk habitat in the area surrounding Mt. Rainier
(Schullery 1984). Puyallup Tribal groups also travelled annually to Mt. Adams, Mt.St.
Helens and various other mountain regions throughout the state for annual huckleberry
picking, and hunting of deer and elk during late summer and early fall (Powatten Mills,
personal correspondence). Camps were set up for the duration of the annual berry
picking and associated hunting in the Mt. Adams, St. Helens, and Indian Wilderness
areas. Thompson also documents local tribal activity around Mt. Rainier as centered on
hunter-gatherer camps near important huckleberry fields at an altitude between 3,000 and
5,000 feet (Thompson 1981). Trade routes were used by Puyallup members throughout
the state. A primary route used to connect to the east side of the Cascade Mountains was
the Naches Trail. It was also common practice for Tribal members to engage in hunting
of deer and elk while travelling. Inter-tribal marriages were also common. These further
strengthened bonds amongst Tribes within the region. The Packwood Pass area, which
extended up the Ohanapekosh across to the Cowlitz Divide that flanks the south and east
sides of Mt. Rainier, was regarded by local tribes as some of the very best hunting
grounds (Schullery 1982, Brown 1920). The presence of native elk and their associated
cultural significance to local tribes may therefore be considered to be well established.
Rocky Mountain elk (Cervus elaphus nelsoni) is a North American subspecies of
elk whose range includes the Rocky Mountains and adjacent mountain ranges, from the
55th parallel southward to the 35th parallel (O’Gara 2002). At the turn of the 20th century
natural resource managers established new policies and regulations to aid in reestablishing the extinct elk herds. This led to Rocky Mountain elk being introduced to
the area around Mt. Rainier, starting in the early 1900s. The intent of the introduction of
Rocky Mountain elk by game managers was to supplement existing remnant populations
of native elk, and to re-establish elk where they had been extirpated. Also, by the 1950s,
forest practice changes such as extensive logging in the Cascades around Mt. Rainier
3
National Park (MRNP), also helped created habitat conditions favorable to elk. This
further boosted local elk populations. Elk herd numbers began to steadily grow. A
census survey conducted by the U.S. Forest Service in September 1962 counted 466 elk
along the eastern border of Mt. Rainier National Park (Schullery 1982, Bender 1962). At
the time, elk were still believed to be very scarce. Park managers not only believed that
very few elk resided in the Park; but they were also under the impression that MRNP
could not possibly support that many animals (Schullery 1982, Bender 1962). Park
managers were skeptical of the elk count, and they conducted their own survey of the
same general area on October 16, 1962. This survey found between 25-30 animals, which
did not support the numbers from the previous survey (Schullery 1982, Bender 1962).
However, Gilbert and Moeller (2008) have shown elk movement across the
landscape to be temporally and spatially dynamic. It is not uncommon to observe
dramatic shifts in habitat use during the onset of the rut (breeding season) in September,
and the rut in October. Elk also have a tendency to use habitat with cover more
frequently during the rut than during the time period leading up to the rut. Typically, elk
will tend to be in larger groups in open landscapes (IE: alpine meadows, shrubs) prior to
the rut. During the rut, the elk tend to fragment into smaller breeding groups. These
groups are frequently found in timber in the sub-alpine areas, which make them more
difficult to detect. With a greater understanding of elk ecology, the disparity in elk
counts between a September and October survey in 1962 is not surprising.
By the 1970s, Washington State Department of Fish and Wildlife (WDFW)
identified ten separate elk herds in Washington State. The South Rainier elk herd is one
of those 10 herds identified, and is the focus of this thesis. Since the mid-late 1900s,
estimates of herd numbers have dramatically fluctuated based on forest practices,
development, and legal and illegal hunting. As already mentioned, extensive logging in
the mid-1900s created large gaps in the forest canopy that stimulated growth of preferred
elk browse species such as: Vaccinium sp., Rubus sp., Salal (Galtheria shallon) Red
elderberry (Sambucus racemosa), Sword fern (Polystichum munitum), and Bear grass
(Xerophyllum tenax (Pursh) Nutt). With the improved habitat conditions for elk, the
carrying capacity in the area likely increased. Elk herd numbers may have increased in
turn. With larger numbers of elk, and the use of less accurate methods for estimating the
numbers of elk, regulations were formulated that allowed extensive harvest of both males
and females of the population. Recent examples of the fluctuating South Rainier elk herd
4
numbers include the 1994 WDFW herd estimate of approximately 3,800 animals,
followed by an estimate in 1998 of only 1,700 animals (WDFW 2002). Liberal state
hunting regulations which included unlimited cow harvest in the mid-1990s (typically
used to reduce population numbers) were employed and resulted in a dramatic reduction
of the population. Current population estimates are made annually in spring of each year
by the Puyallup Tribe. The 2009 estimate was 1,000 animals.
The Puyallup Tribe of Indians are signatory to the Medicine Creek Treaty of
1854. The Treaty guaranteed the Tribe the right to hunt on open and unclaimed lands.
The primary stock of elk currently harvested by the Tribe for ceremonial and subsistence
purposes is the South Rainier elk herd. As a sovereign nation, the Tribe manages their
own resources for wildlife, and sets hunting and harvest regulations based on the best
available science. To better estimate herd numbers, a computer model for estimating elk
abundance, and to monitor population trends was developed by the Puyallup Tribe
(Gilbert and Moeller, 2008). The Tribe conducts annual surveys to collect the raw data to
be used in the computer model for population estimates. The population estimates are
used to steer the hunting season and regulation setting process for the South Rainier herd.
Since 2004, the Tribe has raised approximately $1,000,000 in grant (USFWS and BIA)
and tribal funding for resource management activities benefitting the South Rainier elk
herd. Management activities include: habitat improvement projects on winter and
summer range, land acquisition for elk conservation, population model development,
aerial surveys, mortality/survival estimates, and VHF and GPS collaring of elk.
Migration
Migration can be defined as animal movement, usually periodically, from one
region or climate to another for feeding or breeding (Skovlin et al. 2002, Gove 1969).
Kennedy provided a behavior-oriented definition of migration as: Migratory behavior is
persistent and straightened-out movement effected by the animal’s own locomotory
exertions or by its active embarkation on a vehicle. It depends on some temporary
inhibition of station-keeping responses, but promotes their eventual disinhibition and
recurrence (Kennedy 1985). A variety of elk migratory behaviors have been observed
and well documented with associated hypotheses explaining the behavior. Some of the
variations in elk migration include: elk that do not migrate; migration in response to
changes in forage conditions (Irwin 2002, Graf 1943); and true migration described as
5
migrations as changes of habitat, periodically recurring and alternating in direction,
which tend to secure optimal environmental conditions at all times (Thomson 1926 pp 3).
Examples of elk movement that have been observed but have not been considered true
migration have been differentiated by factors such as: 1) summering areas often are
accessible during winter; 2) movements are not consistent among herds; and 3) the timing
of movements differs among herds (Irwin 2002, McCullough 1969).
Once the migration status of elk based on an established definition of migration
has been determined, variations in migration are still common between and among
regions, sex, and age class of elk, in addition to differences between pregnant and barren
cows (Irwin 2002). Temporal and seasonal variations also influence migration and
timing of migration. Indentifying differences in how migratory and non-migratory elk
use the landscape is important in understanding patterns of habitat use.
Elk migration may be also viewed as dynamic with regard to animal response to
anthropogenic influences. An example of this is non-migratory segments of populations
that have been culled, thus reducing actual numbers of the population that would
represent non-migratory behavior. In many areas, overall patterns of migration have been
obscured because of this type of culling (Irwin 2002). Other examples of elk response to
human influence have been where elk find protection afforded by national parks or
refuges on summer and transitional ranges, and they tend to stay within those sanctuaries
until the hunting season is over or they are driven down by deep snow (Lovaas 1970,
Picton and Picton 1975, Brown 1985). Rocky Mountain elk have also been known to
have changed their migration routes in response to human settlement and hunting (Boyce
1989).
Fidelity
Fidelity is the tendency of an animal either to return to an area previously
occupied or to remain within the same area for an extended period of time (White and
Garrott 1990). In many cases, it is commonly believed that there is substantial variation
in migratory patterns of elk between years. Specifically, the theory held is that there are
increasing numbers of elk year-to-year that have changed their migratory pattern to a
non-migratory habit based on natural selection. In general, female offspring tend to
establish ranges in or adjacent to their mother’s social group (Raedeke et al. 2002).
Many studies have reported strong philopatry of female elk for seasonal and annual
6
ranges (Craighhead et. al. 1972, Rudd et al. 1983, Edge and Marcum 1985, Edge et al.
1986, Smith and Robbins 1994, Raedeke et al. 2002). Strong fidelity, tradition, and
learned behavior among cow elk likely contribute to the stability of social groups
(Raedeke et al. 2002, Edge et al. 1986, Van Dyke et al. 1998.).
Home Range
Home range is considered the area that is traversed by the individual in its normal
activities of food gathering, mating, and caring for young (Burt 1943: 351). Normal
activity is commonly accepted as 95% of the locations of an animal within the entire
home range area (White and Garrott 1990). Home range may be measured using a
variety of different methods yielding varying results. Understanding home range, and the
various aspects of home range, such as variation in range areas between migratory and
non-migratory elk is very important. Baseline knowledge of critical core habitat areas
should be understood before attempts are made to improve existing habitat conditions to
benefit elk herds. In the current study, much of the land use in the winter range area is
potentially not compatible with the presence of elk. Complaints are made annually and
include summer crop damage and livestock fence damage by elk (WDFW Elk Nuisance
Report 2008). Conflicts between land owners and elk occur throughout the year within
the study area (WDFW Elk Nuisance Report 2008). However, elk learn easily from their
experience, and may adjust their ecology and behavior accordingly (Geist 1982). By
using data based management strategies, conflicts on winter range could be reduced
and/or mitigated. Exploring differences at the 50% and 95% MCP scales between
migratory and non-migratory elk may yield results that enhance understanding of how elk
use the landscape depending on their migratory status.
Cow Survival
Annual survival estimates on the reproductive portion of an elk population are
important in managing for sustainable populations of elk. Survival rates may be used to
steer management strategies for the specific herd. Pregnancy rates are also an important
measure of the health of an elk herd.
7
STUDY AREA
Location
The study area includes both summer and winter ranges of the South Rainier elk
herd. Winter range is associated with the upper Cowlitz drainage near the towns of
Packwood and Randle, in eastern Lewis County of Washington (Figure 1). Summer
range areas are bordered by the Cascade Range crest to the east, Mt. Rainier to the North,
the Goat Rocks Wilderness Area to the southeast, Mt. St. Helens to the southwest, and
the eastern shores of Riffe Lake to the west (Figure 1). The entire study area
encompasses approximately 1,028 km2 (102,841 ha). Land ownership is made up of
primarily public land (Gifford Pinchot National Forest, Mount Rainier National Park
(MRNP), Mt. St. Helens, Washington State Department of Natural Resources (WADNR),
WDFW), some industrial timberlands (Port Blakely), land held by the Puyallup Tribe of
Indians, and a small percentage in private land ownership. The majority of critical winter
range is located on private land. The entire study area is located within the Medicine
Creek Treaty traditional use area.
Climate
This sub-region is categorized as a temperate maritime climate which typically
experiences hot dry summers and cool, wet winters. Annual precipitation ranges between
1.47 m (57.8 in) in the low elevation areas and 3.2 m (126 in) in the higher elevation
areas of the range (National Weather Service 2010). Average annual temperature in the
Packwood area is 9.67 C (49.7 F) (National Weather Service 2010). Years 2008 and
2009 experienced La Nina conditions, which resulted in cooler than normal temperatures,
and higher than normal levels of precipitation. Both 2008 and 2009 experienced >1 m (3
ft) of snow on winter range for more than 2 months. Typically, snow accumulation on
winter range does not exceed 0.3m (1 ft) and usually melts within 2 weeks (National
Weather Service 2010).
Topography and Vegetation: Winter Range
Topography in the winter range area is flat to rolling in the river bottoms but
increases steeply in the uplands. The area is dominated by dense coniferous forests of
Douglas-fir (Pseudotsuga menziesia) and Western hemlock (Tsuga heterophylla) in the
8
uplands, open agricultural and rangelands in the valley bottom, and open hardwood
galleries dominated by Big leaf maple (Acer macrophyllum), Black cottonwood (Populus
trichocarpa) and Red alder (Alnus rubra) along the river floodplain (Gilbert and Moeller,
2008). Elevation in the winter range area ranges from 300 m to 2,100 m. The nonmigratory elk in the herd remain
Figure 1. Study Area for elk in the Southwestern Cascades of Washington.
9
generally in the winter range area. The range of the herd falls within the Southern
Washington Cascades Province (Franklin and Dyrness, 1973) and the winter range area
generally falls within the Western hemlock (Tsuga heterophylla) zone (Franklin and
Dyrness, 1973). Detailed plant zone tables are located in Appendix 1.
As stated above, a very small proportion of winter range is within public
ownership (WDFW 2002). Much of the river bottom areas on winter range include land
uses such as agricultural, private timberlands, recreational/undeveloped lands. A smaller
portion of winter range is residential. Human tolerance for elk in these areas is mixed.
Since 2004, local and state jurisdictions on average have received 15 nuisance/damage
complaints annually. In many cases, the State has allowed special hunts during winter,
which has targeted antlerless elk where chronic problems persist. Tribal and state
managers agree that lethal removal of elk is not a long-term solution for nuisance
complaints but the hunts have served to ease local tension and political pressure.
Topography and Vegetation: Summer Range
Summer range forest zones include the Western hemlock (Tsuga heterophylla)
zone, the Pacific silver fir (Abies amabilis) zone, the Mountain hemlock (Tsuga
mertensiana) zone, the upper parkland subzone, and the Alpine zones (Franklin and
Dyrness, 1973). The topography in the mid-elevation areas up to the alpine, are moderate
to steep in most areas. The elevation range spans between 305 m – 1,829 m (1,000 ft –
6,000 ft). Detailed summer range plant zone tables are located in Appendix 2.
Land ownership on summer range areas for the migratory elk in the study group
primarily consists of public lands that include MRNP, Gifford Pinchot National Forest,
and Mount Saint Helens National Monument. Approximately, 66% of the wintering herd
is migratory. Approximately 34% of the wintering herd(s) is non-migratory. The nonmigratory segment of the respective herd is responsible for the damage complaints by
farmers that occur over the summer months. Crop damage is a common complaint filed
with local and state jurisdictions. In response, numerous damage/kill permits for
antlerless elk are issued during the month of August. As a result, cow elk with dependent
calves (2 months old) are likely removed as a part of these damage hunts. Overall
patterns of migration of elk may be obscured by exposure to hunting (Irwin 2002). This
type of management is a cause for concern because it potentially alters the understanding
of elk migration.
10
METHODS
The Puyallup Tribe began the first phase of research, conducted between 1999
and 2000, and placed 36 radio collars on female elk in the study area. At that time, the
wildlife staff was interested in learning more about the ecology of the South Rainier elk
herd. Some areas of interest included: migration patterns, timing of migration, herd
fidelity, and information on calving areas. Analysis of data in that phase of research was
not completed. After a gap of two years, the research was continued under subsequent
United States Fish and Wildlife Service and Bureau of Indian Affairs Grants. Since then,
the Tribal Wildlife Program has maintained approximately 30 cow elk fitted with Very
High Frequency (VHF) radio-collars. Cursory analysis of the data has been completed by
the Tribal Wildlife Program (Puyallup Tribe USFWS grant report).
As a result of initial data analysis and population modeling work (Gilbert and
Moeller, 2008), multiple habitat improvement projects have been funded and land
acquisition for elk conservation on winter range has been completed. In 2004, the Tribe
used relocations from the radio-collared elk to begin work on development of a computer
model (Gilbert and Moeller, 2008) to provide an alternative method for determining elk
numbers due to changes in forest management that made the current methods for
estimating elk abundance unreliable in the more heavily forested areas.
Bureau of Indian Affairs (BIA) funding for 4 GPS collars and associated
equipment was awarded to the Tribe in 2007. Those collars were deployed on 4 adult
cow elk in March 2007. USFWS funding for 11 GPS collars was awarded to the Tribe in
2008. Those collars were deployed on 11 adult cow elk in March 2008.
Elk Captures
In order to place collars on elk, each animal was remotely darted from a Bell 206
B3 (Jet Ranger) helicopter with a Carfentanil citrate (ZOOPHARM, Laramie, Wy) Xylazine hydrochloride (Webster Veterinary Supplies) drug combination during March
2004-2008. During capture efforts, we attempted to select adult cow elk evenly across
the winter range area to avoid oversampling of specific social (family) groups. While the
elk were concentrated on winter range, there was no way of knowing the migratory status
of the captured elk until the onset of the following summer migration. The Carfentanil
citrate-Xylazine hydrochloride drug combination was reversed using Naltrexone (Anazao
Health, Amarillo, TX) and Yohimbine (Anazao Health, Amarillo, TX) once the animal
11
was processed. The darting crew, consisting of two people, were dropped off near the
darted elk and began processing the elk by blindfolding, hobbling, and collecting body
temperature readings. The capture crew of four people were shuttled by the helicopter to
a location near the darted elk. The capture crew then completed processing the darted
elk. Each animal underwent the following procedure: Dart removal, fitted with a GPS or
VHF radio-transmitter collar, blood drawn and sent to lab for pregnancy, disease, and
DNA analysis, body condition assessment, age estimation based on dentition wear, fecal
sample collected for parasite testing, vaccination with antibiotics and vitamin injections.
Age estimates made by the capture crew were based on patterns of tooth eruption and
wear (Quimby and Gaab 1957). The average time the animal was immobilized was
approximately 15 minutes. A veterinarian with extensive elk capture experience was
contracted to work on the captures.
Fecal and serum samples were prepared per veterinary sampling standards and
shipped to Washington State sanctioned laboratories for testing. Disease testing included
testing for the following suite of parasites and diseases: Brucellosis, Blue Tongue,
Coccidia, Liver flukes, Lungworm, Leptospira grippo, Leptospira Hardjo, Leptospira
Ponoma, Leptospira ictero, Leptospira canicloa, Dictocaulus, Anapolysis, Strongyles,
and Capillaria. The elk we captured and collared in 2004 were also tested for gene stock.
Samples were submitted to the WDFW laboratory
Each elk was either fitted with a mortality sensing radio-telemetry collar (MOD500, Telonics Inc., Mesa, Arizona, USA), or a Spread Spectrum Global Positioning
System (GPS) telemetry collar (Gen-3,SST, Telonics Inc., Mesa, Arizona, USA). The
VHF collars had an estimated 3-year battery life. The GPS collars had an estimated 1year battery life. The GPS collars were programmed to record fixes (Latitude and
Longitude) 24 times a day at one-hour intervals while on summer and winter range.
Fixes were recorded 4 times a day at 6-hour intervals during transitional periods in spring
and fall. The collars were also engineered with an activity sensor that records the number
of times the animal moves its head in a feeding posture over a 60-minute period 24 times
each day. For example, if an animal moved its head 1-10 times in an hour it would be
considered to be bedded/resting and/or ruminating. The collars were programmed to
upload the data collected 2 times each week for a 4-hour period via the associated
transceiver (Telonics Inc., Mesa, Arizona) and RAU software (Telonics Inc., Mesa,
Arizona) loaded on a laptop computer. At the end of the data collection period, and
12
towards the end of the unit’s battery life, the GPS collars were programmed to release
from the animals. The collars were then retrieved in the field.
Elk capturing was timed so there was no overlap of a hunting season for up to 30
days due to toxic levels of immobilization drugs. This ensured that no hunter would
ingest any of the unmetabolized drugs. Collared elk did not receive exemption from
being harvested during hunting seasons. Mortalities attributed to harvest were included
in the study and documented and used in the survival analysis. The collars had contact
information and drug warning information attached.
Radio-Telemetry
The battery of the conventional VHF collar was expected to last at least 36
months but often lasted for several additional years. Relocations were obtained 1-4 times
per month. Relocations were obtained from a Cessna 172 aircraft fitted with 2-element
H-antennas (Telonics, Mesa, AZ), using standardized methods described by Mech
(1983). Most relocation surveys were conducted between 08:00 and 14:00 in an attempt
to capture elk when they may still be active. VHF coordinates were recorded with a GPS
receiver (Model Meridian, Magellan Corporation, Santa Clara, CA). VHF coordinates
were evaluated for accuracy by comparing fixes marked with the hand-held GPS unit
with the fix recorded on the GPS collar for the same day and time. Both the VHF and
GPS collared elk were used for analysis in this thesis.
Home Range and Core Use Areas
Locations on a total of 53 unique radio-collared elk were used for home range
analysis using the minimum convex polygon (Hayne 1949) estimator for overall home
range of each individual elk, and an estimate for the average home range for the entire
herd. The MCP estimator was also used to estimate home range area for each elk by
biological year. The fixed kernel method was used for home range estimates for 15 GPScollared elk (Figure 2 and Appendix 3). Home range estimates for the 15 elk at the 95%
and 50% contour levels were generated using the fixed kernel method with the Hawthes
Tools extension for ArcGIS (ArcGIS 9.3, ESRI Inc, Redlands, California, USA).
Approximately 80,000 locations were collected on the GPS-collared elk during the study
period. The large sample size allowed us to run the fixed kernel analysis on the GPS
collared elk. Fixed kernel home range analysis was the preferred method since it
13
provides a more accurate robust reflection of habitat use areas. However, the total
number of locations per year, and life period, for the VHF-collared elk was substantially
smaller. All data were normally distributed and had equal variance between groups.
Paired t-tests (Microsoft Excel 2007, Microsoft, Inc. Redmond, WA) were used to
determine significance in MCP home range estimates between migratory and nonmigratory elk.
Figure 2. Batch fixed kernel density for Elk #30 at the 50, 90 &95% range areas.
14
Cow Elk Survival
Cow elk survival rates were estimated for the radio-collared elk throughout the
period of the entire study using the binomial distribution method (White and Garrott,
1990), and the Kaplan-Meier Method (Kaplan and Meier 1958) with the Pollock
adaptation for staggered entry (Pollock et al. 1989). Estimates were made by biological
year which spanned June 1 – May 31st of the respective year. Annual estimates of
survival rates were made based on these data. No attempt was made to make annual
survival estimates per migratory status of group because sample sizes would have been
too small to provide meaningful information. Total number of migratory and nonmigratory collared elk mortalities combined per biological year ranged from 3-10 per
year.
Habitat Variables
The study area boundaries were divided into two separate sub-units based on
wintering locations of the sub-herds of the collared elk along the upper-Cowlitz river
basin using the minimum convex polygon of all GPS collared elk locations. The GPS
study group of elk consisted of 15 collared elk, four in the Randle unit and 11 in the
Packwood unit. The 15 GPS collared elk yielded >80,000 data points. The two sub-units
are referred to as the Packwood unit (679 km2) located in the most easterly portion of the
study area (Figure 3) and the Randle unit (349 km2), located west of the Packwood unit,
in and around the town of Randle (Figure 4). Habitat variables within each area included
slope, aspect, distance from evergreen/cover, preferred forage (shrub/scrub, meadow, and
pasture), presence of water, and roads. Slope and aspect layers were developed from
1:24,000 scale USGS 10 M Digital Elevation Model (DEM) data. Aspect was divided
into four categories: east/northeast, north/northwest, south/southwest, and
south/southeast. Slope was divided into five categories: 0-1% (flat), 1-15%, 16-30%, 3145%, >46%.
15
Figure 3. Packwood elk habitat analysis area.
National Land Cover Data (NCLD) was obtained and used to create habitat maps
of the study areas. Nineteen habitat variables were initially identified in the study areas.
These were later reduced to seven categories to reflect closely related habitat attributes
for analysis. As already mentioned, land type variables were divided into 7 major
categories: shrub/scrub/meadow/pasture, evergreen forest (cover), water (to include
16
riparian/deciduous areas), and roads. These categories represent critical habitat attributes
that are commonly analyzed in elk habitat suitability studies (Thomas et al 1979, Lyon
1983, Bracken and Musser 1993). Distance class for each category ranged between 2-6
distance classes per habitat type (Appendix 4). For example distance classes to water,
cover, and shrub scrub were: 0-50 m, 51-100 m, 101-200 m, 201-400 m 401-600 m, and
>600 m. Distance class to roads and trails were: 0-100 m, 101-200 m, 201-400 m, 400600 m, and >600 m. Paved roads included only 2 distance classes which were 0-600m
and >600m to roads. Distance classes used in our study were modeled after standard
distances classes used in other elk habitat analysis studies (Skovlin et.al 2002).
17
Figure 4. Randle elk habitat analysis area.
Once the habitat variables were selected, the land cover data set was ground
truthed by comparing aerial digital orthophotos to the land cover image/data set. The
entire image/data set was edited using the aerial orthophotos until a final image was
produced that most accurately reflected current habitat conditions of the study area.
Stream and road layers were obtained from the United States Forest Service (USFS),
18
Washington State Department of Natural Resources (WADNR) Lewis, Pierce, Yakima,
and Skamania Counties. Distance from GPS locations to the selected habitat features
were analyzed by using the near-analysis tool in the ArcGIS toolbox (ArcGIS 9.3, ESRI
Inc. Redlands, CA, USA) for proximity-to-features tool.
Habitat Use, Seasonal Habitat Use, and Time of Day
Habitat use and distances from roads, water, cover, and shrub/scrub were
analyzed at two spatial scales: 95% fixed kernel, and 50% fixed kernel. Paired t-tests
(Microsoft Excel 2007, Microsoft Inc. Redmond, WA) were used to determine if habitat
use varied between individual elk, and between migratory and non-migratory groups
beyond the range that would be expected by chance. Data were analyzed using a
student’s t-test with p≤0.05used to denote significance. Analysis where p=0.10 were also
included in the analysis to provide supplemental insight into the primary analysis. The ttests in this study identify areas where the habitat selected by the elk occurred beyond the
range that would be expected by chance. The focus of this study analyzed differences
between migratory and non-migratory groups of elk. Differences between individual elk
were included in the study, but they were not examined in detail.
Seasonal and time of day variables were also evaluated in the analysis. Elk locations
were divided into four biologically significant seasons: Spring/calving (March-June),
Summer (July-August), Fall/rut (September-November), and Winter (DecemberFebruary). Time of day was also evaluated, and divided into four intervals based on
observed activity: dawn (0500-0900), diurnal (0900-1700), dusk (1700-2100), and
nocturnal (2100-0500). Ordinal regression (SPSS 17.0, SPSS Inc., Chicago, Ill) was used
to determine whether season and season and time of day influenced habitat use between
individual elk and/or migratory versus non-migratory elk. The data set used for the elk
study had taken the physical distances of the elk from given land features (scaler
variables), and grouped then into distance classes (becoming ordinal variables). It was
therefore necessary to conduct ordinal regression (as opposed to linear or non-linear
regression) in modeling the behavior of the elk. Ordinal regression in SPSS allows a
choice from among five different link functions (mathematical transforms) to optimize
the fitting to the data, and calculates a pseudo R2 as a rough estimate of the probability of
picking the correct distance class for a given individual elk, or group of elk, given the
season and/or time of day.
19
RESULTS
Elk Captures
Fifty-three elk were captured and radio-collared during the month of March in
years 2004-2008 respectively. Thirty-nine VHF collars were deployed years 2004-2007,
4 GPS collars were deployed in 2007. The final 11 GPS collars were deployed in March
2008 (Appendix 5). All cow elk were determined to be in relatively good condition by
the project veterinarian (Briggs Hall, DVM) with the exception of the elk captured in
2008. The elk captured in 2008 were in below moderate - poor condition due to
unusually harsh winter conditions (National Weather Service 2010). Special note of
significance were two GPS collared elk mortalities, for which the cause of mortality was
unknown, which occurred in May of 2008. Consequently, the bulk of the data that were
available for habitat use analysis via the GPS data was reduced to 13 GPS collared elk.
No diseases were detected. Genetic testing indicated that the elk were a hybrid
mixture of Rocky Mountain elk (C.e. nelson: translocated elk) and Roosevelt elk (C.e.
roosevelti: native to WA).
Home Range
Home range estimates were made using 53 elk starting where there were > 10
locations (Appendix 6). Comparisons of mean home range size between migratory and
non-migratory elk groups per biological year demonstrated differences between the
groups per biological year (Table 1). Life home range also demonstrated differences
between the migratory and non-migratory elk (Table 1).
20
Table 1. Minimum Convex Polygon (MCP) home range estimates for migratory and
non-migratory elk for years 2004 – 2008 and life home range comparisons for Western
Washington elk in the upper Cowlitz River valley.
Migratory
Status
Mean MCP Km2 Area
Mean MCP Hectares Area
Sample
size (N)
2004
Nonmigratory
8.4
840
3
2004
Migratory
58
5800
14
2005
Nonmigratory
5.4
540
7
2005
Migratory
54
5400
15
2006
Nonmigratory
11
1100
5
2006
Migratory
54
5400
23
2007
Nonmigratory
11
1100
8
2007
Migratory
55
5500
27
2008
Nonmigratory
14
1400
8
2008
Migratory
75
7500
24
20042008
Nonmigratory
20
2000
18
20042008
Migratory
102
10200
36
20042008
Combined
61
6100
54
Year
Home range size was estimated using the Minimum Convex Polygon (MCP) where the
mean area for all elk was 6100 ha (61 km2). Whereas MCP mean for migratory elk was
10200 ha (102 km2) and 2000 ha (20 km2) for non-migratory elk.
Differences were detected between migratory and non-migratory elk for MCP
home range estimates both by individual biological years (BYR), and at the life home
range area scale. Significant differences were detected in BYR 2005 in MCP home range
21
estimates between migratory elk and non-migratory elk (t18 = -6.46, 95%). Differences
were also detected in BYR 2OO6 in MCP home range estimates between migratory elk
and non-migratory elk (t26 = -3.17, 95%). Additionally, differences were detected in
BYR 2OO7 in MCP home range estimates between migratory elk and non-migratory elk
(t30 = -4.64, 95%). Finally, differences were detected in BYR 2OO8 in MCP home range
estimates between migratory elk and non-migratory elk (t28 = -3.167, 95%). Overall, the
most compelling differences were detected in the aggregate MCP life home range
estimates between migratory elk and non-migratory elk (t50 = -5.45, 95%). Differences
may be viewed spatially in Figure 5, Figure 6, Figure 7, and Appendix 16.
22
Figure 5. GPS collared elk data points per migratory status in the Packwood subunit
study area.
23
Migration
The data showed that approximately 66% of the herd was migratory, and 34% of
the herd was non-migratory. The data also showed over the 4-year period of the study
that the radio-collared elk demonstrated high range fidelity (99%). Study results showed
99% site fidelity of cow elk among summer ranges and winter ranges. This study showed
not only range fidelity but considerable variation in migratory routes and summer range
destinations.
Distribution
Packwood Sub-Herd
The sub-herd wintering in the Packwood area, while showing high fidelity to
seasonal range areas, the sub-herd as a whole showed quite a bit of variation in annual
migration routes and summering destinations. An average of 20% of the Packwood subherd migrated annually to areas south of Packwood in the Gifford Pinchot National
Forest, with destinations reaching into the Goat Rocks Wilderness Area. Routes and
destinations included the following: Lake Creek/Snyder Ridge to Johnson Basin, Lake
Creek/Snyder Ridge to Lily Basin, Lake Creek/Snyder Ridge to Lost Lake/Beargrass
Butte, Johnson Creek to Heart Lake Basin, Johnson Creek to Jordan Basin, Johnson
Creek/Deception Creek to Cold Springs Butte, and Smith Creek to Twin Sisters/Castle
Butte (Figure 6).
An average of 46% of the sub-herd migrated to areas north of Packwood into
Gifford Pinchot National Forest, the Tatoosh Wilderness Area, and Mount Rainier
National Park (MRNP). Many of the elk that migrated to MRNP used similar migration
routes into the Park even though their final destinations for summering varied. Routes
and destinations included the following: Cowlitz Muddy Fork to Blue Lake area, Cowlitz
Muddy Fork to Backbone Ridge area, Cowlitz Muddy Fork to Stevens Ridge area
(MRNP), Cowlitz Muddy Fork to Fan Lake Area (MRNP), Cowlitz Muddy Fork to
Cowlitz Park area (MRNP), Cowlitz Muddy Fork to Ohanapekosh Park area (MRNP),
Cowlitz Muddy Fork to Bald Rock area (MRNP), Cowlitz Muddy Fork to Shriner Peak
area, Butter Creek to Johnson Lake area (MRNP), and Skate Creek/Bear Prairie to
Lookout Mountain area (Figure 6).
24
Figure 6. Elk distribution in the Packwood study area. Color coded triangles
represent individual elk.
25
Randle Sub-Herd
Five elk were collared in the Randle area in years 2006 and 2008, one with a
VHF radio-collar and the other four with GPS collars. Three of the five elk were nonmigratory; one of the non-migratory elk was fitted with a VHF collar. The non-migratory
VHF radio-collared elk that wintered in the Randle area along the Cowlitz River showed
range fidelity year to year. Collared elk migrated up the Iron Creek drainage towards Mt.
St. Helens to the Strawberry Mountain area and to the Boot and Grizzly Lake areas
within the Mt. St. Helens National Monument area (Figure 7).
26
Figure 7. Elk distribution in the Randle study area. Color coded triangles
represent individual elk.
27
Survival Rates
Known causes of mortality included legal harvest, auto collision, predator kills,
starvation, and calving complications. Survival rates for 2004 using the bionomial
estimator method were 74%, 2005 rates were 88%, and 2006 rates were 82%. In 2007
the survival rate using the Kaplen-Meier method was 71% and using the same method in
2008, the survival rate was 90%.
Habitat Use
Activity: 50% Range Analysis
Overall, migratory (M) elk were inactive/bedded 51.5% of the time, while nonmigratory (NM) elk were inactive/bedded 53% of the time. The remainder of the time M
elk were active/feeding 48.5% of the time, while the NM elk were active/feeding 47% of
the time. Significant differences were detected between individual elk (which represented
family/social groups of ~ 15 elk) and the migratory group as well as the overall study
group of elk (Figure 8). Non-migratory elk #29 was less active than the other nonmigratory elk (t13 = 0.039, p >.05) and more active than all of the elk in the study group
(t13 = 0.09, p > 0.05). Similarly, M elk #30 was less active than both the migratory group
(t13 = 0.018, p > .05) and the entire study group (t13 = 0.02, p > .05).
28
Figure 8. Activity categories represent number of collar movements per hour for elk in
the combined Randle and Packwood study areas in the Southwest Cascades of
Washington 2007-2008.
Activity 95% Range Analysis
Similar to the results for the 50% range analysis, non-migratoryNM elk were
inactive/bedded slightly more of the time (52%) than their migratory counter-parts (50%).
Differences were detected between two individual elk and their respective migratory
group and the overall study group. Non-migratory elk #29 was less active than the other
non-migratory elk (t13 = 0.011, p > .05) and also less active than all of the elk in the study
29
group (t13 = 0.034, p > .05). Similarly, M elk #30 was less active than both the respective
migratory group (t13 = 0.011, p > .05) and the entire study group (t13 = 0.028, p > .05).
Aspect 50% Range Analysis
Migratory elk spent 11% of their time in habitat that is considered flat, where
NM elk spent 19% of their time in the flat habitat type. Annual breakdowns of percent
use areas per migratory group may be viewed in Table 2. Migratory elk were found to
use the SW aspect significantly less than the NM elk group (t13 = 0.035, p < .10).
Seasonally, M and NM elk spent a greater percentage of their time in southern aspects
during all four seasons. (Appendix 7).
Table 2. Overall annual habitat use of aspects by elk in the combined Randle and
Packwood study areas at the 50% and 95% range scales in the Southwest Cascades of
Washington. 2007-2008.
50% Contour
Kernel
Normalized
Aspect Category
Individual
Flat (-1-0)
NE (0-90)
SE (90180)
SW(180270)
NW(270360)
Migratory
11%
16%
32%
16%
25%
Nonmigratory
19%
11%
24%
28%
18%
Both
13%
15%
30%
19%
23%
95% Contour
Kernel
Normalized
Aspect Category
Individual
Flat (-1-0)
NE (0-90)
SE (90180)
SW(180270)
NW(270360)
Migratory
8%
18%
31%
21%
23%
Nonmigratory
15%
12%
24%
25%
23%
Both
9%
17%
29%
22%
23%
Several individual elk yielded differences in time spent in particular aspects than
their respective migratory group and the entire study group of elk. Elk #28 was found to
30
use the SW aspect less than the elk in its migratory group (t13 = 0.002, p < .10).
Migratory Elk #28 was found to use the NW aspect more than the elk in its migratory
group (t13 = 0.015, p > .05) and the overall study group of elk (t13 = 0.033, p > .05). Nonmigratory Elk #29 used flat terrain more than elk in the entire study group (t13 = 0.036, p
> .05). Non-migratory elk #81 used the flat aspect more than the elk in the entire study
group (t13 = 0.016, p > .05). Migratory elk #84 used the NW aspect less than the elk in
the entire study group (t13 = 0.075, p > .10). Non-migratory elk #419 spent more time in
the SW aspect than the elk in its migratory group (t13 = 0.083, p > .10) as well as the
entire study group of elk (t13 = 0.002, p > .05). Non-migratory elk #420 spent more time
in the NE aspect than the elk in its migratory group (t13 = 0.012, p > .10; she also spent
more time in the NW aspect than her migratory group (t13 = 0.012, p > .05). Migratory
elk #421 used the SW aspect more than the elk in her migratory group (t13 = 0.090, p >
.10). Migratory elk # 422 used the NE aspect less than the elk in the entire study group
(t13 = 0.059, p > .10).
Aspect 95% Range Analysis
In the 95% range analysis, M elk spent 8% of their time in habitat that is
considered flat, where NM elk spent 15% of their time in the flat habitat type. Annual
breakdowns of percent use areas per migratory group may be viewed in Table 2.
Seasonally, migratory and non-migratory elk spent a greater percentage of their time in
southern aspects during all four seasons (Appendix 7).
Again, several individual elk yielded differences in time spent in particular
aspects than their respective migratory group and the entire study group of elk.
Migratory elk #28 was found to use the NW aspect more than the elk in her migratory
group (t13 = 0.035, p > .05) and the overall study group of elk (t13 = 0.028, p > .05). Nonmigratory elk #29 used flat terrain more than elk her entire study group (t13 = 0.065, p >
.10). Non-migratory elk #81 used the flat aspect more than the elk in the entire study
group (t13 = 0.002, p > .05). Non-migratory elk #85 used the SE aspect more than the elk
in her migratory group (t13 = 0.063, p > .10) as well as the entire group of elk (t13 = 0.062,
p > .10). Non-migratory elk #419 spent more time in the SW aspect than the elk in her
migratory group (t13 = 0.066, p > .10) as well as the entire study group of elk (t13 = 0.001,
p > .05). Non-migratory elk #420 spent more time in the NE aspect than the elk in her
migratory group (t13 = 0.075, p > .10) as well as more time in the NW aspect than its
31
migratory group (t13 = 0.023, p > .05). Migratory elk #421 used the NE aspect more than
the elk the entire study group of elk (t13 = 0.076, p > .10). Migratory elk # 422 used the
NE aspect less than the elk in the entire study group (t13 = 0.035, p > .05) as well as the
elk within her migratory group (t13 = 0.094, p > .10).
Cover 50% Range Analysis
Significant differences between the migratory and non-migratory groups were
observed in the 50-100 m distance category where NM elk were in this distance class
more than M elk (t13 = 0.050, p > .05). Migratory elk spent 51% of their time within the
0-50 m distance class of cover where NM elk spent 35% of their time within the same
distance class. Remaining distance class to cover comparisons generally also showed
that NM elk spent more time in the lower distance classes (Table 3). During winter
migratory elk spent 57% of their time within 50 m of cover as compared to nonmigratory elk which spent 34% of their time within 50 m of cover (Figure 9 and
Appendix 8).
32
Figure 9. Distance to cover analysis for migratory and non-migratory elk at the 50%
scale in the Southwestern Cascades of Washington.
Percentage of Observations
Ordinal Regression Model for Cover Distance (Meters) Category (Migratory Elk
/ 50% Contour Kernal)
150%
100%
50%
0%
Winter
Spring
Summer
Fall
Cover Distance Category
0-50
50-100
100-200
200-400
400-600
>600
Percentage of Observations
Ordinal Regression Model for Cover Distance (Meters) Category (NonMigratory Elk / 50% Contour Kernal)
150%
100%
50%
0%
Winter
Spring
Summer
Fall
Cover Distance Category
0-50
50-100
100-200
200-400
400-600
>600
33
Table 3. Overall annual habitat use by elk relative to distance to cover in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008.
Distance from Cover (meters)
Category
50% Contour Kernel
Individual
0-50
50-100
100-200
200-400
400-600
>600
Migratory
51%
17%
21%
9%
2%
0%
Nonmigratory
35%
25%
32%
8%
0%
0%
Both
47%
19%
24%
9%
1%
0%
Distance from Cover (meters)
Category
95% Contour Kernel
Individual
0-50
50-100
100-200
200-400
400-600
>600
Migratory
56%
16%
18%
8%
1%
0%
Nonmigratory
44%
22%
26%
8%
0%
0%
Both
53%
17%
20%
8%
1%
0%
Several individual elk revealed significant differences with respect to distance
class to cover with their respective migratory group as well as the entire study group of
elk. Non-migratory elk #29 spent more time in the 100-200 m distance to cover category
than the entire study group of elk (t13 = 0.087, p > .10). Migratory elk #30 spent more
time in the 50-100 m distance category than her respective migratory group (t13 = 0.017, p
> .05). Non-migratory elk #81 used the 50-100 m distance category more than the elk in
the entire study group (t13 = 0.051, p > .10). Migratory elk #82 was found in the 0-50 m
category more than elk in her respective migratory group (t13 = 0.031, p > .05) and the
study group of elk as a whole (t13 = 0.046, p > .05). The same elk M #82 was found in
the 100-200 m distance class less than elk in her migratory group (t13 = 0.008, p < .05)
and entire study group of elk (t13 = 0.040, p < .05). Non-migratory elk #85 was found in
the 0-50 m distance class more than her migratory group (t13 = 0.051, p > .10). The same
elk, #85 was in the 50-100 m distance class (t13 = 0.079, p < .10) and the 100-200 m
distance class (t13 = 0.079, p < .10) less than elk in her migratory group. Migratory elk
#421 spent less time in the 0-50 m distance category than the other elk in her migratory
group (t13 = 0.044, p < .05) However, M elk #421 spent significantly more time in the
200-400 m distance class than the elk in her migratory group (t13 = 0.002, p > .05) and the
34
elk in the entire study group (t13 = 0.002, p > .05). The same elk (#421) was also found in
the 400-600 m distance category more than the elk in her migratory group (t13 = 0.002, p
> .05), and the elk in the entire study group (t13 = 0.002, p > .05).
Cover 95% Range Analysis
Significant differences between the migratory and non-migratory groups were
observed in the 50-100 m distance category where migratory elk were in this distance
class less than non-migratory elk (t13 = 0.017, p < .05). Migratory elk spent 56% of their
time within the 0-50 m distance class of cover where NM elk spent 44% of their time
within the same distance class. Remaining distance class to cover comparisons may be
viewed in Table 3. Seasonally, during winter migratory elk spent 61% of their time
within 50 m of cover as compared to non-migratory elk which spent 39% of their time
within 50 m of cover (Figure 10 and Appendix 8).
35
Figure 10. Distance to cover analysis for migratory and non-migratory elk at the 95%
scale in the Southwestern Cascades of Washington.
Percentage of
Observations
Ordinal Regression Model for Cover Distance (meters) Category
(Migratory Elk / 95% Contour Kernal)
150%
100%
50%
0%
Winter
Spring
Summer
Fall
Cover Distance Category
0-50
50-100
100-200
200-400
400-600
>600
Percentage of
Observations
Ordinal Regression Model for Cover Distance Category (Meters)
(Non-Migratory Elk / 95% Contour Kernal)
150%
100%
50%
0%
Winter
Spring
Summer
Fall
Cover Distance Category
0-50
50-100
100-200
200-400
400-600
>600
Several individual elk revealed significant differences with respect to distance
class to cover and their respective migratory group as well as the entire study group of
elk. Non-migratory elk #29 spent more time in the 100-200 m distance to cover category
than the entire study group of elk (t13 = 0.041, p > .05). Migratory elk #30 spent more
time in the 50-100 m distance to cover category more than her respective migratory group
(t13 = 0.058, p > .10). Non-migratory elk #81 used the 0-50 m distance to cover category
less than the elk in the entire study group (t13 = 0.050, p < .05). The same elk #81 used
the 50-100 m distance to cover category more than her migratory group (t13 = 0.055, p >
.10) and the entire study group of elk (t13 = 0.042, p > .05). Migratory elk #82 was found
in the 0-50 m category more than elk in her respective migratory group (t13 = 0.072, p >
.10) and the study group of elk as a whole (t13 = 0.072, p > .10). The same elk M #82
was found in the 100-200 m distance class less than the elk in her migratory group (t13 =
36
0.026, p < .05) and entire study group of elk (t13 = 0.081, p > .10). Non-migratory elk
#85 was found in the 50-100 m distance class less than the entire study group of elk (t13 =
0.092, p < .10). Migratory elk #421 spent less time in the 0-50 m distance category than
the other elk in her migratory group (t13 = 0.049, p < .05) However, M elk #421 spent
more time in the 200-400 m distance class than the elk in her migratory group (t13 =
0.001, p > .05) and the elk in the entire study group (t13 = 0.006, p > .05). The same elk
(421) was also found in the 400-600 m distance category more than the elk in her
migratory group (t13 = 0.000, p > .05), and the elk in the entire study group (t13 = 0.000, p
> .05).
Shrub/Scrub 50% Range Analysis
Migratory elk spent 65% of their time in the 0-50 m range of shrub/scrub while
NM elk spent 58% of their time in the same distance class. Remaining distance class to
shrub/scrub comparisons may be viewed in Table 4. Seasonal break downs by
percentage of data points in each distance class may be viewed in Appendix 9.
Table 4. Overall annual habitat use by elk relative to distance to shrub/scrub in the
combined Randle and Packwood study areas at the 50% and 95% range scale in the
Southwestern Cascades of Washington, 2007-2008.
50%
Contour
Kernel
Individual
Distance (Meters) from Shrub/Scrub
Category
0-50
50-100
100-200
200-400
400-600
>600
Migratory
65%
10%
14%
8%
2%
1%
Nonmigratory
58%
11%
17%
14%
1%
0%
Both
63%
10%
15%
10%
1%
1%
95% Contour Kernel
Normalized
Individual
Distance (Meters) from Shrub/Scrub
Category
0-50
50-100
100-200
200-400
400-600
>600
Migratory
62%
9%
14%
10%
3%
3%
Nonmigratory
55%
12%
19%
13%
1%
0%
Both
60%
10%
15%
10%
2%
2%
37
Many individual elk showed significant differences between distance to
shrub/scrub habitat in comparison to their migratory group and the entire study group of
elk. Migratory elk #80 spent more time in the 400-600 m distance from shrub/scrub than
the elk in her migratory group (t13 = 0.000, p > .05) and the entire study group of elk (t13
= 0.000, p > .05). Non-migratory elk # 81 spent less time in the 0-50 m distance class to
shrub/scrub than the elk in the entire study group (t13 = 0.064, p > .10). The same elk #81
spent more time in both the 50-100 m (t13 = 0.018, p > .05) and 100-200 m distance
classes than the elk in the entire study group (t13 = 0.062, p > .10). Migratory elk #83
spent more time in the >600 m distance class to shrub/scrub than the elk in her migratory
group (t13 = 0.000, p > .05) and the entire study group of elk (t13 = 0.000, p > .05). Nonmigratory elk #84 spent less time in the 0-50 m distance class from shrub/scrub than the
migratory elk in her group (t13 = 0.063, p > .10). The same elk #84 spent more time in
the 100-200 m distance category to shrub/scrub than the elk in her migratory group (t13 =
0.008, p > .05) and the elk in the entire study group of elk (t13 = 0.043, p > .05). Nonmigratory elk #420 spent more time in the 200-400 m distance to shrub/scrub category
than the entire study group of elk (t13 = 0.040, p > .05).
Shrub/Scrub 95% Range Analysis
Some differences were detected between the two groups of elk where M elk were
found to be in the > 600 m distance class from shrub/scrub more than NM elk (t13 =
0.056, p > .10). Migratory elk spent 62% of their time in the 0-50 m range of shrub/scrub
while NM elk spent 55% of their time in the same distance class. Remaining distance
class to shrub/scrub comparisons may be viewed in Table 4. Seasonal break downs by
percentage of data points in each distance class may be viewed in Figure 11 and
Appendix 9.
38
Figure 11. Distance to shrub/scrub analysis for migratory and non-migratory elk in the
Southwestern Cascades of Washington.
Percentage of Observations
Ordinal Regression Model for Shrub/Scrub Distance (Meters) Category
(Migratory Elk / 95% Contour Kernal)
150%
100%
50%
0%
Winter
Spring
Summer
Fall
Shrub/Scrub Distance Category
0-50
50-100
100-200
200-400
400-600
>600
Percentage of Observations
Ordinal Regression Model for Shrub/Scrub Distance (Meters) Category (NonMigratory Elk / 95% Contour Kernal)
150%
100%
50%
0%
Winter
Spring
Summer
Fall
Shrub/Scrub Distance Category
0-50
50-100
100-200
200-400
400-600
>600
Many individual elk showed significant differences between distance classes to
shrub/scrub habitat in comparison to their migratory group and the entire study group of
elk. Migratory elk #30 spent less time in the 200-400 m distance class than the elk in her
migratory group (t13 = 0.086, p < .10). Migratory elk #80 spent more time in the 400-600
m distance from shrub/scrub than the elk in her migratory group (t13 = 0.010, p > .05) and
the entire study group of elk (t13 = 0.004, p > .05). Non-migratory elk # 81 spent less
time in the 0-50 m distance class to shrub/scrub than the elk in the entire study group (t13
= 0.053, p < .10). The same elk #81 spent more time in both the 50-100 m distance class
(t13 = 0.009, p > .05) and 100-200 m distance classes than the elk in the entire study
39
group (t13 = 0.033, p > .05). Non-migratory elk #84 spent less time in the 0-50 m
distance class from shrub/scrub than the entire group of elk in the study (t13 = 0.051, p <
.10). The same elk #84 spent more time in the 100-200 m distance category to
shrub/scrub than the elk in her migratory group (t13 = 0.011, p > .05) and the elk in the
entire study group (t13 = 0.091, p > .10). Non-migratory elk #419 spent more time in the
>600 m distance category than the elk in the entire study group (t13 = 0.008, p > .05).
Non-migratory elk #420 spent less time in the 0-50 m distance category than all of the elk
in the entire study group (t13 = 0.093, p > .10). The same elk #420 spent more time in the
200-400 m distance to shrub/scrub category than the entire study group of elk (t13 =
0.031, p > .05). Elk # 420 also spent more time in the 400-600 m distance category than
the elk in her migratory group (t13 = 0.000, p > .05). Migratory elk #421 spent less time
in the 50-100 m distance category than the elk in her migratory group (t13 = 0.088, p <
.10). Migratory elk #422 spent more time in the 400-600 m distance class than both the
migratory group (t13 = 0.019, p > .05) and the entire study group of elk (t13 = 0.002, p >
.05).
Slope 50% Range Analysis
Significant differences were detected between the migratory and non-migratory
groups of elk with regard to slope preferences. The differences that were detected
between the two groups were where the migratory elk utilized 1-15% slope areas less
than the migratory elk (t13 = 0.089, p < .10). Also, migratory elk preferred steeper 1530% slopes (t13 = 0.082, p > .10) and >45% slopes more than non-migratory elk (t13 =
0.053, p > .10). Overall, migratory elk spent 10% of their time within the 0-1% slope
class where NM elk spent 18% of their time within the same slope class. Remaining
slope class comparisons may be viewed in Table 5. Seasonally, migratory elk also spent
a greater percentage of their time on steeper slopes comparatively than non-migratory elk
(Figure 12-13 and Appendix 10).
40
Table 5. Annual overall habitat use by elk of slopes in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern Cascades of
Washington, 2007-2008.
50% Contour Kernel
Normalized
Individual
Percent Slope
Category
0-1
1-15
15-30
30-45
45-25000
Migratory
10%
47%
20%
12%
11%
Nonmigratory
18%
61%
10%
7%
3%
Both
12%
51%
18%
11%
9%
95% Contour Kernel
Normalized
Individual
Percent Slope
Category
0-1
1-15
15-30
30-45
45-25000
Migratory
7%
40%
22%
16%
16%
Nonmigratory
14%
62%
14%
7%
3%
Both
9%
45%
20%
13%
13%
Several individual elk yielded significant differences in their preferences of
particular slope classes than their respective migratory group and the entire study group
of elk. Migratory elk #28 preferred 15-30% slope areas more than the entire study group
of elk (t13 = 0.023, p > .05) and more than the elk in her migratory elk group (t13 = 0.058,
p > .10). Non-migratory elk #29 preferred flat sloped areas of 0-1% grades more than the
elk in the entire study group (t13 = 0.033, p > .05). Non-migratory elk #81 also preferred
flat sloped areas of 0-1% grades more than the rest of the elk in the entire study group (t13
= 0.014, p > .05). Non-migratory elk #85 preferred 1-15% slopes more than the entire
study group of elk (t13 = 0.063, p > .10). Whereas NM Elk #419 preferred 30-45% slopes
(t13 = 0.014, p > .05) and >45% slopes more than the elk in her non-migratory group (t13 =
0.028, p > .05). Migratory elk #421 showed less preference for 1-15% slope areas than
all of the elk in the study group (t13 = 0.079, p < .10). The same elk (#421) showed
greater preference for slope areas >45% than the elk in both her migratory group (t13 =
0.033, p > .05) and the entire study group of elk (t13 = 0.006, p > .05). Migratory elk
#422 was less likely to be found using areas with 1-15% slope than the elk in the entire
study group (t13 = 0.084, p < .10). The same elk (#422) was found to prefer 30-45%
41
slopes more than elk in her migratory group (t13 = 0.008, p > .05) and the entire study
group of elk (t13 = 0.009, p > .05).
Slope 95% Range Analysis
More pronounced differences were detected between the migratory and nonmigratory groups of elk with regard to slope preferences at the 95% range areas. Highly
significant differences were detected between the two groups where migratory elk
utilized 1-15% slope areas less than migratory elk (t13 = 0.004, p < .05). As well, highly
significant differences were detected where migratory elk preferred steeper 15-30%
slopes (t13 = 0.047, p > .05), 30-45% slopes (t13 = 0.008, p > .05) and >45% slopes more
than non-migratory elk (t13 = 0.000, p > .05). Overall, migratory elk spent 7% of their
time within the 0-1% slope class where NM elk spent 14% of their time within the same
slope class. Remaining slope class comparisons may be viewed in Table 5. Seasonally,
migratory elk also spent a greater percentage of their time on steeper slopes
comparatively than non-migratory elk (Figure 12-13 and Appendix 10).
42
Figure 12: Migratory elk habitat selection with slope analysis.
60.00%
l
o
c
a
t
i
o
n
s
Migratory Elk Slope Preferences
50.00%
40.00%
30.00%
Winter
20.00%
Spring
10.00%
Summer
0.00%
Fall
Slope
Figure 13. Non-migratory elk habitat selection with slope analysis.
70.00%
l
o
c
a
t
i
o
n
s
Non-migratory Elk Slope Preferences
60.00%
50.00%
40.00%
30.00%
Winter
20.00%
Spring
10.00%
Summer
0.00%
Fall
Slope
Several individual elk yielded differences in preferences of particular slope
classes than their respective migratory group and the entire study group of elk. Nonmigratory elk #29 preferred flat sloped areas of 0-1% grades more than the elk in the
entire study group (t13 = 0.061, p > .10). Migratory elk #30 showed a preference to flat 01% sloped areas than the other elk in her respective migratory group (t13 = 0.087, p >
.10). Non-migratory elk #81 also preferred flat sloped areas of 0-1% grades more than
the rest of the elk in the entire study group (t13 = 0.002, p > .05). The same elk (#81)
43
showed less preference for 1-15% slope areas than elk in her migratory group (t13 =
0.053, p < .10) and the entire study group of elk (t13 = 0.035, p < .05). Migratory elk #82
showed a greater preference for 1-15% slope areas than the elk in her respective
migratory group (t13 = 0.078, p >.10). Non-migratory elk #85 preferred 1-15% slopes
more than the entire study group of elk (t13 = 0.043, p > .05). Whereas Non-migratory
elk #419 preferred 30-45% slopes (t13 = 0.020, p > .05) and >45% slopes more than the
elk in her non-migratory group (t13 = 0.045, p > .05). Migratory elk #421 showed greater
preference for slope areas >45% than the elk in both her migratory group (t13 = 0.031, p >
.05) and the entire study group of elk (t13 = 0.022, p > .05). Migratory elk #422 was less
likely to be found using areas with 1-15% slope than the elk in the entire study group (t13
= 0.082, p < .10). The same elk (#422) was found to prefer 30-45% slopes more than elk
in her migratory group (t13 = 0.001, p > .05) and the entire study group of elk (t13 = 0.008,
p > .05).
Water 50% Range Analysis
Migratory elk spent 12% of their time in the 0-50 m range of water while NM elk
spent 11% of their time in the same distance class. Remaining distance class to water
comparisons may be viewed in Table 6. Seasonally, migratory elk generally were not
found in distances greater than 600 m to water (8%) whereas non-migratory elk spent
greater percentages of time at distances greater than 600 m on average 28% of the time
(Appendix 11). On average, migratory elk spent 8% of their time at distances greater
than 600 m from water.
44
Table 6. Overall annual habitat use by elk relative to distance to water in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008.
50% Contour Kernel
Normalized
Distance (Meters)from Water
Category
Individual
0-50
50-100
100-200
200-400
400-600
>600
Migratory
12%
11%
22%
27%
16%
12%
Non-migratory
11%
9%
18%
25%
12%
26%
Both
12%
11%
21%
26%
15%
15%
95% Contour Kernel
Normalized
Distance (Meters) from Water
Category
Individual
0-50
50-100
100-200
200-400
400-600
>600
Migratory
11%
10%
20%
26%
16%
17%
Non-migratory
12%
10%
18%
23%
13%
25%
Both
11%
10%
20%
25%
15%
19%
Many individual elk showed significant differences between distance classes to
water habitat in comparison to their migratory group and the entire study group of elk.
Non-migratory elk #29 showed a preference for spending more time in the 200-400 m
distance class to water than all of the elk in the entire study group (t13 = 0.048, p > .05).
The same elk (#29) spent more time in the 400-600 m distance category than other elk in
her respective migratory group (t13 = 0.093, p > .10). Migratory elk #30 spent more time
in the 0-50 m distance category than both her migratory group (t13 = 0.032, p > .05) and
the entire study group of elk (t13 = 0.050, p > .10). The same elk (#30) also spent more
time in the 50-100 m distance category than both her migratory group (t13 = 0.060, p >
.10) and the entire study group of elk (t13 = 0.051, p > .10). Migratory elk #31 spent less
time in the 100-200 m distance category than the other elk in her migratory group (t13 =
0.041, p < .05). The same elk (#31) also spent more time in the 400-600 m distance
category than both her migratory group (t13 = 0.045, p > .05) and the entire study group of
elk (t13 = 0.033, p > .05). Elk #31 also spent more time in the >600 m distance category
than the other elk in her migratory group (t13 = 0.033, p > .05). Non-migratory elk #80
spent more time in the 200-400 m distance category than the other elk in her migratory
group (t13 = 0.086, p > .10). Migratory elk #83 spent less time in the 200-400 m category
than the elk in her migratory group (t13 = 0.096, p < .10). Non-migratory elk #419 spent
45
more time in the >600 m category than both her respective migratory group (t13 = 0.000,
p > .05) and the elk in the entire study group (t13 = 0.000, p > .05). Non-migratory elk
#420 spent more time in the 0-50 m distance category than both the elk in her migratory
group (t13 = 0.017, p > .05) and the entire study group of elk (t13 = 0.039, p > .05). The
same elk (#420) also spent more time in the 50-100 m distance category than the other
elk in her migratory group (t13 = 0.092, p > .10). Finally, M Elk #422 spent more time in
the 400-600 m distance category than all of the elk in the entire study group (t13 = 0.079,
p > .10).
Water 95% Range Analysis
Migratory elk spent 11% of their time in the 0-50 m range of water while NM elk
spent 12% of their time in the same distance class. Remaining distance class to water
comparisons may be viewed in Table 6. Seasonally, migratory elk generally were not
found in distances greater than 600 m from water whereas non-migratory elk spent
approximately 25% of their time at distances greater than 600 m (Appendix 11).
Many individual elk showed significant differences between distance classes to
water habitat in comparison to their migratory group and the entire study group of elk.
Migratory elk #30 spent more time in the 0-50 m distance category than her migratory
group (t13 = 0.061, p > .10). The same elk (#30) also spent more time in the 50-100 m
distance category than the elk in her migratory group (t13 = 0.096, p > .10). Migratory elk
#31 spent less time in the 100-200 m distance category than the other elk in her migratory
group (t13 = 0.036, p < .05). The same elk (#31) also spent more time in the 400-600 m
distance category than both her migratory group (t13 = 0.042, p > .05) and the entire study
group of elk (t13 = 0.021, p > .05). Elk #31 also spent more time in the >600 m distance
category than the other elk in her migratory group (t13 = 0.053, p > .10). Non-migratory
elk #85 spent more time in the 100-200 m distance class than other elk in her migratory
group (t13 = 0.006, p > .05). Non-migratory elk # 419 spent less time in the 50-100 m
distance category than all of the elk in the entire study group (t13 = 0.086, p < .10). The
same elk spent less time in the 100-200 m category than all of the elk in the study group
(t13 = 0.083, p < .10). Elk #419 also spent less time in the 200-400 m category than all of
the elk in the study group (t13 = 0.032, p < .05). In addition, NM Elk #419 spent more
time in the >600 m category than both her respective migratory group (t13 = 0.001, p >
.05) and the elk in the entire study group (t13 = 0.000, p > .05). Non-migratory elk #420
46
spent more time in the 0-50 m distance category than both the elk in her migratory group
(t13 = 0.051, p > .10) and the entire study group of elk (t13 = 0.024, p > .05). The same elk
(#420) also spent more time in the 50-100 m distance category than all of the elk in the
entire study group (t13 = 0.096, p > .10). Finally, M Elk #422 spent more time in the 400600 m distance category than all of the elk in the entire study group (t13 = 0.065, p > .10).
Distance to Paved Roads 50% Range Areas
Overall, migratory elk were found to be within 600 m of paved roads 54% of the
time whereas non-migratory elk were in the same distance class 64% of the time. The
opposite was true for time spent in the >600 m of paved roads where migratory elk spent
47% of their time in the distance class and non-migratory spent 37% of their time in the
>600 m distance class (Table 7). Seasonal break downs by percentage of data points in
each distance class may be viewed in Appendix 12.
Table 7. Overall annual habitat use by elk relative to distance to paved roads in the
combined Randle and Packwood study areas at the 50% and 95% range scale in the
Southwestern Cascades of Washington, 2007-2008.
50% Contour Kernel
Normalized
Individual
Distance (Meters)from
Paved Road Category
0-600
>600
Migratory
54%
46%
Non-migratory
64%
36%
Both
56%
44%
95% Contour Kernel
Normalized
Individual
Distance (Meters)from
Paved Road Category
0-600
>600
Migratory
44%
56%
Non-migratory
57%
43%
Both
47%
53%
47
Individual elk did show some significant differences between preferences in
distance classes to paved roads in comparison to their respective migratory group and the
entire study group of elk. Migratory elk #31 spent less time within 60 m of paved roads
than the entire group of elk in the study (t13 = 0.063, p < .10). The same elk (#31) spent
more time >600 m from paved roads than the elk in the study group (t13 = 0.063, p > .10).
Similarly, NM Elk #419 spent less time within 600 m of paved roads than the other elk in
her migratory group (t13 = 0.086, p < .10) The same elk (#419) spent less time in the
>600 m from paved roads distance class than the elk in her migratory group (t13 = 0.086,
p > .10).
Distance to Paved Roads 95% Range Areas
Overall, migratory elk were found to be within 600 m of paved roads 44% of the
time whereas non-migratory elk were in the same distance class 57% of the time. The
opposite was true for time spent in the >600 m of paved roads where migratory elk spent
56% of their time in the distance class and non-migratory spent 43% of their time in the
>600 m distance class (Table 7). Seasonal break downs by percentage of data points in
each distance class may be viewed in Figure 14 and Appendix 12.
Figure 14: Migratory and Non-migratory elk habitat use and distance to roads in the
Southwestern Cascades of Washington.
Normalized Frequency of Observations of
Elk in Distance (Meters) from Paved Road
Categories (95% Contour Kernel)
Percentage of Total
Observations
70%
Migratory
60%
Nonmigratory
50%
40%
30%
20%
10%
0%
0-600
>600
Distance from Paved Road Category
48
Individual elk did show significant differences between distance classes to paved
roads in comparison to their respective migratory group and the entire study group of elk.
Migratory elk #31 spent less time within 600 m of paved roads than the entire group of
elk in the study group (t13 = 0.075, p < .10). The same elk (#31) spent more time >600 m
from paved roads than the elk in the study group (t13 = 0.075, p > .10). Similarly, NM
Elk #420 spent less time within 600 m of paved roads than the other elk in the entire
study group (t13 = 0.055, p < .10) The same elk (#420) spent more time >600 m from
paved roads than the elk in her migratory group (t13 = 0.055, p > .10).
Distance to Non-Paved Roads 50% Range Areas
Overall, migratory elk were found within 100 m of non-paved roads 7% of the
time where non-migratory elk were in the same distance class 3% of the time. Remaining
distance class to non-paved roads comparisons may be viewed in Table 8. Seasonal
break downs by percentage of data points in each distance class may be viewed in
Appendix 13.
Table 8. Overall annual habitat use by elk relative to distance to non-paved roads in the
combined Randle and Packwood study areas at the 50% and 95% range scale in the SW
Cascades of Washington, 2007-2008.
50% Contour Kernel
Normalized
Distance (Meters)from Non-Paved
Road Category
Individual
0-100
100-200
200-400
400-600
>600
Migratory
7%
7%
15%
19%
53%
Nonmigratory
3%
9%
25%
15%
47%
Both
6%
7%
17%
18%
51%
95% Contour Kernel
Normalized
Distance (Meters)from Non-Paved
Road Category
Individual
0-100
100-200
200-400
400-600
>600
Migratory
8%
8%
14%
15%
55%
Nonmigratory
6%
9%
23%
15%
48%
Both
8%
8%
16%
15%
53%
Individual elk did show significant differences between distance classes to nonpaved roads in comparison to their respective migratory group and the entire study group
49
of elk. Migratory elk #28 spent more time within 100 m of non-paved roads than elk
within her respective migratory group (t13 = 0.000, p > .05) and the entire study group of
elk (t13 = 0.000, p > .05). The same elk (#28) was also found to be within the 100-200 m
distance class of non-paved roads more than the elk in her migratory group (t13 = 0.004, p
> .05) and the entire study group of elk (t13 = 0.017, p > .05). Elk #28 also was found in
the >600 m distance class less than the elk in her migratory group (t13 = 0.094, p > .10).
Migratory elk #30 was found in the 400-600 m distance class more than the elk in her
migratory group (t13 = 0.047, p > .05) and the entire study group of elk (t13 = 0.048, p >
.05). Migratory elk #83 was found in the 200-400 m distance class less than the elk in
her migratory group (t 13= 0.099, p < .10). Non-migratory elk #84 was also found in the
200-400 m distance class less than the elk in her migratory group (t13 = 0.099, p < .10).
The same elk (#84) was also in the >600 m distance class less than the other elk in her
migratory group (t13 = 0.095, p < .10. Finally, NM elk #420 showed differences where
she was within 100 m of non-paved roads more than the elk in her migratory group (t13 =
0.001, p > .05). The same elk (#420) spent more time in the 100-200 m distance class
than the elk in her migratory group (t13 = 0.027, p > .05) and the elk in the entire study
group (t13 = 0.029, p > .05). Elk #420 also spent more time in the 200-400 m distance
class than the elk in her migratory group (t13 = 0.005, p > .05) and the elk in the entire
study group (t13 = 0.085, p > .10).
Distance to Non-Paved Roads 95% Range Areas
Overall, migratory elk were found within 100 m of non-paved roads 8% of the
time where non-migratory elk were in the same distance class 6% of the time. Remaining
distance class to non-paved roads comparisons may be viewed in Table 8. Seasonal
break downs by percentage of data points in each distance class may be viewed in
Appendix 13.
Individual elk did show significant differences between distance classes to nonpaved roads in comparison to their respective migratory group and the entire study group
of elk. Migratory elk #28 spent more time within 100 m of non-paved roads than elk in
her respective migratory group (t13 = 0.010, p > .05) and the entire study group of elk (t13
= 0.011, p > .05). The same elk (#28) was also found to be within the 100-200 m
distance class of non-paved roads more than the elk in her migratory group (t13 = 0.046, p
> .05). Migratory elk #30 was found in the 400-600 m distance class more than the elk in
50
her migratory group (t13 = 0.014, p > .05) and the entire study group of elk (t13 = 0.047, p
> .05). Non-migratory elk #81 was found in the 400-600 m distance class more than the
elk in the entire study group (t13 = 0.082, p > .10). Migratory elk #83 was found in the
>600 m distance class more than the elk in her migratory group (t13 = 0.058, p > .10).
Non-migratory elk #84 was found in the 200-400 m distance class less than the elk in her
migratory group (t13 = 0.027, p < .05). The same elk (#84) was in the >600 m distance
class more than the other elk in her migratory group (t13 = 0.088, p > .10). Nonmigratory elk #85 was in the >600 m distance class more than all of the elk in the study
group (t13 = 0.062, p > .10). Finally, NM elk #420 showed differences where she was
within 100 m of non-paved roads more than the elk in her migratory group (t13 = 0.000, p
> .05). The same elk (#420) spent more time in the 100-200 m distance class than the elk
in her migratory group (t13 = 0.012, p > .05) and the elk in the entire study group (t13 =
0.006, p > .05). Elk #420 also spent more time in the 200-400 m distance class than the
elk in her migratory group (t13 = 0.060, p > .10) and the elk in the entire study group (t13 =
0.000, p > .05). In addition, elk #420 was found in the >600 m of non-paved roads less
than the elk in the entire study group (t13 = 0.021, p > .05).
Distance to Trails 50% Range Areas
Overall, migratory elk spent 1% of their time within 100 m of trails, where nonmigratory elk spent 0% of their time in the same distance class. Remaining distance class
to trails comparisons may be viewed in Table 9. Seasonal break downs by percentage of
data points in each distance class may be viewed in Appendix 14.
51
Table 9. Overall annual habitat use by elk relative to distance to trails in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the SW Cascades of
Washington, 2007-2008.
50% Contour Kernel
Normalized
Distance (Meters) from Trail
Category
Individual
0100
100-200
200-400
400-600
>600
Migratory
1%
1%
2%
2%
94%
Non-migratory
0%
0%
0%
0%
100%
Both
1%
1%
2%
2%
95%
95% Contour Kernel
Normalized
Distance (Meters) from Trail
Category
Individual
0100
100-200
200-400
400-600
>600
Migratory
2%
2%
4%
3%
89%
Non-migratory
0%
0%
1%
1%
98%
Both
2%
1%
3%
3%
91%
Individual elk did show significant differences between distance classes to trails
in comparison to their respective migratory group and the entire study group of elk.
Migratory elk #421 was found in the 100-200 m distance class more than the elk in the
entire study group (t13 = 0.053, p > .10). The same elk (#421) was found in the 400-600
m distance class more than the elk in her migratory group (t13 = 0.031, p > .05) and the
entire study group of elk (t13 = 0.004, p > .05). Elk #421 was less likely to be found in
the >600 m distance class than the elk in the entire study group (t13 = 0.078, p < .10).
Migratory elk #422 was found within 100 m of trails more than both the elk in her
migratory group (t13 = 0.000, p > .05) and the entire study group of elk (t13 = 0.000, p >
.05). The same elk (#422) was found in the 100-200 m distance class (t13 = 0.004, p >
.05), 200-400 m distance class (t13 = 0.000, p > .05), 400-600 m distance class (t13 =
0.047, p > .05) and the >600 m distance class (t13 = 0.002, p > .05) more than the elk in
her migratory group. In addition, elk #422 was also found in the following distance
categories more than the elk in the entire study group of elk: 100-200 m distance class
(t13 = 0.000, p > .05), 200-400 m distance class (t13 = 0.000, p > .05), 400-600 m distance
class (t13 = 0.008, p > .05) and the >600 m distance class (t13 = 0.000, p > .05).
52
Distance to Trails 95% Range Areas
Significant differences were detected between the migratory and non-migratory
groups of elk in relation to habitat used near trails. Differences were detected between
the two groups where migratory elk were found within 100 m more often than their nonmigratory counterparts (t13 = 0.033, p > .05). Migratory elk were also found more often
within the following distance categories than non-migratory elk: 100-200 m distance
class (t13 = 0.055, p > .10), 200-400 m (t13 = 0.049, p > .05), 400-600 m (t13 = 0.041, p >
.05). However, migratory elk were found in the >600 m distance class less often than
non-migratory elk (t13 = 0.041, p < .05). Overall, migratory elk spent 2% of their time
within 100 m of trails, where non-migratory elk spent 0% of their time in the same
distance class. Remaining distance class to trails comparisons may be viewed in Table 9.
Seasonal break downs by percentage of data points in each distance class may be viewed
in Appendix 14.
Individual elk showed significant differences between distance classes to trails in
comparison to their respective migratory group and the entire study group of elk.
Migratory elk #28 was found in both the 200-400 m distance class (t13 = 0.080, p > .10)
and 400-600 m distance classes more than all of the elk in the entire study group (t13 =
0.091, p > .10). Non-migratory elk #420 was found in the 200-400 m distance class more
than the elk in her migratory group (t13 = 0.000, p > .05). The same elk (#420) was found
in the >600 m distance class less than the elk in her migratory group (t13 = 0.000, p <
.05). Migratory elk #421 was found in the 0-100 m distance class more than the elk in
the elk in her migratory group (t13 = 0.069, p > .10) and the entire study group (t13 =
0.016, p > .05). The same elk (#421) was found in the 100-200 m distance class more
than the entire study group of elk (t13 = 0.037, p > .05). She (#421) was found in the 400600 m distance class more than all of the elk in the entire study group (t13 = 0.057, p >
.05). Elk #421 was less likely to be found in the >600 m distance class than the elk in the
entire study group (t13 = 0.052, p < .10). Migratory elk #422 was found within 100 m of
trails more than the entire study group of elk (t13 = 0.036, p > .05). The same elk was
found in the 100-200 m distance class (t13 = 0.036, p > .05) and 200-400 m distance class
(t13 = 0.033, p > .05) more than the entire group of elk in the study. She (#422) was also
found in the 400-600 m distance class more than the elk in the entire study group (t13 =
0.073, p > .10). Elk #422 was less likely to be found in the >600 m distance category
than the rest of the elk in the entire study group (t13 = 0.040, p < .05).
53
Habitat Use with Season and Season and Time as Predictors
Regression analysis was performed to determine if season and/or season and time
of day predicted habitat use by migratory elk versus non-migratory elk. The same
analysis was conducted to view differences in habitat use of individual elk.
Group Analysis: Season and/or season and time of day did influence habitat
selection per migratory group. At both the 50% and 95% scale levels season and season
and time of day predicted habitat use per slope category (% gradient) for migratory elk.
At the 50% scale, ordinal logistic regression analysis yielded a significant model that
predicted seasonal habitat use of migratory elk to utilize slopes with steeper gradients
with a Nagelkerke pseudo R2 of 0.223, and at the 95% scale with Nagelkerke pseudo R2
value of 0.248. Similarly, at the 50% scale, ordinal logistic regression analysis yielded a
significant model that predicted seasonal habitat use in combination with time of day to
predict migratory elk utilization of slopes with steeper gradients with a Nagelkerke
pseudo R2 of 0.232, and at the 95% scale with Nagelkerke pseudo R2 value of 0.253.
At the 50% scale, season and season/time of day predicted distance to trails for
migratory elk. Ordinal logistic regression analysis yielded a significant model that
predicted seasonal habitat use of migratory elk relative to distance to trails with a
Nagelkerke pseudo R2 of 0.231. Similarly, at the 50% scale, ordinal logistic regression
analysis yielded a significant model that predicted season and time of day habitat use
relative to distance to trails of migratory elk with a Nagelkerke pseudo R2 of 0.231.
Finally, at the 95% scale, season and season/time of day predicted distance to paved
roads for migratory elk. At the 95% scale, ordinal logistic regression analysis yielded a
significant model that predicted seasonal habitat use of migratory elk relative to distance
to paved roads with a Nagelkerke pseudo R2 of 0.239. Also, at the 95% scale, ordinal
logistic regression analysis again yielded a significant model that predicted season and
time of day habitat use of migratory elk relative to distance to paved roads with a
Nagelkerke pseudo R2 of 0.242.
Individual Elk Analysis: One elk (M #82) demonstrated predictability with
regard to season and season/time and aspect of habitat used (Appendix 15). Nine elk
showed predictability with regard to season and season/time and slope of habitat used
(Appendix 15). Eight of the nine elk were migratory. Very few elk (2 migratory elk)
showed predictability with regard to season and season/time and distance to cover of
54
habitat used (Appendix 15). Seven migratory elk showed predictability with regard to
season and season/time and distance to non-paved roads of habitat used (Appendix 15).
Eight migratory elk showed predictability with regard to season and season/time and
distance to paved roads of habitat used (Appendix 15). Six elk showed predictability
with regard to season and season/time and distance to shrub habitats used (Appendix 15).
Five of the six elk were migratory elk. Six elk showed predictability with regard to
season and season/time and distance to trail habitats used (Appendix 15). Five of the six
elk were migratory elk. Three migratory elk showed predictability with regard to season
and season/time and distance to water habitats used (Appendix 15).
DISCUSSION
Rural mountain areas in the Pacific Northwest are rapidly being discovered and
developed as vacation and/or second homes. In addition, these fertile river valley areas
such as the upper Cowlitz River Valley, continue to be home to various agricultural
practices and livestock ranching. As was the case with the herd in this study and many
other elk herds in North America, critical winter range areas are being reduced in size
because of human impacts. The results of our study confirm that non-migratory elk not
only prefer relatively flat areas, but are prone to be found in open habitats (pasture/clear
cuts) at greater distances from cover (safety), and use habitat in closer proximity to paved
roads than their migratory herd counterparts. As a result, the non-migratory elk in our
study appear to have become habituated to the presence of humans and human activity.
While individual elk often had strong behavioral differences compared to the rest
of their group, migratory and non-migratory elk primarily differed in use of flat vs. steep
topography, slope preference, abundance near trails, and season/time of day.
The differences detected between individual elk and their respective migratory
group and the overall study group of elk were beyond the scope of the study. However, it
is important to recognize the differences, as further analysis could show linkages to
genetic and or/behavioral factors that may impact the behavior of individual family/social
groups of elk within an elk herd.
Home Range
Home ranges on the 53 elk were estimated using the minimum convex polygon
(MCP) method for home range analysis and comparisons. Although, other methods for
55
estimating home range are used to yield greater precision, the MCP method is the oldest
method that has been commonly used (White and Garrott 1990). For our purposes of
comparing home range size between migratory and non-migratory elk, the simplicity,
flexibility of shape, and ease of calculation of MCP was ideal. The study areas for both
the Randle and Packwood group also used the MCP for the entire sub-herds to determine
study area boundaries.
The purpose of the home range area comparisons between migratory and nonmigratory elk was to identify differences, if any, in how the two segments of the herd
were using the landscape. Elk home range areas central and west of the Rocky
Mountains have been reported to span anywhere from 1 km2 – 90km2 in size (Schroer
1987). Overall, the collared elk (both migratory and non-migratory) in our study had a
combined mean life home range of 61 km2 (6100 ha), migratory elk life home range was
102 km2 with a range of 54-102 km2 annually over the 2004-2008 year period. The nonmigratory life home range was 20 km2 with a range of 5.4-20 km2 over the 2004-2008
year period. Relative to other studies the home range estimates of this study were similar
to range areas of other studies (Jenkins 1980, Pope 1994, Millspaugh 1995, Cole 1996).
It is difficult to compare home range areas among different studies since the results will
vary based on the method of estimation used as well as the influence of sample sizes on
the results. Much analysis and comparisons of home range estimation methods have been
conducted (Pope 1994, Witmer 1981, White and Garrott). Still, the non-migratory range
areas of this study are very similar to the results of other studies that focused on nonmigratory elk. Pope (1994) focused his study on non-migratory elk which had a mean
home range of 7.65 km2 over an approximately 14 month period. Other studies on nonmigratory elk using varying methods and sample sizes ranged between 4 km2 (Witmer
1981), 10 km2 (Jenkins 1980), and 3 km2 (Franklin et al. 1975). Many of these studies
analyzed data over ~ 12 month periods. Our study included five separate biological years
of data for analysis and comparison and also included life home range estimates of the
combined five years of data separated out per migratory status. Our non-migratory range
areas per biological year yielded similar results to the studies cited above. Mean nonmigratory range sizes included the following: 2004 = 8.5 km2, 2005=5.4 km2,
2006=11km2, 2007=11km2, 2008=14km2, life home range=20km2. Mean migratory
range sizes included the following: 2004=58km2, 2005=54km2, 2006=54km2,
2007=55km2, 2008=75km2, life home range=102km2. Considering the fact that
56
migratory elk travel greater linear distances to reach high-elevation summer range areas,
intuitively we expected that home range areas of migratory elk would be significantly
larger and different than non-migratory elk. The results of our home range analysis
provide evidence that home range areas between migratory and non-migratory elk were
significantly different for the study group of elk.
Migration and Distribution
Anecdotal inferences have been made to explain elk behavior of the South
Rainier elk herd (Bradley 1982, WDFW 2002). It was previously thought that a clear
delineation could not be made between summer and winter ranges in the South Rainier
elk herd area (WDFW 2002). Elk that resided in areas outside of Mt. Rainier National
Park were thought to have become non-migratory because of logging practices that
opened up expanded habitats favorable to elk (WDFW 2002). Other studies have
demonstrated that it has not been uncommon that elk herds may typically have a portion
of the population that is migratory and a portion that is non-migratory (Martinka 1969,
Craighead et al. 1972, Boyd 1970). This study showed much more variation in elk
distribution than previous understandings described. The high fidelity rate of the elk in
our study is consistent with other studies where elk showed high range fidelity from year
to year (Smith and Robbins 1994, Schwartz and Mitchell 1945). The elk wintering in the
upper Cowlitz Basin between Randle and Packwood were previously thought to have
either been non-migratory because of forest practices that had changed outside of MRNP
or if migratory, migrated to Mount Rainier National Park (WDFW 2002). The results of
this study showed that migratory elk were ceasing most opportunities to access quality
forage in the sub-alpine areas surrounding the Packwood and Randle study areas.
Survival Rates
Survival rates were similar, but mostly lower in comparison to survival rates of
other studies in Oregon (Pope 1994, Stussey 1993). Factors contributing to mortality that
may have resulted in lower survival rates included vehicle collision 16%, wounding/
legal harvest/illegal hunting 32%, predator kills 16%, starvation 20%, and high
proportions of unknown causes 24%. Legal and illegal hunting and wounds associated
with hunting accounted for the highest annual percentage of mortalities. Legal hunting of
both male and female elk is allowed during their respective seasons. In response to
chronic damage complaints in the agricultural segment of the range, state officials have
57
created damage hunt areas (IE: Game Management Units 503 and 516) where female elk
harvest is allowed. Additionally, ≤ 40 female elk are legally harvested by tribal hunting
annually. Illegal hunting (poaching) has become a chronic problem for elk in the study
area. Poaching has been listed in the South Rainier Elk Herd Plan as the primary limiting
factor of the herd (WDFW 2002). Severely reduced State and local governmental
budgets have resulted in little, to no, enforcement coverage to address localized poaching
problems. With regard to road-kill, the winter range area of the herd has the highest
incidence of vehicle-elk collisions in the entire state of Washington (Meyers et al. 2008).
The percentage of elk killed by cougars is not unusually high, however, if state
sanctioned cougar hunts allowed the use of hounds, the local cougar population
impacting the herd would be dramatically reduced. The use of hounds to hunt cougars is
prohibited by the State of Washington in the range area of the study. In the case of
moralities attributed to starvation, it is not uncommon to have winter-kill numbers in the
15% range. Our mean % of starvation mortalities for the study period was 20%, slightly
higher than the acceptable range (Skovlin et al. 2002). The proportion dying from
starvation should be viewed carefully and noted that, during two of the four years of the
study we experienced La Nina weather conditions. With the La Nina weather system, we
received excessive amounts of snow in low elevation areas for extended periods of time.
Not only did the extreme winter weather make the elk more vulnerable to legal and
illegal hunting, food sources were dramatically reduced. Because of the small sample
size for overall annual mortalities (3-10 per year), differences between migratory and
non-migratory elk were not analyzed.
Habitat Selection
Aspect
Many studies have shown evidence that elk prefer specific aspects that vary
seasonally. Skovlin and others (2002), found that in winter, elk prefered upper southfacing slopes that, because of wind, sun angle (radiation) or shade pattern were the first to
become bare of snow. Overall, both migratory and non-migratory elk spent a higher
percentage of their time in southern aspects during all four seasons of the year that was
analyzed. Witmer (1981) also found similar results in his study of elk in western Oregon.
The literature also suggests that elk prefer northern aspects during summer months
(Skovlin et al. 2002). Both the migratory and non-migratory groups of elk in this study
58
did not show a preference for north-facing slopes during summer. The migratory elk in
this study, showed a preference for south facing slopes regardless of season. In summer
the migratory elk mostly spent their time above 5000 ft where temperatures were cooler
and still within the elevation range of sub-alpine habitat areas where cover/shade were
also abundant. This likely provided relief from warmer temperatures. It could also be that
the sub-alpine areas used by the migratory elk in this study remain lush with abundant
vegetation throughout the summer regardless of aspect. Although some of the historic
studies that have documented patterns of habitat use by elk have shown elk prefer
northerly aspects during summer, these studies may have been in areas with arid climates
where elk must seek north aspects in search of abundant vegetation (Nelson and Burnell
1975, Julander and Jeffrey 1964). More recent studies have shown even more variation
to habitat use and preferences that deviate from what had been considered normal
patterns such as southern aspect preference in winter and north-facing preferences in
summer (Bracken and Musser 1993). The non-migratory elk in our study spent
approximately twice as much of their time in flat aspect habitat areas than the migratory
elk. This is not surprising since the river bottom areas that encompass both summer
range for non-migratory elk and winter range for the entire herd consists of dense riparian
vegetation that provide abundant water, food, and cover. In addition, the flat range areas
used by the non-migratory elk in summer include agricultural lands and residential areas
with an appealing array of vegetation. Although the regression analysis to detect aspect
preferences by season did not show significant correlations, it was possible to view a
large data set (>80,000 data points) and observe where the elk were spending their time
feeding and bedding. Migratory elk spent 11% of their time in habitat that is considered
flat, where NM elk spent 19% of their time in the flat habitat type, NE aspect: M=16%,
NM=11%, SE aspect: M=32%, NM=24%, SW aspect: M=16%, NM=28%, NW aspect:
M=25%, NM=18% (Table 2). Migratory elk were found to use the SW aspect
significantly less than the NM elk group (t13=0.035, p<.10). Based on the results of other
studies I expected to see clear patterns of aspect preferences for both summer and winter
seasons. The results demonstrated variation in aspect preferences as other studies have
done (Pope 1994). Based on the percentage of data point locations, the results also
showed differences between aspect preferences of migratory and non-migratory elk.
59
Slope
Elk have been known to use steeper slopes in summer than in winter (Julander
and Jeffrey 1964, Zahn 1974, Marcum 1975, Leege et al. 1975). Although not identified
as such, many of the studies that document elk habitat use with slope preferences ranging
between 15% - 50% appear to be describing migratory herds, and/or migratory segments
of particular elk herds. The results showed that migratory elk preferred steeper 30-45%
and >45% slopes more than non-migratory elk at both the 50% and 95% scales. In
addition, the results showed that season predicted habitat use of migratory elk use of
habitat slopes with steeper gradients. These results are similar to Marcum (1975) who
found that elk on summer range preferred moderately steep slopes – 27% to 58%.
Studies in north-central Idaho, also found that elk on summer range tend to use 20-40%
slopes up to 60% as seasons progressed (Leege et al. 1975, Hershey and Leege 1982).
Typically, slopes > 20% correlate with sub-alpine and alpine habitat types in our study
area. Sub-alpine and alpine habitat in summer provides the best quality and quantity of
forage, in addition to abundant cover for migratory elk. The disparity in slope
preferences between migratory and non-migratory elk in our study demonstrates optimal
habitat use based on behavior. Non-migratory elk in this study preferred 1-15% slopes
more than migratory elk. The areas encompassing summer range that are 1-15% in slope
are river bottom areas rich in riparian forage, with abundant cover and water. The areas <
15% also include agricultural areas in the river bottom, which are also desirable to elk.
The adjacent slopes >15% slopes are dense evergreen forest areas of Gifford Pinchot
National Forest. The dense canopy cover of trees in this area does not allow sunlight to
penetrate the forest floor to stimulate under-story growth of plant communities that elk
prefer for browse. In other words, the forage in the > 15% slope areas of the nonmigratory elk range would be of lower quality and quantity. Tree overstory and canopy
coverage are primary determinants of understory herbage productivity (Skovlin et al.
2002). High degrees of canopy closure often equate to little or no elk food under the
canopy (Skovlin et al. 2002).
Cover
Cover is considered to be an important component to elk survival for both the
thermal benefits contributing to temperature regulation and hiding or escape cover.
Studies in Washington State do not support the hypothesis that elk require forest cover in
60
summer to maintain body temperatures, at least in Pacific coastal climates (Skovlin et al.
2002, Merrill 1991).
I analyzed my data with a view to determine if cover and distances to cover were
important per migratory group primarily for security/hiding and escape cover. Migratory
elk in this study spent 51-56% of their time within the 0-50 m distance class of cover
where the non-migratory elk spent 35-44% of their time within the same distance class.
Seasonally, during winter migratory elk spent 57%-61% of their time within 50 m of
cover as compared to non-migratory elk which spent 34%-39% of their time within 50 m
of cover. In addition, the results of t tests showed that migratory elk preferred to be
within 50 m of cover in winter more than the non-migratory elk. An explanation for this
behavior could be that the migratory elk were less habituated to humans and human
presence than the non-migratory elk. Therefore, while on winter range, migratory elk
tended to be within 50 m of escape cover more than non-migratory elk. In summer, there
was no disparity between the percentages of data points per migratory group per distance
classes. This could have had to do with the fact that while migratory elk were on summer
range, being in close proximity to escape cover was not as important for a couple reasons.
First, much of summer range was within National Park, Monument, and/or Forest Service
lands where human disturbance was reduced or limited. Second, there were no hunting
seasons during summer, unlike winter where hunting seasons did occur annually.
Shrub
The greatest percentage of the study data points for both migratory and nonmigratory elk fell within the 50 m distance class of the shrub/scrub habitat types. This
type of habitat may be referred to as an ecotone where different types of vegetation are
juxtaposed, which include high frequencies of early successional communities which are
important components of elk habitat (Skovlin et al. 2002). Levels of elk activity have
been known to decrease with increased distance from the interface of forest and nonforest communities (Marcum 1975, Winn 1976, Leckenby 1984). Studies in the Wasatch
Mountains of Utah demonstrated that both frequency of plant species and herbage
biomass at an edge was two times greater than 46 m into a meadow (Winn 1976). This
data showed migratory elk preferred ecotone/edge habitats (0-50 m) >83% of the time in
summer and >52% of the time throughout the rest of the year. One non-migratory elk fell
within this distance class >59% of the time in summer and >32% of the time the rest of
61
the year. Greater than 96% of our data points for both migratory and non-migratory elk
fell within 600 m of the shrub scrub habitat type for all seasons. Also of interest was that
89% of the data points for both migratory groups fell within the 400 m distance class
during summer. Leckenby (1984) found that at least 80% of elk use in summer forage
areas occurred within 274 m edge ecotone areas in the Blue Mountains of Oregon. In the
50% range areas, migratory elk spent 65% of their time in the 0-50 m range of
shrub/scrub while NM elk spent 58% of their time in the same distance class.
Water
Studies from various regions of the northwest suggest elk prefer summer habitats
within 800 m of water (Bracken and Musser 1993, Jeffrey 1963, Mackie 1970, Marcum
1975). Greater than 78% of the elk locations were within 600 m of water in summer for
both migratory and non-migratory elk. Even though my data is consistent with other
findings, many of these studies appear as though they were in climates that tended to be
semi-arid. What appeared to be somewhat unusual was that I had the greatest frequency
of elk found in distances greater than 600 m during summer than other seasons
throughout the year. The greater number of elk points in this distance class during
summer could be the result of temporal differences rather than an overall statement of
water preferences. The summer of 2008 was unusually cool which could have slowed
snow melt in upper alpine and sub-alpine areas utilized by the migratory elk. In that case,
snow melt in these areas often times provides temporary water sources in the form of
shallow alpine depressions where the snow has melted. Since these water sources are not
included in the water inventory they would not show up in the analysis. It is clear that
water sources and associated riparian habitats are important to elk in this study based on
the data points. Riparian habitat not only provide natural travel corridors, they have
greater plant diversity, and possess different microclimates from surrounding areas due to
increased humidity, consequently providing relief from temperature extremes (Oakley et
al. 1985, Thomas et al. 1979) The data showed that both migratory segments of the herd I
studied had very similar preferences for water and riparian habitats. Habitat use in
proximity to water may fluctuate temporally, however, this physiological need of elk to
utilize these habitats appeared to transcend migratory behavior.
62
Roads and Trails
Of all the factors related to logging, the construction of roads and the subsequent
vehicle traffic on those roads has proved to be the most significant modification of elk
habitat (Lyon and Christenson 2002). Not only do roads have a tendency to collect debris
and impede elk movement they also facilitate human access/contact with elk. Elk have
been known to avoid human disturbance and/or roads (Irwin and Peek 1979, Lyon 1979,
Hershey and Leege 1982). The development of many roads have been sighted in
topographic locations that were previously used as wildlife travel routes (Skovlin et al.
2002). As a result, many migration routes have been fragmented and/or have accelerated
rates of auto-collisions with wildlife crossing paved roads have occurred. In fact, SR 12
that bisects the winter range has the highest rate of auto-collisions with elk in Washington
State (Meyers et al. 2008). My analysis viewed elk habitat use and its associated
distances to roads in three categories which were: distance to paved roads, non-paved
roads, and trails. This discussion has therefore been broken down by road type category
as follows.
Paved Roads
Overall, non-migratory elk spent a greater percentage of their time within 600 m
of paved roads than migratory elk. One caveat to note is that State Route 12 runs parallel
to the Cowlitz River which runs through the heart of the winter range area and is mostly
within 600 m of the river. During the winter months the elk are typically found in the
river bottom areas between the Packwood sub-area and the Randle area. In light of that
fact, it does not seem unusual that migratory and non-migratory elk would be using
winter habitat areas similarly. The other seasons showed greater variation between the
migratory groups, primarily during summer. Many of the winter range areas that are
within 600 m of paved roads also fall within no-shoot zones. It could also be that the
herd as a whole did not experience security issues when using habitat areas within 600 m
of paved roads in winter because many of these areas are designated no-shoot zones.
Migratory elk spent 13-18% of their time within 600 m of paved roads during summer
whereas non-migratory elk spent 45-55% of their time within the same distance class.
Explanation for the differences during summer could be that there are very few paved
roads, if any, within 600 m of the sub-alpine and alpine habitat areas utilized by
migratory elk. Since the non-migratory elk remain in areas more populated by humans
63
with associated development and infrastructure (roads etc.), it would be an intuitive
assumption to predict non-migratory elk may be more habituated to the presence of
humans and roads, and as a result be found more often in closer proximity to paved roads
than migratory elk. However, migratory elk were also found to avoid the 0- 600 m
distance class of paved roads both during spring and fall seasons more than their nonmigratory counterparts. The data show migratory elk avoiding habitat areas within 600 m
of paved roads more than non-migratory elk, with the exception of the winter season.
The results of the data must be viewed carefully to avoid potential misleading cursory
summations of what the data may be demonstrating.
Non-Paved
Overall, both migratory and non-migratory elk avoided non-paved roads to a
greater extent cumulatively throughout the year. What was also clearly demonstrated in
the data, was that both migratory and non-migratory elk avoided using habitat within 200
m of non-paved roads >80% of the time. This supports the literature that states elk avoid
areas nearest roads (Lyon and Christensen 2002). Non-paved roads are the most travelled
roads/paths for vehicles to access hunting grounds on public and non-public lands in our
study area. It is therefore not surprising to see that elk avoid habitat areas within 200 m
of the non-paved roads and that they are in fact found at distances >600 m most of the
time. Also, most of the non-paved roads within our study area were outside of most noshoot zones that prohibit hunting which stands to reason that both segments of the herd
would be wary to place themselves within shooting distance of the non-paved roads.
Seasonally, habitat use patterns were fairly similar between the migratory and nonmigratory elk groups.
Trails
There were statistically significant differences between the migratory and nonmigratory elk at the 95% home range scale. However, it must be noted that trails within
the study area were only relative during summer months, and mostly only present in the
migratory elk habitat areas. Therefore, the analysis has been included but should be used
carefully when considering behavioral differences between migratory and non-migratory
elk in this area. No differences were detected between the migratory and non-migratory
groups of elk in relation to habitat used near trails in the 50% range areas. However, at
64
the 95% scale, differences were detected between the migratory and non-migratory
groups of elk in relation to habitat used near trails.
65
MANAGEMENT IMPLICATIONS
The South Rainer elk herd plan (2002) calls for more information on the
movements and wintering areas of the herd. This study identified three separate
wintering areas of elk considered part of the South Rainier elk herd. The three wintering
areas were along the upper-Cowlitz River in the Packwood area, upper-Cowlitz River in
the Randle area, and the lower Cispus drainage just south of Randle near the town of
Cispus. The Cispus wintering area was combined with the Randle range area because of
its close proximity to the Randle group. To start with, the land ownership in the Cispus
drainage area consisted of a combination of small ranches and homesteads, industrial
timberlands, and portions of the Gifford Pinchot National Forest. Very few, if any elk
damage complaints have been reported in this area. However, elk damage complaints in
both the Packwood and especially in the Randle range areas along the Cowlitz have been
common. One of the biggest challenges for the sustainability of this herd is landowner
tolerance of elk on winter range. This study may be used to identify ideal areas for
conservation of winter range. The defining boundary that we used between the
Packwood and Randle range areas was the Cora Bridge located where SR 12 crossed the
Cowlitz River between Packwood and Randle. This study showed that there was very
little, if any mixing between the Packwood and the Randle groups. One elk that
primarily wintered in the Packwood Range area (95%) was known to move southwest of
the Cora Bridge into the Randle sub-area 5% of the time in winter. The non-migratory
segment of the herd in the Randle range area have been known to cause summer crop
damage and as a result have been targeted by state officials for removal (WDFW Issue
10/42, WA Fish and Wildlife Commission Minutes, 2009).
The results of this study provide detailed information on the herd with added
insight on how the herd uses the range. As human encroachment into elk range continues
to increase, planning for, protecting and securing critical habitat such as winter range is
imperative. Responsible management of wildlife needs to be closely tied to practicing
ethical stewardship, and a responsibility to live with wildlife rather than resorting to
lethal removal when the wildlife we encounter become inconvenient. Land acquisition
with a goal of creating winter refuge areas is needed in both the Packwood and Randle
range areas. Elk in these two locations have become a popular roadside attraction to
tourists travelling through the area. Development of elk refuges in these locations could
benefit elk by preserving critical habitat, and serve to promote local tourism by
66
incorporating elk viewing and interpretive information of the local elk and other local
fauna.
Home range areas between the migratory and non-migratory elk were
significantly different. As predicted, the home ranges of the migratory elk were much
larger than the non-migratory elk. Critical habitat for non-migratory, migratory, and the
elk herd as a whole have been identified as a part of this study. Some of the other
differences between the migratory and non-migratory elk with regard to habitat use
preferences were illuminating. Migratory elk were found in closer proximity to cover
more often than non-migratory elk. Migratory elk also selected habitat areas at greater
distances to paved and non-paved roads than non-migratory elk. These data reinforce the
view that non-migratory elk tend to be more habituated to humans and human
disturbance. Land managers should continue to seek opportunities to increase forage
enhancement projects on surrounding U.S. Forest Service lands, WA DNR lands, and
industrial timberlands. In addition, an effort should be made to work with landowners in
damage-prone areas of winter range to develop incentives for conservation easements to
enhance and maintain elk habitat, in addition to allowing elk use of their property.
These results contribute to the body of knowledge describing home range sizes of
elk in the region of our study, and habitat preferences with regard to distances to roads,
cover, and use of aspect and sloped habitats per season. The elk in this study avoided
habitat areas within 600 m of paved roads the majority of the time and avoided habitat
areas within 200 m of non-paved roads the majority of the time. Management activities
including road closure opportunities in critical winter habitat areas should be pursued to
minimize disturbance to elk during winter. Cover for security was important to nonmigratory elk and even more so for migratory elk. Proximity to cover during summer
was not as important to elk as it was during other seasons, especially during winter. If
future habitat improvements are made on winter range, consideration must be made in
providing escape cover within distances of 200 m of the habitat enhancements.
Differences in behavior and habitat use were detected between migratory elk and
non-migratory elk. Based on this analysis, non-migratory elk do appear to have become
more habituated to the presence of human activity. The results of this study demonstrate
similar migratory patterns in terms of proportions of elk that are migratory and nonmigratory within the same herd (Craighead et al. 1972, Martinka 1969). With an interest
67
to manage and maintain the herd in a holistic way, it is important to avoid short-sighted
management decisions in response to damage complaints that, in many instances could
result in elimination of non-migratory segments of populations. Targeting segments of a
population such as the non-migratory portion of a herd obscures the natural migration
pattern and understanding of the respective herd (Irwin 2002). The non-migratory elk
tended to use habitat areas that were relatively flat (62% of the time) indicating that flat
river bottom areas were important habitat year around. The non-migratory elk tended to
use habitat areas in the river-bottoms as well as adjacent upland areas on both north and
south facing slopes. The summer range areas used by the migratory elk are fairly
abundant in quantity and quality of forage. Although the non-migratory portion of the
herd is relatively small in numbers, it could negatively impact forage availability for the
entire herd in winter. On ranges occupied by elk during summer and winter, summer use
of important forage plants by elk can reduce forage supplies during winter (Martinka
1969). However, studies of domestic cattle light spring grazing may have improved
forage quality in winter range areas (Anderson and Scherzinger 1975). Protection of
winter range areas remains of particular concern to both WDFW and Tribal wildlife
managers. The concern is not so much non-migratory elk degrading conditions on what
will also be winter range by the collective herd, as much as conflicts with existing
agricultural land users, and increasing development within the winter range areas.
Planning for the sustainability of the herd must focus on issues that have been raised
around winter range. Securing of funds to address chronic damage to agricultural lands
incurred by non-migratory elk in summer would be useful. In addition, funding for elkproof fencing in chronic damage prone agricultural lands would also help. Long-term
solutions would include land acquisition on winter range with a goal of creating elk
refuges both in the Randle and Packwood areas. Although very similar in the way both
migratory and non-migratory elk use the landscape, the data show non-migratory elk
using primarily river-bottom areas in closer proximity to human activity. The nonmigratory elk are seizing opportunities to exploit the best habitat possible, which is
frequently in agricultural areas. Core home range areas were in flat habitat areas with
regard to aspect 62% of the time.
Wildlife over and under passes have been successful in reducing wildlife
mortalities as the result of auto-collisions in other portions of the state of Washington (I90 corridor near Snoqualmie Pass) and in areas of Canada near the town of Banff. Use of
68
these data to plan for wildlife crossings bisecting SR 12 would help reduce elk mortalities
and auto-collisions in the winter range area.
The three primary goals of the WDFW South Rainier elk herd plan state the
following: to manage the elk herd for a sustained yield; to manage elk for a variety of
recreational, educational, and aesthetic purposes including hunting, scientific study,
cultural and ceremonial uses by Native Americans, wildlife viewing and photography;
and to manage and enhance elk and their habitats to ensure healthy, productive
populations. Tribal wildlife managers also share these goals for the South Rainier elk
herd. To achieve these goals in planning for the sustainability of the elk herd, legal
harvest must also continue to be carefully regulated using the best available science to
steer management decisions.
Finally, although this study contributes to a greater understanding of the
movements and habitat preferences of the South Rainier elk herd, and ultimately to a
larger body of knowledge on elk ecology, further studies on the herd could be useful. For
example, supplemental research with a focus on bull habitat preferences would be
especially useful in understanding the entire elk herd. Also, radio-collaring additional
cow elk in the Randle area will be important to continue to gather more information on
the habitat use and distribution of the migratory elk that winter in this area.
69
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74
APPENDICES
75
Appendix 1. Winter range plant list for Western Hemlock zones (Franklin and
Dyrness, 1973).
Western Hemlock zone
Overstory species
Pseudotsuga menziesia
Tsuga heterophylla
Acer macrophyllum
76
Understory species
Taxus brevifolia
Holodiscus discolor
Acer circinatum
Berberis nervos
Gaultheria shallon
Vaccinium parviflora
Rubus ursinus
Herb layer
Linnaea borealis
Trillium ovatum
Adenocaulon bicolor
Chimaphila umbellate
Polystichum munitum
Oplopanax horridus
Appendix 2. Summer range plant lists for Pacific Silver and Mt.
Hemlock zones (Franklin and Dyrness, 1973).
Pacific Silver fir zone
Overstory species
Abies amabilis
Abies procera
Tsuga heterophylla
Pseudotsuga menziesii
Understory species
Vaccinium alaskaense
V. Ovalifolium
Berberis nervosa
Acer circinatum
Rubus lasiococcus
Gaultheria shallon
Herb layer
Cornus Canadensis
Clintonia uniflora
Pyrola secunda
Achlys triphylla
Xerophyllum tenax
Linnea borealis
Tiarella unifoliata
Mt. Hemlock zone
Overstory species
Tsuga mertensiana
Abies lasiocarpa
A. Amabilis
Chamaecyparis nootkatensis
Pinus monticola
P. albicaulis
Picea engelmannii
Understory species
Vaccinium membranaecum
V. alaskaense
V. Ovalifolium
Rhododendron albiflorum
Menziesia ferruginea
Rubus lasiococcus
Sub-alpine meadow
Understory species
Phyllodoce empetriformis
Cassiope mertensiana
Vaccinium deliciosum
Phlox diffusa
Herb layer
Valeriana sitchensis
Lupinus latifolius
Veratrum viride
Polygonum bistortoides
Aster ledophyllus
A. alpigenus
Ligusticum grayi
Castilleja parviflora
Anemone occindentialis
Potentilla flabellifolia
Arnica latifolia
Luetkea pectinata
Antennaria lanata
Grass and Sedge species
Festuca viridula
Carex spectabilis
C. nigricans
77
Appendix 3. Collar 82 Batch fixed kernel density at the 50, 90 &95% range areas
78
Appendix 3. Collar 83 Batch fixed kernel density at the 50, 90 &95% range areas
79
Appendix 3. Collar 84 Batch fixed kernel density at the 50, 90 &95% range areas
80
Appendix 3. Collar 85 Batch fixed kernel density at the 50, 90 &95% range areas
81
Appendix 3. Collar 86 Batch fixed kernel density at the 50, 90 &95% range areas
82
Appendix 3. Collar 419 Batch fixed kernel density at the 50 & 95% range areas
83
Appendix 4. Habitat Attributes and distance classes for habitat use analysis of female elk
in southwest Washington.
Habitat Attribute
Distance (m) Class
Water
0 – 50
51-100
101-200
201-400
401-600
>600
Trails
0 – 100
101 – 200
201-400
400-600
>600
Non-paved Road
0 – 100
101-200
201 – 400
400-600
>600
Paved Roads
0 – 600
>600
Cover
0 – 50
51-100
101-200
201-400
401-600
>600
Shrub/scrub
0-50
51-100
101-200
201-400
401-600
>600
Slope
84
0-1 = 1
1-15=2
15-30=3
30-45=4
45-25000=5
Habitat Attribute
Aspect
Distance (m) Class
-1-0=1
0-90=2 (NE)
90-180=3(SE)
180-270=4(SW)
270-360=5(NW)
Activity
0-10=1
10.1-100=2
85
Appendix 5. All Elk. Radio frequencies, capture dates and total number of locations of radiocollared elk in southwest Washington, 2004-2008.
Name
86
Capture Date
# of Locations
Alma
March 2005
61
Amelia
March 2006
39
Andrea
March 2007
25
Anise
March 2008
6689
Annie
March 2006
42
Arnica
March 2008
6627
Becky
March 2005
11
Carrie
March 2005
51
Cheyenne
March 2007
30
Christine
March 2006
10
Cindy
March 2005
64
Cinnamon
March 2008
6890
Clove
March 2008
6793
Cynthia
March 2005
49
Echinacia
March 2008
6720
Eleanor
March 2006
42
Erika
March 2005
49
Hilary
March 2006
11
Jane
March 2006
28
Kate
March 2005
62
Kathie
March 2005
42
Kendra
March 2007
6177
Liz
March 2005
61
Marilu
March 2005
47
Mary
March 2005
64
Mary Ellen
March 2007
5471
Nevada
March 2006
42
Nutmeg
March 2008
6482
Roslyns
March 2007
4873
Sage
March 2008
6876
Sally
March 2005
63
Sheila
March 2005
63
Susan
March 2005
46
Therese
March 2006
42
Appendix 6. Elk name, life home range area, and migratory status of radio-collared elk
in southwest Washington, 2004-2008.
Name
MCP Area Km2
Migratory Status
Akirst
45.57
Non-migratory
Bridget
25.62
Non-migratory
Elma
19.35
Non-migratory
Gertrude
7.66
Non-migratory
Heather
25.88
Non-migratory
Janie
5.69
Non-migratory
Jennifer
10.09
Non-migratory
Judee
12.48
Non-migratory
Judy
19.53
Non-migratory
Lavender
5.63
Non-migratory
Martina
17.85
Non-migratory
Nicky
21.39
Non-migratory
Pilar
11.44
Non-migratory
Rachel
9.42
Non-migratory
Rosa
79.52
Non-migratory
Ruth
19.78
Non-migratory
Thyme
16.72
Non-migratory
Alison
1.80
Non-migratory
Alma
122.02
Migratory
Amelia
103.45
Migratory
Andrea
125.86
Migratory
Anise
92.22
Migratory
Annie
275.10
Migratory
Arnica
31.45
Migratory
Becky
20.67
Migratory
Carrie
106.21
Migratory
Cheyenne
37.06
Migratory
Christine
57.61
Migratory
Cindy
221.87
Migratory
Cinnamon
45.52
Migratory
Clove
42.70
Migratory
Cynthia
137.69
Migratory
Echinacia
177.79
Migratory
87
MCP Area Km2
Migratory Status
Eleanor
109.34
Migratory
Erika
66.22
Migratory
Jane
61.05
Migratory
Kate
118.72
Migratory
Kathie
107.25
Migratory
Kaye
270.84
Migratory
Kendra
48.03
Migratory
Liz
113.34
Migratory
Marilu
99.32
Migratory
Mary
95.72
Migratory
MaryEllen
83.37
Migratory
Nevada
76.36
Migratory
Nutmeg
79.47
Migratory
Roslyn
96.62
Migratory
Sage
70.20
Migratory
Sally
76.40
Migratory
Sheila
73.44
Migratory
Susan
82.96
Migratory
Therese
96.60
Migratory
Bettina
198.73
Migratory
Name
88
Appendix 7. Habitat use of aspects by elk in the combined Randle and Packwood study
areas at the 50% and 95% range scales in the Southwest Cascades of Washington. 20072008.
Migratory
Elk
Aspect
Winter
50%
Winter
50%
Spring
50%
Spring
50%
Summer
50%
Summer
50%
Fall 50%
Fall
50%
Flat
10%
10%
10%
10%
11%
11%
13%
13%
NE
16%
42%
15%
42%
17%
40%
20%
36%
SE
32%
48%
32%
48%
34%
49%
36%
50%
SW
16%
16%
16%
15%
NW
26%
27%
23%
17%
Nonmigratory
Elk
Aspect
Winter
50%
Winter
50%
Spring
50%
Spring
50%
Summer
50%
Summer
50%
Fall 50%
Fall
50%
Flat
19%
19%
17%
17%
21%
21%
15%
15%
NE
11%
29%
10%
32%
12%
26%
9%
36%
SE
24%
52%
22%
51%
25%
53%
21%
50%
SW
28%
29%
28%
29%
NW
18%
22%
14%
27%
89
Migratory
Elk
Aspect
Winter
95%
Winter
95%
Spring
95%
Spring
95%
Summer
95%
Summer
95%
Fall 95%
Fall
95%
Flat
7%
7%
7%
7%
8%
8%
8%
8%
NE
17%
42%
17%
41%
19%
29%
19%
39%
SE
30%
50%
30%
50%
32%
52%
32%
52%
SW
20%
20%
20%
20%
NW
25%
24%
20%
20%
Nonmigratory
Elk
Aspect
Winter
95%
Winter
95%
Spring
95%
Spring
95%
Summer
95%
Summer
95%
Fall 95%
Fall
95%
Flat
17%
17%
12%
12%
14%
14%
14%
14%
NE
14%
33%
10%
42%
12%
38%
12%
38%
SE
25%
50%
21%
46%
23%
49%
23%
49%
SW
25%
25%
26%
26%
NW
19%
32%
26%
26%
90
Appendix 8. Habitat use by elk relative to distance to cover in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern Cascades of
Washington, 2007-2008.
Migratory
Elk
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category
Category
Category
Category
Category
0-50
57%
47%
36%
43%
61%
52%
49%
61%
50-100
16%
17%
17%
18%
15%
17%
17%
15%
100-200
18%
23%
28%
25%
16%
20%
22%
16%
200-400
7%
10%
14%
11%
7%
9%
10%
7%
400-600
1%
2%
3%
3%
1%
2%
2%
1%
>600
0.00
0.00
0%
0%
0%
0%
0%
0%
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category
Category
Category
Category
Category
0-50
34%
44%
33%
35%
39%
51%
46%
53%
50-100
25%
24%
24%
25%
22%
21%
21%
20%
100-200
33%
26%
26%
32%
29%
22%
25%
21%
Distance
to Cover
Nonmigratory
Elk
Distance
to Cover
200-400
9%
6%
6%
8%
10%
6%
7%
6%
400-600
0.00
0.00
0.00
0%
0%
0%
0%
0%
>600
0.00
0.00
0.00
0%
0%
0%
0%
0%
91
Appendix 9. Habitat use by elk relative to distance to shrub/scrub in the combined Randle
and Packwood study areas at the 50% and 95% range scale in the Southwestern Cascades
of Washington, 2007-2008.
Migratory
Elk
Distance
to Shrub
scrub
Winter
50%
Spring
50%
Summer
50%
Fall
50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category Category
Category Category
Category
0-50
59%
65%
88%
63%
52%
61%
83%
52%
50-100
11%
10%
4%
10%
11%
10%
5%
11%
100-200
17%
14%
5%
15%
17%
14%
6%
17%
200-400
10%
8%
2%
8%
12%
10%
4%
12%
400-600
2%
2%
0%
2%
4%
3%
1%
4%
>600
1%
1%
0%
1%
4%
3%
1%
4%
Winter
50%
Spring
50%
Summer
50%
Fall
50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Nonmigratory
Elk
Distance
to Shrub
scrub
Category Category
Category Category
Category
0-50
61%
32%
63%
57%
56%
49%
59%
48%
50-100
10%
14%
10%
11%
12%
13%
11%
13%
100-200
16%
27%
15%
18%
18%
21%
17%
22%
200-400
12%
26%
11%
14%
13%
15%
11%
16%
400-600
1%
1%
1%
1%
1%
2%
1%
2%
>600
0%
0%
0%
0%
0%
1%
0%
1%
92
Appendix 10. Habitat use by elk of slopes in the combined Randle and Packwood study
areas at the 50% and 95% range scale in the Southwestern Cascades of Washington,
2007-2008.
Migratory
Elk
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% Slope
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category
Category
Category
Category Category
0-1
14%
11%
1%
5%
11%
10%
1%
3%
1-15
56%
52%
15%
37%
53%
51%
14%
29%
15-30
18%
21%
23%
28%
21%
22%
20%
27%
30-45
8%
10%
26%
17%
10%
11%
27%
22%
45-25000
4%
6%
34%
13%
6%
7%
38%
19%
Nonmigratory
Elk
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% Slope
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category
Category
Category
Category Category
0-1
16%
27%
22%
10%
14%
18%
14%
8%
1-15
62%
61%
62%
56%
63%
63%
63%
56%
15-30
11%
7%
8%
16%
14%
11%
14%
20%
30-45
8%
4%
6%
13%
7%
5%
7%
11%
45-25000
3%
1%
2%
5%
3%
2%
3%
5%
93
Appendix 11. Habitat use by elk relative to distance to water in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern Cascades of
Washington, 2007-2008.
Migratory
Elk
Distance
to Water
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category Category
Category
Category Category
Category Category
Category
0-50
14%
13%
3%
14%
14%
14%
4%
10%
50-100
13%
12%
4%
13%
12%
13%
5%
9%
100-200
25%
24%
11%
25%
23%
24%
13%
20%
200-400
27%
28%
25%
27%
26%
26%
25%
28%
400-600
13%
14%
28%
14%
13%
13%
22%
17%
>600
1%
1%
28%
1%
11%
11%
31%
16%
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Nonmigratory
Elk
Distance
to Water
Category Category
Category
Category Category
Category Category
Category
0-50
10%
17%
8%
12%
13%
13%
10%
14%
50-100
8%
12%
7%
9%
10%
10%
8%
11%
100-200
18%
21%
15%
19%
18%
18%
16%
0.19
200-400
25%
24%
25%
25%
23%
23%
23%
0.23
400-600
13%
10%
14%
12%
13%
13%
14%
12%
>600
26%
32%
32%
23%
24%
24%
29%
22%
94
Appendix 12. Habitat use by elk relative to distance to paved roads in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008.
Migratory
Elk
Winter
50%
Spring
50%
Summer
50%
Spring
50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
Distance to
Paved Roads
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category
Category
Category
Category
Category
0-600
65%
61%
18%
40%
61%
55%
13%
33%
>600
35%
39%
82%
60%
39%
45%
87%
67%
Winter
50%
Spring
50%
Summer
50%
Spring
50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category
Category
Category
Category
Category
0-600
655%
82%
50%
59%
61%
67%
45%
56%
>600
35%
18%
50%
41%
39%
33%
55%
44%
NonMigratory
Elk
Distance to
Paved Roads
95
Appendix 13. Habitat use by elk relative to distance to non-paved roads in the combined
Randle and Packwood study areas at the 50% and 95% range scale in the Southwestern
Cascades of Washington, 2007-2008.
Migratory
Elk
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
Fall 95%
95%
% in
% in
% in
% in
% in
% in
Category
Category
0-100
7%
8%
4%
10%
0.10
12%
4%
9%
100-200
7%
8%
4%
9%
0.09
10%
4%
9%
200-400
15%
17%
11%
19%
0.16
18%
8%
16%
400-600
20%
21%
15%
23%
0.17
18%
10%
17%
>600
52%
46%
65%
39%
0.49
42%
74%
49%
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
% in
% in
% in
% in
% in
% in
Category
Category
0-100
3%
5%
3%
4%
6%
7%
6%
5%
100-200
8%
13%
7%
10%
9%
11%
9%
7%
200-400
24%
34%
21%
28%
23%
26%
24%
19%
400-600
15%
17%
14%
16%
14%
16%
15%
13%
>600
49%
30%
55%
43%
49%
40%
46%
56%
% in
% in
Distance to
Non-paved
Roads
Nonmigratory
Elk
Category Category Category Category Category Category
Summer
Fall 95%
95%
Distance to
Non-paved
Roads
96
% in
% in
Category Category Category Category Category Category
Appendix 14. Habitat use by elk relative to distance to trails in the combined Randle and
Packwood study areas at the 50% and 95% range scale in the Southwestern Cascades of
Washington, 2007-2008.
Migratory
Elk
Distance
to Trails
Winter
50%
Spring
50%
Summer
50%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Category
Category
Category
Category Category
Category
Category Category
0-100
0%
0%
3%
1%
1%
1%
4%
2%
100-200
0%
0%
3%
1%
1%
1%
4%
2%
200-400
0%
0%
9%
5%
2%
3%
9%
5%
400-600
0%
0%
8%
5%
1%
2%
6%
4%
>600
99%
99%
77%
88%
95%
93%
77%
87%
Winter
50%
Spring
50%
Summer5
0%
Fall 50%
Winter
95%
Spring
95%
Summer
95%
Fall 95%
% in
% in
% in
% in
% in
% in
% in
% in
Distance
to Trails
0.00
0.00
0.00
0.00
Category
Category
0-100
0.00
0.00
0.00
0.00
0%
0%
0%
0%
100-200
0.00
0.00
0.00
0.00
0%
0%
0%
0%
200-400
0.00
0.00
0.00
0.00
1%
0%
1%
0%
400-600
0.00
0.00
0.00
0.00
1%
0%
1%
0%
>600
0.00
0.00
0.00
0.00
97%
100%
98%
100%
NonMigratory
Elk
Category Category
97
Appendix 15. R2 values for Individual elk by habitat attribute at both the 50% and 95%
scales.
Regression with Nagelkerke Pseudo-R2 > .2
Freq50
Habitat
Attribute
Aspect
Slope
Collar ID
0.199
82M
0.203
28M
0.342
0.454
31M
0.407
0.383
0.333
82M
0.266
0.285
83M
0.665
0.528
84M
0.568
0.432
86M
NonPaved
0.32
419N
0.348
421M
0.257
28M
0.294
421M
0.259
28M
31M
0.428
0.366
0.385
0.345
81N
0.207
82M
0.737
83M
0.208
86M
0.205
0.484
80M
98
Pseudo R2
28M
80M
Cover
Freq95
0.543
0.411
Regression with Nagelkerke Pseudo-R2 > .2
Freq50
Habitat
Attribute
Paved
Collar ID
Trail
0.467
80M
0.341
82M
0.226
83M
0.576
0.477
84M
0.598
0.527
0.588
421M
0.575
0.372
422M
0.293
0.257
80M
0.233
83M
0.417
0.306
84M
0.329
0.334
85N
0.26
421M
0.206
0.308
422M
0.263
0.292
28M
0.621
31M
0.299
84M
0.27
419N
Water
Pseudo R2
30M
86M
Shrub
Freq95
0.36
421M
0.226
422M
0.566
28M
0.288
0.352
83M
0.653
421M
0.203
0.484
99
Appendix 16. GPS collared elk data points per migratory status in the Randle subunit
study area.
100
