Migration Routes and Stopovers of North Puget Sound Snow Geese

Item

Title

Eng Migration Routes and Stopovers of North Puget Sound Snow Geese

Date

2017

Creator

Eng Stevick, P. Frank

Subject

Eng Environmental Studies

extracted text

Migration Routes and Stopovers of North Puget Sound Snow Geese

by
P. Frank Stevick

A Thesis
Submitted in partial fulfillment
of the requirements for the degree
Master of Environmental Studies
The Evergreen State College
June 2017

©2017 by P. Frank Stevick. All rights reserved.

This Thesis for the Master of Environmental Studies Degree
by
P. Frank Stevick

has been approved for
The Evergreen State College
by

________________________
John Withey, Ph. D.
Member of the Faculty

________________________
Date



iv

ABSTRACT
Migration Routes and Stopovers of North Puget Sound Snow Geese

P. Frank Stevick
Snow geese are one of the most abundant waterfowl species in North America and have
been studied for decades. However, their migratory routes and stopovers remain largely
unresolved due to their remoteness. Advancements in satellite tracking methods have
allowed researchers to study regional snow geese population migration patterns in much
more detail then presently possible. Washington State Department of Fish and Wildlife
captured seven lesser snow geese (Chen caerulescens caerulescens) in North Puget
Sound (NPS) Washington, USA, fitted them with PTT model satellite tags, and tracked
them from 27 February 2013 to 5 October 2014. Study geese were tracked using two
different migratory routes from their wintering grounds in the Fraser-Skagit region along
the border of Washington and British Columbia (BC), Canada to their breeding ground
on Wrangel Island, Russia. Five of the study geese used a Pacific coastal route to fly from
the Fraser-Skagit region to upper Cook Inlet in Alaska (AK), USA before cutting across
the SW corner of the AK, crossing the Bering Sea, and migrating along the northern coast
of the Chukchi Peninsula in Eastern Siberia to Wrangel Island. The second group used an
inland route and headed east towards Alberta, Canada prior to heading NW across the
Northern and Yukon Territories in Canada, across AK, the Bering Sea, along the coast of
Eastern Siberia and to Wrangel Island. Study geese were tracked to staging regions, areas
were geese were on the ground for 7 or more days, in the Fraser River delta in BC and the
Stikine River delta, AK. Stopovers regions, areas where geese were on the ground for 2
to 7 days, were located in the Stikine, Knik, Serpentine, and Bering River deltas in AK
and the Fraser and Klinakline Rivers in BC. Continuous satellite telemetry provided
original data for managers and researchers for spring migrations in the remote regions of
BC and AK and confirmed that Midcontinent and NPS snow geese migration routes
overlap. Further research is needed to determine if Midcontinent lesser snow geese are
using the same wintering grounds as NPS lesser snow geese, while current stopover and
staging data will help managers in their conservation efforts for the NPS population of
lesser snow geese.



v

Table of Contents
CHAPTER ONE: INTRODUCTION TO THE THESIS ...…………….…………….1
CHAPTER TWO: LITERATURE REVIEW………………………………………….4
Introduction……………………………………………………….……………….4
Snow geese Overview……….…………………………………………………….7
Taxonomy…………………………………………………………………….….10
Wrangel Island Lesser Snow Geese……………………………………….….….11
North Puget Sound Population of Snow Geese……….….……………….….….16
Migration and Timing……………………………………………………………18
Management of the Northern Puget Sound Lesser Snow Geese…...……………23
Research Needs……………………...…………………………………………...28
Conclusions………………………………………………………………………31
CHAPTER THREE: MANUSCRIPT…...……………………………………………33
Abstract………………………………………………….……………………….34
Introduction………………………………………………………………………35
Study Area……………………………………………………………………….41
Methods…..............................................................................................................41
Results……………………………………………………………………………44
Discussion…..........................................................................................................49
Management Implications……………………………………………………………….53
CHAPTER FOUR: DISCUSSION AND CONCLUSION……...……………….….55
WORKS CITED………………………………………………………….……………89



vi

List of Figures

Figure 1. Wrangel Island main snow geese nesting colonies….…….………….………63
Figure 2. Wrangel Island snow goose population trends 1975-2015…….……….…….64
Figure 3. North Puget Sound snow geese migration stopovers and winter ranger……...65
Figure 4. Wrangel Island snow geese migration routes……………...…………………66
Figure 5. Goose 127446 migration………………………………….…………….…….71
Figure 6. Goose 127446 stopover sites……………………………….…………………72
Figure 7. Goose 127447 migration……………………………………………………...73
Figure 8. Goose 127447 stopover sites………………………………………………….74
Figure 9. Goose 127448 migration……………………………………………………...75
Figure 10. Goose 127448 stopover sites…………………………...……………………76
Figure 11. Goose 127449 migration……………………………….……………………77
Figure 12. Goose 127449 stopover sites…………………………...……………………78
Figure 13. Goose 127450 migration……………………….……………………………79
Figure 14. Goose 127450 stopover sites………………………………...………………80
Figure 15. Goose 127452 migration route…………….….………….….………………81
Figure 16. Goose 127452 rest stops………………………………….………………….82
Figure 17. Goose 127454 migration…………….………...……….……………………83
Figure 18. Goose 127454 rest stops……………………………………...………………84
Figure 19. Stopover and staging area of all study geese: Analysis one……….…………85
Figure 20. Stopover and staging area of all study geese: Analysis two…………………86
Figure 21. North Puget Sound wintering areas……………………….……….…………87



v

List of Tables

Table 1. Table of commonly referenced location with GPS coordinates……………….67
Table 2. North Puget Sound captured lesser snow geese release sites………….……....68
Table 3. North Puget Sound PTT tracking statistics………. .…………….……….…....69
Table 4. North Puget Sound Satellite tagged lesser snow geese transmitter program….70
Table 5. North Puget Sound stopover sites………………………………………………88



vi

Acknowledgements

I would like to thank the individuals who provided sound council, encouragement, and
support while I worked towards the completion of my thesis and the Master of
Environmental Studies (MES) degree. This thesis would not have been possible without
them, and I am grateful for their advice.
I would like to thank the faculty and staff from The Evergreen State College MES
program. Dr. Erin Martin provided sound councel during my first year of the MES
program by encouraging me to find a thesis project that combined my experience and
interests, which ultimately lead me to this work. I would also like to thank Ted Whitesell
for his constructive feedback on my thesis prospectus and literature review during my
first year of the MES program. Having these two pieces written in their entirety helped
immensely in setting up the foundation for the rest of the thesis in year two of the MES
program. I would also like to thank my thesis reader and advisor, Dr. John Withey. His
timely advice and constructive feedback was instrumental toward the completion of this
thesis. Lastly, I would like to thank the MPA/MES writing assistants Rhianna Hruska and
Paige Anderson for looking over countless drafts of my writing and providing
constructive feedback for revisions.

I would like to thank Washington’s Department of Fish and Wildlife, in particular Joe
Evenson and his colleagues, for providing the data for this thesis.
To my family and friends. I want to thank my parents who have always supported me on
all my education endeavors over the years. To my neighbors and preschool parents who
offered their time and homes for childcare, you gave me the additional time I needed to
work on my thesis and provided a safe place for Eliot to play while I worked, thank you.
You saved me countless hours of working through the night to meet thesis deadlines.
Lastly, I want to thank my wife, Bethany, who encouraged me throughout the thesis
process and helped me stay motivated. I couldn’t have asked for a better partner and I
look forward to reclaiming our time together after so many evenings away from home.

Thank you all



vii

CHAPTER ONE: INTRODUCTION TO THE THESIS

Introduction
Snow geese (Chen caerulescens) are one of the most abundant species of waterfowl in
North America (Baldassarre 2014). Their numbers have increased dramatically in recent
decades as they adapted quickly to using agricultural fields for forage and favorable
nesting conditions on Arctic breeding grounds (Boyd and Cooke 2000; Hupp et al. 2001;
Pacific Flyway Council 2006). For example, the lesser snow geese (Chen caerulescens
caerulescens) population breeding on Wrangel Island, Russia has increased 25% from
2015 to 2016 and the Midcontinent population of lesser snow geese has increased 300%
since the 1970s (Abraham and Jefferies 1997; U.S. Fish and Wildlife Service 2016).
However, prior to the 1970s, some populations of snow geese were in danger of
becoming extinct. The Wrangel Island population of lesser snow geese was one of these
(Kerbes and Meeres 1999; Pacific Flyway Council 2006). Wrangel Island lesser snow
geese are currently in recovery, but they still represent the last major snow goose
population breeding in Asia and wintering in North America, making the population a
high priority for the Russian, Canadian, and U.S. wildlife service’s (Kerbes and Meeres
1999; Pacific Flyway Council 2006).
The Wrangel Island lesser snow goose population became a concern in the early
1970s when the population dropped to historically low levels from decades of
overharvesting (Baranyuk 1999, Kerbes et al. 1999, Baldassarre 2014). In an effort to try
and protect the species and promote recovery, the Soviet Union made Wrangel Island into
a State nature preserve (Bousfield and Syroechkovsky 1985). This helped protect the



1

population by eliminating goose hunting and egg harvesting. Removing most of the
introduced reindeer prevented them from destroying goose-nesting grounds. However, in
the 1980s the lesser snow goose population had dropped by 50% to fewer than 110,000
geese and did not appear to be recovering (Kerbes and Meeres 1999). It was during this
period that an international effort was made by Russia, Canada, and the United States to
improve and update information on the distribution, survival, and size of Wrangel Island
lesser snow geese in order to form better management and conservation plans to preserve
the species (Kerbes and Meeres 1999). Their objectives were met through the use of
cooperative neck banding programs from 1987 to 1992, the analysis of recovered leg
bands from the 1950s to 1980s, and annual ground surveys of nesting colonies (Kerbes
and Meeres 1999). However, these methods still left large gaps in the literature on
migration routes and behaviors of Wrangel Island lesser snow geese leaving their
breeding grounds and entering North America to winter. As a result, managers and
researchers have made a large number of assumptions about Wrangel Island lesser snow
geese migration. A different approach was needed to address these research gaps.
Preliminary satellite studies done by Takekawa et al. (1994) and Baranyuk and
Takekawa (1998) were the first to more accurately document the migration routes of the
Wrangel Island lesser goose population using a non-banding method. Their work found
previously undocumented migration routes, stopovers, and staging areas, which
addressed some of the research gaps from earlier banding studies. This data will help
managers maximize their conservation efforts by setting aside reserves, refuges, and
nature preserves, along with modifying hunting regulations to reduce hunting pressure
during key migration times and at key staging locations. However, further studies have



2

yet to be contributed to the literature with similar findings from satellite transmitter data
for the Wrangel Island lesser snow geese population. This thesis is designed to reproduce
their work with the Northern Puget Sound population of lesser snow geese and address
the following question: What are the migration patterns and behaviors of the Northern
Puget Sound population of lesser snow geese (Chen caerulescens caerulescens) in
response to temporal changes in the environment over the course of the 2013 migration
season based on satellite telemetry?
Satellite telemetry provides a useful and unbiased tool for describing migration
routes and identifying stopover sites over the large uninhabited wilderness areas
commonly found in our study area. The data provided by modern satellite telemetry
technology will be more accurate and precise then previous studies on Wrangel Island
lesser snow geese, which will enable future researchers to be more efficient with their
research efforts in future studies. For example, by locating new staging or stopover areas
with satellite data and then using traditional leg or neck banding techniques to
strategically band large numbers of lesser snow geese, researchers can begin to answer
questions about populations intermixing between flyways, conduct local movement
pattern studies (e.g., feeding patterns, inter and intra species behaviors), population shifts
due to climate change and mortality assessments. Using traditional banding methods is
cheaper than using an equivalent number of satellite tags, which allows researchers to
increase sample size at the same cost and to increase the statistical power of the study.
Hence, identification of new stopover sites and associated migration patterns of lesser
snow geese is important for the current and future management of the lesser snow geese
population.



3

This thesis introduces the reader to the general topic and the specific research
question being addressed within the thesis in Chapter 1. Chapter 2 is the literature review,
which covers the Wrangel Island lesser snow goose population and research gaps in the
literature up to 2016. Chapter 3 is a manuscript formatted for the Journal of Wildlife
Management to be submitted after the completion of this thesis and Chapter 4 is a more
in-depth discussion and conclusion section than was covered in the manuscript in Chapter
3.

CHAPTER TWO: LITERATURE REVIEW

Introduction
Snow geese are one of the most abundant species of waterfowl in North America
(Baldassarre 2014). Their numbers have increased dramatically over the last few decades,
since snow geese adapted quickly to use agricultural fields for forage and favorable
nesting conditions on Arctic breeding grounds (Boyd and Cooke 2000; Hupp et al. 2001;
Pacific Flyway Council 2006). In some areas of the United States, the population of snow
geese has increased by 300% since the 1970s; however, the average rate of population
growth over the last ten years has ranged from 4-8% per year depending on the
population (Abraham and Jefferies 1997; U.S. Fish and Wildlife Service 2016). In
Washington State, flocks of thousands of lesser snow geese migrating south from their
breeding grounds on Wrangel Island, Russia can often be seen landing at their wintering
grounds on the Fraser and Skagit River deltas along the British Columbia (BC) border.
This population of lesser snow geese represents the last major snow goose population


4

breeding in Asia and the primary Russian goose population that winters in North
America, making the species a high priority for the Pacific Flyway Council and the
subject of long standing international cooperative management and conservation efforts
with Russian, Canadian, and U.S. wildlife service’s (Pacific Flyway Council 2006).
Since the early 1970s, when this population was at a historical low, the Russian
and Canadian wildlife services used banding studies and nesting ground surveys to track
population changes over time. The ground surveys show the current population of
Wrangel Island snow geese in recovery, but a recovering population presents a new set of
problems and questions for researcher and managers. For example, large flocks of
recovering snow geese are now overbrowsing their natural resources (mainly marsh
plants in estuaries) in some areas and shifting their foraging habits to agricultural
products, creating conflicts between conservationists, managers, and agricultural land
owners. In addition, the effects of overbrowsing can have deleterious effects on the
ecological processes, species and habitats in the marshes where snow geese are feeding.
However, ground surveys do not give any insight as to how snow geese are dispersing
from their breeding grounds and migrating south to their wintering grounds in North
America. The Canadian Wildlife services have conducted banding studies of Wrangel
Island snow geese and determined that 90% of the Wrangel Island lesser snow goose
population uses a coastal migration route to transit from their breeding grounds on
Wrangel Island to their wintering grounds in North America (Armstrong et al. 1999). The
other 10% was shown to take an inland route down through Alaska (AK) into their
wintering grounds in the southern regions of North America. Further research into the
migration patterns of the Wrangel Island lesser snow geese to gather more accurate and



5

precise data is difficult due to the remoteness and inaccessibility of the wilderness areas
in Canada, AK, and the Bering Sea the geese fly over.
The Washington State Department of Fish and Wildlife (WDFW) captured
Wrangel Island lesser snow geese overwintering in the Puget Sound region of the state in
2013 and implanted them with satellite transmitters. By utilizing satellite transmitters
WDFW was able to overcome the limitations of previous traditional banding studies and
collect migration data in the remote and inaccessible areas of Canada, AK, the Bering
Sea, and Eastern Siberia. Data collected from the geese implanted with a satellite
transmitter will provide a wealth of original data on Wrangel Island snow geese
migration patterns that can be used for management purposes or serve as baseline data for
future studies investigating migration patterns of lesser snow geese.
The purpose of this literature review is to summarize what is currently known
about the migration routes, stopovers, and management of Wrangel Island lesser snow
geese wintering in North Puget Sound. The first section will be an overview of the
general migration behaviors and patterns relevant to all snow geese populations. The next
section explains the taxonomy of snow geese and why the literature uses two scientific
names for the lesser snow geese covered in this thesis. The subsequent section will
describe the causes and consequences of changes in the Wrangel Island snow geese
population specifically and the known migration destinations of this population. The
following sections will focus on the WI population wintering in North Puget Sound and
management practices for the population. The last portion of the literature review will
outline the current research gaps, how this research will fill some of these gaps and make
suggestions for future research.



6

Snow Geese Overview
Snow geese are one of the world’s most abundant waterfowl species and one of the most
well studied birds in North America (Baldassarre 2014). Their distribution is split into
two seasons. During the breeding season (spring and summer months), much of their time
is spent in northern latitudes in order to find mates, raise chicks, and molt (losing and
then regrowing their flight feathers) in large colonies on the open tundra. During this
time, they range across the Arctic from Wrangel Island (Eastern Russia), the coastal
regions and islands in the Arctic and subarctic areas across AK and Canada, to the
northwestern coast of Greenland (Bechet et al. 2004; Baldassarre 2014). During the fall,
snow geese undertake extensive migrations from their Arctic breeding grounds to the
mid-Atlantic seaboard states, the lower Mississippi River valley, the Gulf Coast, the
Central Valley of California, and northern Mexico (Baldassarre 2014). Upon completing
their fall migration south, snow geese will spend the remainder of the fall and winter
months foraging to build fat reserves for the return trip back to the breeding grounds in
the spring.
There are two recognized subspecies of snow geese. The widely distributed and
smaller lesser snow goose (Chen caerulescens caerulescens) and the larger and more
discretely distributed greater snow goose (Chen caerulescens atlantica), whose breeding
distribution is restricted to a few islands in the Canadian high Arctic (Cooke et al. 1995;
Baldassarre 2014). Lesser snow geese are grouped into three populations groups based on
their breeding and or wintering grounds as follows: the Wrangel Island population, the
Western Arctic population, and the Midcontinent population (Hines et al. 1999;



7

Baldassarre 2014). A fourth population, the Western Central Flyway population, is solely
based on the winter distribution (Baldassarre 2014). Lesser snow geese breeding and
wintering grounds are not specific to only lesser snow geese. Intraspecies overlap with
other waterfowl populations is common. For example, Ross’ Geese (Chen rossii) and
lesser snow geese have overlapping wintering grounds in the Midcontinent and Western
Central Flyway populations.
All populations of snow geese have extensive migration, from their breeding
grounds to their wintering grounds. Round trips in excess of 11,000 kilometers (km) are
not uncommon, especially for those breeding on WI. In general, nonbreeding snow geese
will depart the breeding grounds first from mid to late August followed by the adults with
their young in early September (Bellrose 1980; Baldassarre 2014). The geese will head
south along either the Central or Pacific flyway to their wintering grounds in
southwestern United States and northern Mexico, making stops along the way to rest and
forage. Migration typically begins in the early daylight hours or from dusk to early
nocturnal hours, with flocks of thousands of individuals and family groups flying
continuously day and night until the geese reach their destination (Bellrose 1980;
Baldassarre 2014). Flocks of migrating snow geese have been observed flying at speeds
between 64 to 80 kilometers per hour (km/h) and tracked by radar at altitudes of 1,500 to
3,000 meters (m) with some reports from pilots observing snow geese as high as 6,100 m
(Bellrose 1980; Baldassarre 2014). However, most snow geese have a flying altitude
between 600 and 900 m during migration (Bellrose 1980; Baldassarre 2014). Alisauskas
et al. (2011) using band data from several lesser snow geese colonies in the central and
eastern Arctic from 1989 to 2006 during the fall migration, determined these flocks travel



8

through Canada at a rate of 25 km per day and 22 km per day through the United States.
During their migrations, snow geese fly in an undulating pattern with individuals
staggered at different heights throughout a U-shaped formation rising and descending
slightly. Snow geese rarely use the well-formed V formations of Canadian geese.
However, both formation styles have been showed to save energy on long flights (Denny
2016).
The timing of these migrations is dependent on population and location, but some
general associations for snow geese departing their wintering areas (northbound geese)
correlate with maximum daily temperature. For example, in Texas (1972-1977) lesser
snow geese initiated migration after temperatures exceeded 28 degrees Celsius (C) for
five days (Flicker 1981; Baldassarre 2014). Departures from wintering grounds have not
been strongly correlated with minimum temperatures, relative humidity, sky cover,
precipitation, surface wind, or atmospheric pressure (Blokpoel 1974; Blokpoel and
Gauthier 1975; Baldassarre 2014). For geese arriving at their wintering grounds
(southbound geese), evidence suggests their arrival could be correlated to higher mean
April temperatures melting snow earlier and thus increasing access to forage (Cooke et al.
1995). The most likely scenario is a combination of environmental factors (e.g.,
atmospheric pressure, wind, and sky cover), such as the passage of a frontal system,
initiates the start of a migration (Blokpoel and Gauthier 1975; Blokpoel 1974;
Baldassarre 2014).



9

Taxonomy
The lesser snow goose (Chen caerulescens caerulescens, Linnaeus, 1758) belongs in the
family Anatidae (ducks, geese, and swans) and in the Anserini tribe of True Geese
(Cooke et al. 1995). The Anserini tribe is divided into two genera: Chen (white and grey
geese) and Branta (black geese; Clements et al. 2016). Snow geese (Chen caerulescens)
are one of 11 recognized species within the genus Chen with two recognized subspecies:
the greater snow goose (Chen caerulescens atlanticus [Kennard 1927]) and the lesser
snow goose (Clements et al. 2016; Ottenburghs et al. 2016). The two subspecies can be
identified (as adults) by differences in size, range, and plumage color morphs. The lesser
snow goose is the smaller of the two subspecies, but has a much wider distribution across
North America and its plumage comes in two color phases: dark, sometimes referred to
as the, “blue phase” or “blue goose”, and all white. The greater snow goose is larger, its
plumage stays white all year and its distribution is limited to a few islands in the eastern
Arctic (Baldassarre 2014).
Within the literature, the lesser snow goose will also be found under the genus
Chen instead of Anser. Both scientific names (Chen caerulescens caerulescens and Anser
caerulescens caerulescens) are synonymous with the common name of lesser snow goose
and are genetically the same subspecies of goose. The difference in the naming
convention is explained by authors and journals choosing to classify the lesser snow
geese by the amount of white in their plumage instead of their genetic makeup (Cooke et
al. 1995; Ogilvie and Young 2002). The argument for using Chen over Anser is primarily
based on research that shows there is a high degree of hybridization within the Anser
genus resulting in genetic similarities in the different species and therefore the amount of



10

white in a bird’s plumage is a preferred tool for its classification and identification (Avis
et al. 1992; Quinn 1992; Cooke et al. 1995). In this literature review, I follow the
American Ornithological Society’s convention of using Chen (American Ornithologists'
Union 1983).

Wrangel Island Lesser Snow Geese
The Wrangel Island population of lesser snow geese (hereafter referred to as snow geese
or goose) represents the last major snow goose population breeding in Asia and the
primary Russian goose population that winters in North America. This distinction makes
it a high priority for the Pacific Flyway Council and the subject of long standing
international cooperative management and conservation efforts with Russian, Canadian,
and U.S. wildlife service (Kerbes and Meeres 1999; Pacific Flyway Council 2006). The
remaining breeding populations of snow geese in Russia are thought to still be nesting in
low densities of around 100 to 300 birds along the northern coastal mainland around the
Chukochya and Kolyma River deltas (Coordinates for geographical locations can be
found in table 1; Figure 3 [Baranyuk 1999; Kerbes, et al. 1999; Hines 1999; Baldassarre
2014]). Over the course of the last century, starting around the mid-1800s, the hundreds
of thousands of snow geese that breed all along the Russian Arctic coast from the Lena
River to the Northeastern coast of Siberia and wintered in Japan were extirpated by
humans (Takekawa et al. 1994; Baldassarre 2014). In 1976, Wrangel Island was
designated a State Nature Preserve by the Russian Ministry of Natural Resources and
Environment in order to manage and protect this last remaining population of snow geese
(Bellrose 1980).



11

Wrangel Island is an 800,000 hectares (ha) island located at 71˚ N off the coast of
northeastern Siberia and is surrounded by the Arctic Ocean (to the north), the Eastern
Siberian Sea (to the east), and the Chukchi Sea (Figure 1). Since 1969, the main nesting
colony of snow geese on the island have used the same 2,600 ha area, along the middle
reaches of the Tundra River valley in the Severnye Mountains (Pacific Flyway Council
2006). Starting in 1970, an annual survey was conducted on nesting adults using
systematic ground surveys of the main colony in the Tundra River valley (Bousfield and
Syroechkovsky 1985; Kerbes et al. 1999). The surveys were conducted in July after
broods had left the nest by running an 8m wide and 250 m long transects, spaced 200
meters apart through the entire colony, and counting current year nests (done by counting
the presence or absence of the egg membrane). The number of nests (two birds per nest)
were then averaged by transect and multiplied by the total area of the main colony to
estimate the islands breeding population of adults. The nonbreeding birds tend to
congregate in areas of low nest density prior to the arrival of breeding adults in July
(typically early June), which allows observers to make estimates of the populations using
binoculars and spotting scopes from nearby vantage points. The observers then add the
population number to the nesting survey for a total population estimate of the islands
Snow geese.
The results from these surveys show (Figure 2) a decline in the population of
snow geese from 1970 to 1975 from 150,000 to 56,000 (total population, breeding, and
nonbreeding), a recovery during the 1980s back up to a total population level close to
100,000 and then a decline again in the early 1990s to an average of around 65,000 total
birds (Bousfield and Syroechkovsky 1985; Kerbes et al. 1999). The population has



12

steadily been recovering since then, with the current population estimated to be around
300,000 birds (Pacific Flyway Council 2006; U.S. Fish and Wildlife Service 2016). This
amounts to an annual average growth rate of 8% over the last ten years, with a 25%
increase in population from 2015 to 2016 when the total population increased from
240,000 to 300,000 birds (U.S. Fish and Wildlife Service 2016). Key factors affecting
breeding population success and growth for Wrangel Island snow geese (WISG) are the
amount of snow cover in spring and the date by which the snow cover is cleared or
melted, allowing birds access to forage (Bousfield and Syroechkovsky 1985; Kerbes et al.
1999). These factors suggest that this population is growing from the effects of climate
change due to warmer annual temperatures melting snow earlier in the year (Hupp et al.
2001; Aubry et al. 2013). The effects of a warmer climate have also been linked to an
increasing population of younger snow geese in the total population relative to historical
total populations in the 1970s, 1980s, and early 1990s as reproductive success increases
by year (Pacific Flyway Council 2006). The consequences, if any, have not been well
studied.
The increasing WISG population is leading to changes in behavior relative to
traditional habitat and foraging resources. For example, WISG historically foraged
primarily in intertidal estuaries in the Fraser (BC, Canada) and Skagit River deltas
(Washington, USA) on plants such as the American three-square bulrush (Scirpus
americanus) during their migrations prior to the late 1970s (Boyd 1995; Boyd and Cooke
2000). After the 1970s, agricultural products (e.g. cereal, rye, barley, wheat, oats, etc.)
became more available to a growing WISG’s population that has increasingly become
reliant on agricultural products as available estuarine plants decreased due to



13

overbrowsing. Over the last twenty years, an increasing WISG population has gradually
shifted from their traditional feeding habitats in the marshes in the Skagit and Fraser
River deltas to more inland agricultural resources to increase their winter energy reserves
for the upcoming spring migration to the breeding grounds on Wrangel Island (Pacific
Flyway Council 2006).
Wrangel Island snow goose population dynamics are complex, with at least three
subpopulations migrating to different locations to overwinter (Kerbes and Meeres 1999;
Baldassarre 2014). The first is located in the Fraser and Skagit River deltas. The second
is located in the Central Valley in California. A third and much smaller population of
around 1,600 birds also winter along the lower Columba River in Washington and
Oregon. In the 1960s and 1970s, most (78% - 90%) of the WISG population wintered in
California and 10% wintered in the Fraser and Skagit River delta regions (Hines et al.
1999; Baldassarre 2014). Hines et al. (1999) used neck and leg banding field studies to
show the population had shifted to a more northern winter distribution in the 1980s with
47% of the WISG population wintering in California and 52% wintering in the Fraser and
Skagit River deltas. Boyd (1995) confirmed a northern shift in the WISG population
using data from annual photos. These photos show an estimated number of geese
wintering in the Fraser and Skagit deltas with 22% of the population wintering there in
1968 and increasing to 56% in 1992. A study by Boyd and Cook (2000) showed the ratio
has shifted to 60% in the Fraser-Skagit region by 2000. This result suggests a continuing
trend in the foreseeable future, but further studies are needed to confirm this trend from
2000 to present day.



14

Hines et al. (1999) hypothesized that this northward shift in the population hit its
peak in the 1970s, when snow geese first started to feed on agricultural fields and no
longer had to entirely depend on salt marsh habits. The timing of this peak shift in habitat
has been correlated to four consecutive years of reproductive failure due to weather
resulting in a depressed population and reduced competition for resources during their
migration southward (Hines et al. 1999). With limited competition during stopovers in
areas such as the Fraser-Skagit region on route to the Central Valley in California, many
birds opted to remain in the area rather than continue further south. However, with
contemporary populations on Wrangel Island at 300,000 birds, the population has
recovered and surpassed the WISG population estimates of the 1970s. With current high
populations of snow geese on Wrangel Island, the recovered population would imply that
the competition of the resources in the Fraser-Skagit region have increased since the
1970s and more birds would be heading further south to take advantage of the resources
in the Central Valley, a geographically larger area then the Fraser-Skagit region with
potentially more resources and less competition. However, the population trend through
2000 shows this assumption to be incorrect. There must be some other mechanism at
work to allow such a large population of snow geese to winter in the Fraser-Skagit region
and a continued northward shift in habitat from the Central Valley. Much of the current
literature suggests a connection to additional anthropogenic resources, such as increased
agricultural capacity and production of grains along with earlier snow melts, as possible
mechanisms for reducing competition through easier access to forage and further
increases in the carrying capacity of the Fraser-Skagit region (Boyd 1995; Hines et al.
1999; Boyd and Cooke 2000; Hupp et al. 2001; Kraege et al. 2008). Additional works by



15

Armstrong et al. (1999) and Williams et al. (2008) suggest a high philopatry rate of 74%
to 97% as an additional mechanism for the northward shift from the southern wintering
areas to the Fraser-Skagit region. Their findings suggest snow geese tend to stay in the
same wintering and breeding areas, unless various stochastic events, such as reproductive
failure due to weather, force them to leave the wintering and breeding grounds (e.g. in the
1970s). Once the geese moved to new northern wintering areas with improved weather,
their reproductive success increased on Wrangel Island. The total numbers of snow geese
increased at a higher rate in the Fraser-Skagit region than in the Central Valley due to the
new influx of previously displaced snow geese and their offspring. 74-97% of this group
stayed, further explaining the observed mechanism for the observed northward shift from
the Central Valley to the Fraser-Skagit region, but further studies are needed to verify
these findings.

North Puget Sound Population of Snow Geese
The North Puget Sound (NPS) population of snow geese winter in the Skagit and Fraser
River deltas along the western border of the United States and Canada. This population of
snow geese is referred to by its geographic location as the Fraser-Skagit region
population and not by the NPS moniker, but either name refers to the same population. I
use the term “NPS population” to refer to snow geese residing in Washington, since
wildlife managers in the State of Washington follow this convention.
The NPS population historically wintered on Fir Island and in the northern
portions of Port Susan Bay (Figure 3) and exhibited red facial staining due to the high
iron content of the intertidal marsh plants (bulrush; Scirpus americanus) they were



16

consuming (Boyd 1995; Baranyuk et al. 1999). Prior to the 1970s, this was a reliable
method to tell the NPS population apart from those continuing further south to the
wintering grounds in the Central Valley of California (Baranyuk and Syroechkovsky
1994; Baranyuk et al. 1999; Pacific Flyway Council 2006). After the 1970s, agricultural
products, such as cereals, became more readily available and many geese started to utilize
these alternative food sources, reducing the reliability of using face staining for visual
identification of the NPS population compared to the Central Valley population (Boyd
and Cooke 2000; Pacific Flyway Council 2006). However, body size can be used as an
alternative to face staining due to the NPS population’s significantly larger body sizes
(for both sexes) compared to wintering Central Valley geese using the Fraser-Skagit
region as a staging area (Baranyuk 1999; Pacific Flyway Council 2006). Since the early
1970s, the population of NPS snow geese has increased from a low of 12, 356 in 1974 to
an average of over 68, 517 between 2001 and 2005. Many NPS snow geese now seek
new food sources and expand their wintering range from traditional marshland habitats
on Fir Island and Port Susan Bay to more inland agricultural habitats in Skagit,
Snohomish, and Island counties in Washington (Pacific Flyway Council 2006).
The annual population growth prior to 2001 shifted from an average of 5% to an
average of 7% from 2001 to 2004. This rapid increase in population has put pressure on
snow geese foraging resources in the Puget Sound region and changed the composition of
the NPS population of snow geese (Pacific Flyway Council 2016). For example, in some
areas of the Skagit River delta, intense grubbing by large numbers of snow geese have
resulted in low biomass levels in the upper bulrush zone due to overgrazing and to the
introduction of invasive cordgrass (Spartina spp.) from the eastern United States (Boyd



17

1995). Furthermore, as natural resources in the Skagit River delta become less available
due to overgrazing, snow geese relied more heavily on anthropogenic resources, such as
agricultural crops, and urban green spaces to sustain a larger NPS population,
exacerbating depredation of these resources, and at times, causing friction between
farmers, hunters, birders, and managers (J. Everson, personal communication, November
1, 2016). WDFW has established a sanctuary on Fir Island (the 200 ha Hayton Reserve)
and planted it with winter cover crops favored by snow geese, which (as of 1996) has
mitigated friction between farmers and geese damaging their agricultural crops (Pacific
Flyway Council 2006).
The rapid reproductive success of the NPS population has also altered its
composition to include a larger component of younger birds than is typical in flocks prior
to the 1970s (Pacific Flyway Council 2006). The consequences of such reproductive
success remains unknown, but increased population growth can be expected as larger
cohorts of younger snow geese reproduce. However, the mechanisms of reproductive
success are well understood. They are linked to low predation rates on Wrangel Island,
early snow melts, and mild weather conditions on Wrangel Island, which allows snow
geese to produce larger clutches of eggs with lower mortality rates than in harsher
weather conditions (Hupp et al. 2001; Baranyuk 2005; Pacific Flyway Council 2006).

Migration and Timing
Wrangel Island snow geese undertake some of the longest migrations of any goose
population in North America with some individuals observed migrating 14,000 km
annually (Armstrong et al. 1999; Baldassarre 2014). However, the average distance for



18

the WISG population traveling to their furthest southern reaches of their breeding
grounds is 11,000 km (Armstrong et al. 1999). Much of what researchers currently know
about the migration patterns and behaviors of WISG relies heavily on various banding
studies (e.g., neck bands and leg bands), with the exception of a few studies using the
same satellite data from Takekawa et al. (1994). This section will review the available
literature and highlight some of the research gaps this study will address starting with the
fall migration from WISG.
Wrangel Island snow geese depart for their fall migration in late August and
follow one of two routes (Figure 4) to reach the wintering or staging grounds in the
Fraser-Skagit region or the Central Valley in California. The first birds start to arrive in
late September and continue to grow in numbers through October and early November
(Baldassarre 2014). Ninety percent of the WISG population follows the coastal routes to
reach the Fraser-Skagit staging region before 60% of the WISG take off again to head
further south to the Central Valley (Bellrose 1980; Armstrong et al. 1999; Hines et al.
1999; Boyd and Cooke 2000). These “coastal routes” are vaguely described in the
literature mainly due to the remoteness of the geographical areas WISG fly over, limiting
the amount of access researchers have to this population. For example, Bellrose (1985)
describes the primary fall migration route of WISG as, “across the Gulf of Alaska to
make landfall near the mouth of the Columbia River and on to Summer Lake, Oregon,
and the Klamath Basin”.
Syroechkovsky and Litvin (1986) used neck band data from the 1970s to
hypothesize that WISG did not use offshore migration routes and geese wintering in the
Central Valley migrated there via the Canadian prairies through Alberta and



19

Saskatchewan. Syroechkovsky and Litvin’s hypothesis was also vague on how geese
migrate from an island to the mainland without crossing any major bodies of water
(Syroechkovsky and Litvin 1986; Hines et al. 1999). Hines et al. (1999) and Armstrong et
al. (1999) using neck band and leg band data were able to describe that 10% of the WISG
population migrated from Wrangel Island through the Canadian prairies and then south to
the Central Valley of California, validating some of Syroechkovsky and Litvin’s (1986)
hypothesis. However, Hines et al. (1999) and Armstrong et al.’s (1999) hypothesis still
lacked specifics on coastal routes used by WISG to get to the Fraser-Skagit region. This
lack of specificity for fall migration routes is mainly due to the remoteness and
inaccessibility of the migration routes in northern Canada, AK, and northeastern Siberia
prohibiting any observer networks from reporting the locations of neck or leg banded
geese during their migrations through these areas (Armstrong et al. 1999). New methods
were required to gather more detailed information on WISG migration patterns to help
policy makers and managers make more informed decisions. In the early 1990s satellite
transmitter technology had advanced far enough to allow installation of satellite
transmitters on larger birds such as geese without interfering with their flight
characteristics. This allowed researchers to collect real time data in remote and
inaccessible areas such as the wilderness areas of Canada and AK.
Takekawa et al. (1994) and Baranyuk and Takekawa (1998) were the first to use
GPS satellite tags to track the fall migration of WISG. Their results characterized the fall
migration from Wrangel Island as a combination of long stopovers and rapid distance
flights between stops (Table 1 and Figure 3). The first major stopovers are on the
Chukotka (also known as Chukchi) Peninsula on the Siberian mainland (in the Cape



20

Billings and Koluthin Bay area). From there, the geese cross the Bering Sea to St.
Lawrence Island and then to the Seward Peninsula to stage within the Yukon-Kushokwin
River delta and the northern portions of the Alaskan Panhandle, which has large staging
areas at the mouth of the Yukon River and the southern coast of Norton Sound (Baranyuk
and Takekawa 1998; Pacific Flyway Council 2006; Baldassarre 2014). After leaving the
Yukon-Kushokwin River delta region, some flocks fly down to the Fraser-Skagit region
following the Alaskan and BC coastlines with some migrants making a final stop at the
mouth of the Stikine River near Wrangell, AK before heading on to their wintering areas
in the Fraser-Skagit deltas (Pacific Flyway Council 2006). Baranyuk and Takekawa
(1998) go on to describe that half of the satellite tagged coastal migrants headed toward
the Fraser-Skagit region along the coast turned east when they came near the Canadian
Queen Charlotte Islands (now known as the Haida Gwaii Islands) and crossed the Rocky
Mountains to the staging areas near Edmonton, Alberta in Canada (Konrad 1993). These
results contradict leg and neck bands studies by Armstrong et al. (1999), Hines et al.
(1999), and observations made by the Pacific Flyway Council (2009), which conclude
that 10% of the WISG population not using the coastal route migrate from Wrangel
Island and fly an inland route through the Northern Territories in AK, Alberta and
Saskatchewan, Montana, and eastern Oregon to the Klamath Basin and California (Figure
4). This thesis will contribute to their work by providing additional satellite tag data to
create a more accurate and precise picture of how WISG migrate to and from the FraserSkagit region.
The NPS population of snow geese over winter from late September to late
January, with geese departing for the spring migration in February and March (Pacific



21

Flyway Council 2006; Baldassarre 2014). The spring migrations are generally
hypothesized to follow the fall migrations in reverse order with some exceptions from the
Central Valley (Armstrong et al. 1999; Hines et al. 1999; Pacific Flyway Council 2006;
Baldassarre 2014). Researchers from the Pacific Flyway Council (2006) have
documented the NPS population of snow geese as having major staging areas in AK on
the Stikine River delta and the upper Cook Inlet with geese starting to arrive in late April.
In early May, the lower reaches of the Yukon River become available for staging areas
and become more populated with snow geese as food availability increases due to
melting snow and ice in the region (Armstrong et al. 1999; Hupp et al. 2001). After
feeding in staging areas in AK, the flocks travel to mainland Siberia and then on to
Wrangel Island to start breeding in late May with most of the various populations from
California, Oregon, and the Fraser-Skagit region arriving by June (Pacific Flyway
Council 2006; Baldassarre 2014). For the Central Valley, 74% of the WISG population
(other populations of snow geese also use these wintering grounds, e.g., Midcontinent
snow geese) wintering there follow an inland route starting in mid-April, with staging
areas on Freezeout Lake in Montana and in the Canadian prairies (Pacific Flyway
Council 2006; Baldassarre 2014). The rest of the WISG populations wintering in the
Central Valley follow a coastal route. Once any snow geese enter NW Canada, AK, and
Siberia the amount of available data decreases as the remoteness and inaccessibility for
humans increases, with the exception of satellite data (Armstrong et al. 1999; Pacific
Flyway Council 2006).
The timing of the Fraser-Skagit region snow geese has not been extensively or
specifically described in the literature. Temporal data has often been collected in



22

conjunction with other metrics for a snow goose study. However, general trends have
been noted from this surplus data collection on WISG populations and noted previously
in this literature review. Most snow geese follow the northward progression of snow melt
and use stopover areas shortly after open water and bare ground first becomes available
during spring migrations; suggesting that migration can be correlated to weather events
(Ryder 1967; Cooper 1978; Lincoln 1979; Raveling 1979; Wege and Raveling 1983;
Hupp et al. 2001). Specifically, for the WISG population, Armstrong et al. (1999) was
able to show seasonal variability for arriving times of wintering snow geese in the FraserSkagit region and the Columbia River with a leg and neck band data set from 1987 to
1992. The cause of this variability remained unexplored, but could be due to weather
events (e.g. maximum or minimum temperatures). More specific temporal mechanisms
and layover information during migration remain to be explored for the NPS.

Management of Northern Puget Sound Lesser Snow Geese
The management of the WISG population of snow geese is a complicated, international
multiagency affair because WISG migrate through three different countries within at least
two disjunctive regions of the Pacific Flyway (Syroechkovsky and Litvin 1986; Ely et al.
1993). Starting in Russia and traveling south, the WISG breeding grounds are managed
by the Russian Federation under the ministry of Natural Resources and Ecology. Since
1976, access to the island has been restricted to protect the unique natural systems by the
Soviet Union when it became a State Natural Reserve (Bousfield and Syroechkovsky
1985). Prior to 1976, the island contained at least two human settlements, a military radar
installation, and a large herd of introduced reindeer that were destroying large areas of



23

snow goose nesting areas. Post 1976, almost all human settlements were abolished and
the reindeer herd was reduced to 1,000 managed animals to protect the snow geese
nesting areas. In 1997, the reserve expanded to include 12 nautical miles of the
surrounding water and was extended to 24 miles in 1999. Hunting is currently not
allowed on the island, except for subsistence hunting for the local population and a
rotating staff of researchers visiting the island to conduct the annual snow geese ground
nest survey during the breeding season (Bousfield amd Syroechkovsky, 1985).
The Canadian Wildlife Service handles all matters of wildlife that belong to the
Canadian government, including the protection and management of all migratory birds
and associated habitats (Hines et al. 1999; Kerbes et al. 1999). The Canadian Wildlife
Service has conducted the bulk of the various snow geese banding studies used in this
literature review. In the United States, migratory bird management falls under the federal
jurisdiction of U.S. Fish and Wildlife Service, with individual states co-managing their
local waterfowl species and populations (U.S. Fish and Wildlife Service 2016).
In addition to state, federal and provincial management agencies, several
international administrative councils provide guidance for migratory bird populations
(Barlow 2016). These councils are divided up by flyway (Atlantic, Mississippi, Central,
and Pacific) and are composed of the director or an appointee from the public wildlife
agency in each state and or province in the western U.S., Canada, and Mexico (Barlow
2016). The Wrangel Island breeding population and the NPS population wintering in the
Fraser-Skagit region fall into two biological and administrative flyways due to their
migration routes and habitats, but in practice are placed in the Pacific Flyway with
additional representation from the Russian Federation, including representatives from the



24

Russian Academy of Sciences and the curator of the Wrangel Island Nature Preserve
(Barlow 2016; Pacific Flyway Council 2006). The intent of the council is to provide
general recommendations to the agencies for cooperative management plans, identify
common goals to foster continued collaborations for data collection and analysis, and
emphasize research needs (Barlow 2016). These management plans are non-binding
recommendations and do not require agencies to commit to specific goals or actions; but
are highly encouraged to do so (Barlow 2016). As a result, WDFW works in concert with
the U.S. Fish and Wildlife Service and the Pacific Flyway Council to manage its
migratory bird resources.
Management topics that are of increasing concern to managers of snow geese on
Wrangel Island and the Fraser-Skagit region include: overpopulation, shifting wintering
areas, overgrazing of marshlands, setting appropriate harvest rates, and population
dynamics (J. Everson, personal communication, October 1, 2016). For example, as the
population of WISG grows, managers have increasingly observed the overgrazing of
marshland habits, which over time will have deleterious effects on the aquatic and
terrestrial species using these ecological niches. In Washington, this could mean negative
impacts on endangered salmon populations using these wetlands as nurseries on their way
out to sea.
Furthermore, there is mounting evidence that the Canadian Arctic tundra is being
adversely affected around geese breeding areas from overgrazing. The large colonies of
geese forage by grubbing and pulling up the vegetative biomass, degrading soil quality
and increasing the rate of erosion (Abraham and Jefferies 1996; Abraham et al. 2005;
Abraham et al. 2012; Baldassarre 2014). Since snow geese exhibit a high degree of



25

fidelity towards their breeding areas, the problem of overgrazing will continue as geese
nest in the same Arctic locations for the foreseeable future (Willams et al. 2008;
Baldassarre 2014). Studies related specifically to the WISG population and overgrazing
have yet to be conducted, but inferences can be made about its effects from the Canadian
Arctic tundra studies and applied to the management regime for WISG and NPS
population to try to avoid a similar situation.
When populations of snow geese are considered too high by managers, various
methods of harvesting (e.g., sport and subsistence hunting and egg collecting by native
peoples) are initiated to control the population. The Pacific Flyway Council (2016)
recommends harvesting activities will be designed to meet the management plan
objective level of 120,000 geese per year for the entire population of WISG. Individual
agencies need to come up with harvesting regulations in their areas of jurisdiction. For
the Fraser-Skagit region, the council recommends the WISG population be maintained
between 50,000 and 70,000 geese (Pacific Flyway Council 2016). If the total WISG
population falls below 60,000, the council recommends all harvesting activities cease.
For the Fraser-Skagit region, the council recommends no harvesting activities if the
WISG population falls below 30,000. Other more extreme methods besides hunting are
available to regulate WISG populations levels, such as trapping geese on the breeding
grounds (birds are molting during this period and cannot fly). There is an emphasis on
increasing harvesting rates using hunting as the most desirable solution for reducing
overabundant populations (Johnson and Ankney 2003; Alisauskas et al. 2011).
WDFW has managed the growing NPS population of snow geese by increasing
harvest opportunities, while also facilitating a growing number of non-hunters seeking to



26

view the snow geese (J. Everson, personal communications, October 15, 2016). However,
increased harvest rates of the NPS population has not extinguished agricultural
depredation issues or decreased overgrazing of marsh vegetation communities (J.
Everson, personal communications, October 15, 2016). WDFW continues to invest
significant management resources to address these and other management concerns such
as the following (J. Everson, personal communications, October 15, 2016):
•

Establishing a snow goose reserve on Fir Island to mitigate damage to
private agricultural lands.

•

Developing a lease and cover crop program to provide winter forage for
snow geese to mitigate damage to private agricultural lands.

•

Implementing a special snow goose quality hunting program to increase
hunter harvest opportunity and maximize harvest rates.

•

Manipulating annual hunting regulations to maximize harvest rates while
balancing landowners’ concerns.

•

Facilitating a working group composed of landowners, hunters, and nonconsumptive users on Fir Island to address these issues and provide
recommendations to the Fish and Wildlife Commission.

•

Investigating whether the NPS population is a discrete population to avoid
overharvesting of non-NPS population snow geese.

•

Investigating migration routes, phenology, staging areas, stop-over
locations, and timing of migrations throughout the flyway(s).



27

The end goal of these management efforts is to avoid reactive management and
move forward with proactive and adaptive management strategies for Snow geese and
other species in a sustainable manner (J. Everson, personal communications, October 15,
2016).

Research Needs
After reviewing the literature, a number of research needs, gaps, and access to the
literature were identified. First, the Russians have conducted over three decades of
research on the WISG population; but most of the articles are published in Russian peerreviewed articles in Cyrillic, making it inaccessible for many English speakers to access
the primary literature sources. Only a few articles (e.g., Bousfield and Syroechkovsky,
1985) have been translated into English and are available to researchers outside of
Russia. However, I was able to access various conference notes that many of the Russian
researchers attended in English speaking countries summarizing their work.
Second, when the population of snow geese on Wrangel Island reached an all-time
low in the mid-1970s, a concentrated effort was made over the next 30 years by the
Russians and Canadians to understand population dynamics using leg and neck banding
data. All of the research done by Armstrong et al. (1999), Hines et al. (1999), and Kerbes
et al. (1999a and 1999b) included various banding methods. For studying migration
routes in the remote and inaccessible areas in Canada, AK, and the Bering Sea, these
methods have limitations. For example, banding relies on a network of observers to
report the location of banded birds. In remote areas such as the Canadian and Alaskan
wilderness with large geographical regions inaccessible to humans, gathering enough



28

data points to create an accurate migration route is difficult. Furthermore, leg and neck
band data can be of limited use for migration research because assumptions need to be
made on how birds fly from point A to point B. This is often assumed to be a straight
line, and therefore the shortest route between data points (Armstrong et al. 1999).
However, this is not always the case do to geographical interferences, such as mountains,
which birds may choose to fly around instead of over. As a result, as the distance
increases between relocations, it becomes more difficult to account for local changes in
behavior to weather (e.g., wind), topography (e.g., mountain ranges), and anthropogenic
impacts (e.g., hunting pressure). Lastly, distances for all birds need to be measured in
curved lines rather than straight lines, similar to airline miles due to the curvature of the
earth.
A more efficient method of tracking snow goose migration includes using radio
(VHS transmitters), radar tracking, or satellite tracking. These methods have been used to
study other populations of snow geese in the Central, Mississippi, and Atlantic flyways,
but rarely for the Pacific Flyway populations (Blokpoel 1974; Blokpoel and Gauthier
1975; Ely et al. 1993; Baranyuk and Takekawa 1998). Again, the remoteness and
inaccessibility of the Canadian, Alaskan, and Siberian wilderness makes it difficult to
apply the radio and radar tracking methods. Radio tracking requires personnel to either
set up remote radio listing posts or be near the radio tag (within line of sight), which is
difficult to do in areas with no roads or trails (Denny 2016). Radar tracking of snow
geese has also been done using ground based radar at airports, but in the vast wilderness
areas of the Pacific Flyway, provides little infrastructure for the use of other radar
resources such as weather radars, aircraft radar, and portable radars (Blokpoel 1974;



29

Denny 2016). However, radio transmitters can be useful to study local movements and
are currently in use by WFDW to study the NPS population (J. Everson, personal
communications, October 15, 2016 & unpublished data).
Satellite data collection for WISG was been conducted twice by Dr. John
Takekawa, who was with U.S. Fish and Wildlife at the time in 1991 (Konrad 1993). His
data set was collected in 1991 (30 tagged geese on Wrangel Island) and again in 1992 (24
tagged geese on Wrangel Island) using transmitters mounted on geese using back packs.
Most of his data set remains unpublished with the exception of two. One is published
only in Russian (Baranyuk and Takekawa 1998) and the other is a combined study using
satellite tags and VHS radio tags to study the movement patterns of snow geese in the
Yukon-Kuskokwin delta in AK (Ely et al. 1993). These studies were referenced in the
Pacific Flyway Counsil 2006 report to describe the migratory path of the WISG
population. Since then, no other studies have been done using satellite tags to study
WISG or NPS snow geese populations using satellite telemetry.
Current satellite technology has advanced considerably since the 1990s. For
example, satellite transmitters are now small enough to be implanted in geese to avoid
damaging the transmitter (e.g., geese biting of the antennas) with little harm on the bird’s
flight characteristics. In addition, battery life has increased, which allows tags to transmit
for longer periods, resulting in better data collection coverage. Given that only one set of
data has been collected using satellite tags to study the migration patterns of WISG (only
for fall migration), the sample size is small (n=54) in comparison to the total current
population of 300,000 geese currently breeding on Wrangel Island. Further studies are



30

needed to add to this data set in order to complete a more accurate picture of the
migration patterns of these geese.

Conclusion
The WISG geese population is recovering, but as their population increases, additional
biological, migratory, and population data is needed to make informed management
decisions. Past studies using banding and ground surveys have provided enough data to
make accurate population estimates, but not enough data to accurately hypothesize the
migration routes and stopovers of the WISG population. Accurately documenting
stopovers and migration routes of WISG populations will help managers know where to
maximize the conservation efforts when setting aside reserves, refuges, and nature
preserves. In addition, predicting the timing of the migration routes and stopovers would
allow managers to regulate hunting seasons more precisely, so hunting pressure would
not interfere with key migration times and staging areas. Satellite data, since it does not
require observers on the ground, will help to create a more accurate picture of WISG
migration and timing.
Preliminary satellite studies done by Takekawa et al. (1994) and Baranyuk and
Takekawa (1998) were the first studies conducted using satellite telemetry to more
accurately document the migration routes of the WISG population. Their findings
contradicted previous banding hypothesis on WISG migration, which suggests further
replication of these satellite studies are needed. This study aims to contribute to the
WISG satellite data set to create a more accurate and precise picture of their migration
patterns and stopovers by increasing the sample size of tagged geese in the literature.



31

Researchers and managers will be able to strategically band geese in newly discovered
stopovers and staging areas, which allows these stakeholders to answer questions about
populations intermixing between flyways, local movement pattern studies (e.g., feeding
patterns, inter and intra species behaviors, etc.), and population shifts and mortality
assessment. Furthermore, researchers can place their observer networks strategically in
these newly discovered (and accessible) WISG areas to report snow goose locations and
reduce the amount of human effort and research money it would take to study WISG
population dynamics and migration. For example, technologies that are currently on the
market, such as Radio-frequency identification (RFID) tags, will enable collection of
local movement data remotely: once an RFID tag (the size of a grain of rice) is injected
into an individual goose, the data can be transmitted from the field, up to a satellite, and
down to a researcher’s desktop. The next step is to initially gather enough data to locate
these remote staging and stopover areas to further study the WISG population patterns.



32

CHAPTER THREE: MANUSCRIPT

Formatted and prepared for: Journal of Wildlife Management
This manuscript is a preliminary draft submitted to fulfill graduation requirements for
The Evergreen State College Master of Environmental Studies program. The following
document has not been edited, reviewed, or otherwise endorsed by any of the listed coauthors and serves only to exemplify the potential final journal submission.
June 15, 2017
P. Frank Stevick
The Evergreen State College
2700 Evergreen Pkwy, Olympia, Washington 98505
206-240-7432
Stepau22@evergreen.edu

Migration Routes and Stopovers of North Puget Sound
Snow Geese
P. Frank Stevick,1 Department of Environmental Studies, Graduate Program on the
Environment, The Evergreen State College, 2700 Evergreen Pkwy, Olympia, Washington
98505
John C. Withey,2 Department of Environmental Studies, Graduate Program on the
Environment, The Evergreen State College, 2700 Evergreen Pkwy, Olympia, Washington
98505
Joseph R. Evenson,3 Washington Department of Fish and Wildlife, 600 Capital Way
North, Olympia, WA 98501, USA



33

ABSTRACT Snow geese have been extensively studied in the North America, but data
on migration routes, stopover sites, and staging areas remains incomplete for many snow
geese populations due to the remoteness of the regions they migrate over and the limited
access available for observer networks to report snow goose locations using traditional
banding methods. Advancements in satellite tracking methods have allowed researchers
to study snow geese migration patterns in detail in these remote and inaccessible regions.
Seven lesser snow geese (Chen caerulescens caerulescens) were captured in North Puget
Sound (NPS) Washington, USA, fitted with PTT model satellite tags, and tracked from
27 February 2013 to 5 October 2014. Study geese were tracked using two different
migratory routes from their wintering grounds in the Fraser-Skagit region along the
border of Washington State and British Columbia (BC), Canada to their wintering ground
on Wrangel Island (WI), Russia. Five of the study geese used a Pacific coastal route to fly
from the Fraser-Skagit region to the upper Cook Inlet in Alaska (AK), USA before
cutting across the SW corner of the AK, crossing the Bering Sea, and migrating along the
northern coast of the Chukchi Peninsula in Eastern Siberia to WI. The second group used
an inland route and headed east towards Alberta, Canada prior to heading NW across the
Northern and Yukon Territories in Canada, across AK, the Bering Sea, along the coast of
Eastern Siberia and to WI. Study geese were tracked to staging regions, areas where
geese were on the ground for 7 or more days, in the Fraser River delta in BC and the
Stikine River delta, AK. Stopovers regions, areas where geese were on the ground for 2
to 7 days, were located in the Stikine, Knik, Serpentine, and Bering River deltas in AK
and the Fraser and Klinakline Rivers in BC Continuous satellite telemetry provided



34

original data for managers and researchers for spring migrations in the remote regions of
BC and AK and confirmed that Midcontinent and NPS snow geese migration routes
overlap. Further research is needed to determine if Midcontinent lesser snow geese are
using the same wintering grounds as NPS lesser snow geese, while current stopover and
staging data will help managers in their conservation efforts for the NPS population of
lesser snow geese.

KEY WORDS Anser caerulescens caerulescens, Chen caerulescens caerulescens, lesser
snow goose, migration routes, North Puget Sound, satellite telemetry, stopover, Wrangel
Island.

Wrangel Island, Russia, lesser snow geese (Chen caerulescens caerulesnes) undertake
some of the longest migrations of any goose population in North America with some
individuals migrating 14,000 km annually (Armstrong et al. 1999; Baldassarre 2014).
However, the average distance for a Wrangel Island (WI) lesser snow goose (hereafter
referred to as WISG) migrating to their furthest southern reaches of their wintering
grounds in the southern U.S. is 11,000 km (Armstrong et al. 1999). During their long
migrations to and from their breeding grounds on WI to North America, WISG make
extensive use of stopovers and staging areas between long distance destinations to feed,
rest, and take shelter (Baranyuk and Syroechkovsky 1994; Boyd 1995; Boyd and Cooke
2000; Lok et al. 2011). However, the largely remote and inaccessible areas of the Alaska
(AK), USA and western Canadian wilderness areas that WISG migrate over make it



35

difficult to gather data on migration, stopovers, and staging areas for managers,
researchers and policy makers (Kerbes et al. 1999).
The WISG population is composed of 2 subpopulations breeding in one mixed
colony, but winter in geographically separate areas in North America (Bousfield and
Syroechkovsky 1985; Willams et al. 2008). In late August both subpopulations depart for
their fall migrations and follow one of two routes (Figure 4) to reach their wintering or
staging grounds in the Fraser-Skagit region along the BC and Washington boarder
(48°38'40"N, 122°32'33"W) or the Central Valley in California (Armstrong et al. 1999).
The first snow geese start to arrive in the Fraser-Skagit region in late September and
continue to grow in numbers through October and early November (Baldassarre 2014).
Ninety percent of the total WISG population migrates down the Pacific coast to reach the
Fraser-Skagit region to either winter there or use it as a staging area to continue further
south to the Central Valley (Bellrose 1980; Armstrong et al. 1999; Hines et al. 1999;
Boyd and Cooke 2000). Sixty percent of the WISG population that lands in the FraserSkagit region will use the area as a staging area before continuing south by various routes
to reach their wintering grounds in the Central Valley (40°12'N, 122°12'W) and form the
second of the two WISG subpopulations. Forty percent of the WISG population that
migrated down the Pacific coast will overwinter in the Fraser-Skagit region and form the
first WISG subpopulation we refer to as the North Puget Sound (NPS) population
(Armstrong et al. 1999; Hines et al. 1999; Kerbes et al. 1999). The remaining 10% of the
WISG population that does not migrate down the Pacific coast crosses the Bering Sea to
AK down through the Canadian prairies and then south to the Central Valley
(Syroechkovsky and Litvin 1986; Armstrong et al. 1999).



36

The NPS population of snow geese ovoverwinterfrom late September to late
January, with geese departing for the spring migration in February and March (Pacific
Flyway Council 2006; Baldassarre 2014). The spring migrations are generally
hypothesized to follow the fall migrations in reverse order with some exceptions from the
Central Valley (Armstrong et al. 1999; Hines et al. 1999; Pacific Flyway Council 2006;
Baldassarre 2014). Researchers from the Pacific Flyway Council (2006) have
documented the NPS population of snow geese as having major staging areas (Figure 3)
in AK on the Stikine River delta (56°33'28"N, 132°24'35"W) and the upper Cook Inlet
(61° 3'17"N, 150° 7'45"W) with geese starting to arrive in late April. In early May, the
lower reaches of the Yukon River (62°36'29"N, 164°53'3"W) become available for
staging areas and become more populated with snow geese as food availability increases
due to melting snow and ice in the region (Armstrong et al. 1999; Hupp et al. 2001).
After feeding in staging areas in AK, the flocks travel to mainland Siberia and then on to
WI (71°13'46"N, 179°25'39"W) to start breeding in late May with most of the various
populations from California and the Fraser-Skagit region arriving by June (Pacific
Flyway Council 2006; Baldassarre 2014).
The WISG population represents the last major snow goose population breeding
in Asia and the primary Russian goose population that winters in North America, making
WISG a high priority for international cooperative management and conservation effects
(Kerbes and Meeres 1999; Pacific Flyway Council 2006; Baldassarre 2014). The WISG
population became an ecological concern in the early 1970s when the population dropped
by over 50% from 150,000 geese to fewer than 60,000 with no signs of recovery
(Bayanyuk 1992; Kerbes and Meeres 1999). This lead to an extensive international effort



37

by Canada, Russia, and the U.S. to improve and update information needed to conserve
and manage the species in the 1980s. Neck and leg band data was used to address WISG
population size (Kerbes et al. 1999); distribution (Hines et al. 1999); mortality (Hines et
al. 1999); and routes and timing of migration (Armstrong et al. 1999). However, observer
networks for these banding studies had a difficult time gathering specific information on
WISG migration routes and stopovers due to the inaccessibility and remoteness of the
Canadian and AK wilderness areas WISG fly over. A new method was needed to gather
more specific migration data on routes, stopover locations, and staging areas for
managers to make more informed decisions about the WISG population.
Takekawa et al. (1994) and Baranyuk and Takekawa (1998) used GPS satellite
tags to track the fall migration of WISG and gather more specific migration data than
banding studies would have collected (Kerbes et al. 1999). Their results characterized the
fall migration from WI as a combination of long stopovers and rapid distance flights
between stops (Konrad 1993). The first major stopovers for the WISG are on the
Chukotka (also known as Chukchi) Peninsula on the Siberian mainland (Cape Billings
[69°51'0"N, 176°8'0"E]; (Figure 3). From there, WISG cross the Bering Sea to St.
Lawrence Island (63°24′0″ N, 170°10′0″W) and then to the Seward Peninsula, AK to
stage within the Yukon-Kushokwin River delta (60°20'25"N, 163°44'25"W) and the
northern portions of the AK Panhandle. The Yukon-Kushokwin River delta and the
northern AK Panhandle have large staging areas at the mouth of the Yukon River
(62°39'35"N, 165°5'14"E) and the southern coast of Norton Sound ([63°34'43"N,
162°31'13"E] Konrad 1993; Baranyuk and Takekaw 1998; Pacific Flyway Council 2006;
Baldassarre 2014). After leaving the Yukon-Kushokwin River delta region, some WISG



38

flocks fly down to the Fraser-Skagit region following the AK and BC coastlines with
some migrants making a final stop at the mouth of the Stikine River near Wrangell, AK
before heading on to their wintering areas in the Fraser-Skagit deltas (Pacific Flyway
Council 2006). Baranyuk and Takekawa (1998) described that half of the satellite tagged
WI coastal migrants (N>25 snow geese) headed toward the Fraser-Skagit region along
the coast and turned east when the flock was near the Canadian Queen Charlotte Islands
(Haida Gwaii [53°21'43"N, 132°15'22"W]) and crossed the Rocky Mountains to the
staging areas near Edmonton, Alberta in Canada (Konrad 1993). These results contradict
leg and neck bands studies by Armstrong et al. (1999), Hines et al. (1999), and
observations made by the Pacific Flyway Council (2009) researchers; which concluded
that 10% of the WISG population not using the Pacific coastal route migrate from WI and
fly an inland route through the Northern Territories in AK, Alberta and Saskatchewan,
Montana, and eastern Oregon to the Klamath Basin and winter in the Central Valley of
California.
An understanding of WISG stopovers and migration is critical for WISG
conservation and management. For example, spring stopover sites often provide the
energy and nutrients required for reproduction in migratory waterfowl on route to their
nesting grounds (Reed at al. 2004; Schmutz et al. 2006; Lok et al. 2011). Specifically, for
the WISG population locating stopovers and staging areas with high quality forage is
important to their survival during their exceptional long migrations and for WISG
management. For the NPS population many of these sites remain unknown due to the
remoteness of their migration route(s) and therefore make it necessary to use satellite
tagging methods to locate stopover sites and migration routes (Armstrong et al. 1999).



39

Previous works using satellite data have primarily focused on southbound migrations of
the WISG migration and not the NPS population migrating northward (Baranyuk and
Takekawa 1998; Takekawa et al. 1994). Thus, making original baseline satellite
telemetry provided from this study important for NPS snow geese managers. Data on new
NPS population stopovers and migration routes could be used for conservation purposes,
such as the setting aside of refuges in critical foraging areas or heavily used stopovers or
staging areas. Furthermore, regulators could use this data to update polices for harvesting
opportunities. For example, hunting seasons and locations could be changed so harvest
pressure does not alter the behavior of NPS geese to interfere with important migration
routes, stopover sites, and staging areas that would further increase NPS mortality during
migration. Managers could use the satellite data to define snow geese populations more
clearly by studying migration routes and looking for any overlap in flyway use. This is
especially important to prevent overharvesting of specific populations such as the WISG
wintering in the Fraser-Skagit region, where snow geese from multiple populations and
different flyways use local foraging resources (Kerbes et al. 1999).
Satellite telemetry provides an unbiased and valuable tool for describing
migration routes and locating stopover areas in the remote regions of the Pacific coast,
western Canada, AK, and Eastern Siberia. The specific objectives of this research were to
1) identify stopover sites used by the NPS population for both spring and fall migrations,
2) locate any additional wintering areas used by the NPS population outside of the FraserSkagit region, 3) intensify typical NPS migration routes for both spring and fall
migration, and 4) ascertain if the NPS population uses other flyways other than the
Pacific flyway during migrations.



40

STUDY AREA
We considered all territory in the Pacific Northwest, western Canada, AK, the Bering Sea
and Eastern Siberia as the study areas and examined stopover sites and migration routes
within these regions. The Pacific coastal regions of the Pacific Northwest, western
Canada, and AK are composed of large, complex archipelagos adjacent to a mountainous
mainland. These landforms create an extensive network of protected waterways and
complex nearshore habitats for a wide variety of bird species in the Pacific Flyway (Lok
et al. 2011). The interior region of the study area in western Canada is composed mainly
of boreal forests, arctic tundra, and prairie lands, while much of our study area in AK is
arctic tundra and boreal forests. The Bering Sea and Eastern Siberia region contains
various islands devoid of trees, which formed through the subduction of tectonics plates
and volcanism in the region, including WI on which the main colonies of the WISG
population breed.

METHODS
Capturing and Tracking
We captured 7 female snow geese on their wintering grounds in Washington between
February and March, 2013; using a general-purpose net gun after the conclusion of the
goose hunting season (Mehlmin and Shaiffer 1980). Individual geese were captured in
three wintering locations in Skagit (48°19'22"N, 122°21'49"W), Snohomish (48°12'36"N,
122°21'12"W), and Island Counties (48°13'28"N, 122°26'31"W) using a human spotter
on the ground to locate and identify lesser snow goose flocks and to guide the capture
crew into position to use the net gun. We tagged only females, as this study was part of a



41

larger study on female breeding, nesting habits and chick raising. When multiple geese
were captured in a single net only one female was selected at random to avoid tagging
multiple members of the same family group. Individuals selected for tagging were sexed,
weighted leg banded and implanted with satellite platform terminal (PTT) transmitters
(Model IMPTAV-2640, 42g, Telonics, Inc.) in the abdomen by a veterinarian following
established surgical protocols (Korschgen et al. 1996; Table 2). Marked birds were
released after a recovery period of less than 2 hours. We programmed PTTs with duty
cycles to be more frequent (4 hours on and 27 hours off) during the spring and fall
migrations and less frequent (2 hours on and 77 hours off) for non-migratory periods in
the winter and summer (Table 4). This study was part of larger study on female breeding,
nesting habits and chick raising and as a result, our sample of geese were required to be
female as part of these efforts.
The Argos location and data collection system was used to monitor snow geese
movements. Argos estimates PTT locations from the Doppler shift in transmitter
frequency received by National Oceanic and Atmospheric Administration (NOAA)
weather satellites as WISG approach and then move away from an individual PTT (Argos
Inc. 1996). The accuracy of each location is then classified based on the satellite-to-PTT
geometry during each satellite pass; the number of transmissions (messages) received per
satellite pass and the stability of the PTT transmission frequency (Argos Inc. 1996; Miller
et al. 2005). The Location Classes (LC) 3, 2, 1, and 0, are rated by the Argos data system
as <150, 150-350, 350-1000, and > 1,000 m, respectively. Accuracy was not provided for
LC A (3 messages received), LC B (2 messages), and LC Z (latitude and longitude often
provided if >1 message received). Location data usually included at >1 usable location



42

per bird in a transmission day. One Selected Location was chosen for each bird per
transmission day according to the nine criteria described in Miller et al. (2005), which
favors locations for LC 3, 2, and 1 (Lok et al. 2011). The Selected Location method by
Miller et al. (2005) was chosen to account for multiple locations per bird per transmission
day not being independent as well as Argos expression of accuracy as the probability that
67% of the locations will fall within stated limits; which therefore might make some
high-quality locations inaccurate and some poor-quality locations very accurate (Britten
et al. 1999; Hatch et al. 2000; Hays et al. 2001).
Data Analysis
We used Movebank (www.movebank.org), a free online infrastructure available
to all researchers for storage, managing, sharing and analyzing animal movement data, to
spatially and temporally filter PTT marked snow geese data from the Argos data system
to select only stopover and migration locations in our study area (Kranstauber et al. 2011;
Douglas et al. 2012). Data was further filtered using Movebank and Argos sorting and
filtering algorithms to look for and omit outliers in the PTT data. Outliers would include
incorrect locations caused by equipment or data processing problems, locations for which
the error is too large to properly analyze, or LC data that should not be considered part of
an animals track (e.g., negative altimeter reading for a migrating goose; Argos Inc. 1996).
Data was then transferred to ArcGIs 10.4 via a shapefile to process location data with the
Selected Location method to determine migration stopovers, staging areas, and migration
routes. We further defined our Selection Location method for staging and stopover sites
to favor LC 3, 2, 1 data as defined in Miller et al. (2005) and added the additional criteria
of including only LC data with a recorded zero m altitude reading. We defined a stopover



43

event as the act of an individual stopping at a site for a rest or foraging during migration
(Lok et al. 2011). Spatially, we defined stopover events as a series of ³2 consecutive
Selected Locations within 10 km of each other. For temporal use of sites, we classified
stopover events as a short stopover if the site was used for 2-7 days, and as a staging
stopover if the site was used for >7 days (Warnock and Bishop 1998; Lok et al. 2011).
Given the limited number of tagged individuals, we considered use by 2 individuals
(n=7) adequate to represent selection and use of a stopover or staging site. We increased
our Selection Location criteria for the migration analysis to also favor LC 0 data,
including any records of zero m in altitude meeting the 9 criteria set forth in Miller et al.
(2005), due to limited sample size and the very large geographical area covered in this
study. Locations for migration data was sorted by migration season (e.g., spring =
northward movements and fall = southward movements) and mapped.

Results
Satellite location data quality
PTT tag performance
PTT tag performance varied by goose during this study (Table 3), but none of the 7 PTT
tags were able to transmit for the intended 3 years of the PTT program (Table 4). Goose
127447 was able to transmit the longest from 27 February 2013 to 17 September 2014 for
330 days of satellite tracking (Table 3). Goose 127452 transmitted for the least amount of
time from 28 February 2013 to 30 April 2013 or 108 days of satellite tracking (Table 3).
Goose 127449 had fewer satellite tracking days (80), but transmitted longer temporally
by calendar year (88 verses 62 calendar days; Table 3) then Goose 127452. The largest
date set of combined Location Class’s (LC 3, 2 ,1,0, A, B and Z) came from goose



44

127447 (n=1924; Table 3). Reasons for termination of PTT signals are unknown (e.g.,
mortality, damaged PTT tag, negative effects of PTT tag on goose or PTT battery life)
with the exception of goose 12752, which was reported deceased to WDFW. Cause of
death was unreported.
Migration routes and chronology of migration
Characteristics of NPS migration routes and chronology
All 7 of the tagged geese survived the tagging process, over wintered in the Fraser-Skagit
region (Figure 3), and were able to start their migration back to the WI breeding grounds
(Figure 1). Geese 127446, 127448, 127449, and 127454 migrated up the Pacific coast
from the Fraser-Skagit region to upper Cook Inlet, AK before cutting across the
southwestern (SW) quadrant of the AK mainland to the Yukon River delta or Norton
Sound. PTT transmissions from goose 127449 ended at the Yukon River delta (Figure
11) after 26 May 2013 (Table 3), during the spring migration of season 2 of the PTT
program (Table 4). Goose 127448 (Figure 9) migrated past the Seaward Peninsula of AK,
across the Bering Sea to the Kolyuchinskaya Bay region of Russia, through northeastern
Siberia to the Cape Billing region, and then crossed the Eastern Siberian Sea to the
breeding grounds on WI. PTT signal was lost after 5 October 2013 (Table 3) during the
fall migration (Table 4). The Kolyuchinskaya Bay region was a common waypoint used
by 5 of the study geese. Goose 127446, after reaching the Norton Sound region, flew over
the Seward Peninsula of AK and across the Bering Sea straight to WI (Figure 5). After
the breeding season on WI, goose 127446 migrated south, for the fall migration of 2013
(Table 4), to Cape Billings, along the Eastern Siberian coast past Kolyuchinskay Bay,
across the Bering Sea, through SW AK, across the Gulf of Alaska, over Haida Gwaii, and



45

then followed the Pacific coast back to the wintering grounds in the Fraser-Skagit region
for the second time. Goose 127446, along with geese 127450, 127447, and 12754,
crossed the Gulf of Alaska during fall migrations, but not during the spring migration,
while using a more western Pacific coastal route then on the northbound (fall) migrations
(Figures 5, 7, 13, and 17). Goose 127446 accumulated the most days of satellite tracking
(342; Table 3), with the last signal received on 24 May 2014 near Upper Cook Inlet, near
Anchorage, AK during its second spring migration (Table 4). Goose 127454 was tracked
overwintering twice in the Fraser-Skagit region before the signal was lost after 7 June
2014 (Table 3) after Goose 127454’s second spring migration back to WI (Table 4).
Goose 127454 followed a coastal route to upper Cook Inlet, similar to geese 127449,
127448, and 127446, during the first spring migration (Table 4) and was the only goose
to have a large aggregation of LC data points (n=41) along the northwestern shoreline of
the Seward Peninsula before crossing the Bering Sea to Kolyuchinskaya Bay, Russia
(Figure 17). The migration plot suggests that goose 127454 crossed the Gulf of Alaska
twice, once during the second spring migration and once during the first fall migration,
but there are no data points between the Stikine River delta and landfall south of the
upper Cook Inlet to support the second spring Gulf of Alaska crossing (Figure 17).
Goose 127447 used a combination of inland and coastal routes. Goose 127477
used a Pacific coastal route similar to 127446, 127448, 127449, and 127454 during the
second spring migration (Table 4), but followed a more inland migration on the first
spring migration on the east side of the Canadian coastal mountain range (Figure 7). This
goose took an unusual migration path compared to the rest of the study group during the
first spring migration by flying west towards Norton Sound and then switching direction



46

northward and flying over Kotzebue Sound, AK and then crossing the Bering Strait to
Kolyuchinskaya Bay. The signal was lost on 17 October 2014 after tracking this goose
for 4 migratory seasons (Table 4) near Cape Billings, Russia, and after two breeding
seasons on WI and two wintering seasons in the Fraser-Skagit region.
Goose 127450 was the only goose to migrate to the Canadian prairies near
Alberta before proceeding to the Northwest and Yukon Territories and onward to WI
(Figure 13). This migration route would suggest there is some intermixing of the NPS
snow goose and the Midcontinent populations along the Pacific and Midcontinent
flyways, and at the wintering grounds in the Fraser-Skagit region. However, the second
spring migration track from the Fraser-Skagit region would suggest the migration route
towards Alberta during spring is not an annual occurrence, since the last data point on the
migration track would indicate a more coastal route then the previous spring migration.
Goose 12752 had the shortest migration track, using an inland Pacific coastal
route before heading east (Figure 15), when the signal terminated during the spring
migration of 2013 (Table 4). The last few data points for this goose would have had it
heading towards Alberta and the Canadian prairies, possible indicating further mixing of
the Pacific and Midcontinent flyway populations.
Stopover and Staging areas
Sites used as stopovers
Seven sites were identified as stopover sites (2-7-day usage) using the Selective Location
method with LC’s 1, 2, 3 data at an elevation of zero meters (n=1008). All of these
locations were located along the coastline of North America. Commonly used stopover
sites were the Fraser River delta, Bering River delta, and upper Cook Inlet (Figure 19).



47

Geese that used the upper Cook Inlet (127446, 127447, 127448, and 127454)
congregated in or near the Knik River delta. Less common sites were the Stikine and
Klinaklini River deltas in BC, and the Artic Lagoon and Serpentine River delta on the
Seward Peninsula in AK (Table 5). Geese that used a combination of both common and
less common stopover sites were 127447 (Figure 8), 127448 (Figure 10), and 127454
(Figure 18), over the course of the study period. Single stopover site users were geese
127449 (Figure 12), 127450 (Figure 14), and 127452 (Figure 16). The only goose that
used two stopovers was Goose 127446 (Figure 6).
Another stopover analysis was run using the Selective Location method with LC’s
1, 2, 3 data at an elevation of zero meters and changing the temporal setting to include all
data, i.e., not selecting data that only fell into the 2-7 or >7 days categories. This
increased the available stopover and staging area data by 61.5% (n=1639 instead of
n=1008), but represents 24.5% of the total LC data collected (n=6690); an increase of
9.5% in the amount of data used from the entire LC data set (Table 3) from the previous
analysis. The results show an increased use of the coastal regions and offshore areas, such
as the Chukchi Sea, the Bering Sea, and the Gulf of Alaska (Figure 20). Common
stopovers sites such as the Stikine River delta, the Bering River delta, and the upper Cook
Inlet show increased use, but for less time (>2 days), than the previous analysis. Other
less common sites along the coasts of the Seward Peninsula (i.e., Shishmaref Inlet,
Kotzebue Sound and Port Clarence) are short term stopover sites (>2 days) for geese.
New areas of use include the Kolyuchinskaya Bay, the coastal regions of Eastern Siberia
up to Cape Billings, and the offshore regions of Yakutat, AK (Figure 20). No inland
stopover or staging areas were documented using either of the two analyses.



48

Staging and wintering areas
The geese in this study used two staging areas: the Fraser River delta in BC and the
Stikine River delta (Figure 19). The Fraser River delta was used during both the fall
(September through October 2013) and spring (April 2013) migrations by geese 12746,
12748, and 127454 (Table 5). The Stikine River delta was used only during the spring
migration (April through May 2013, mainly around Dry and Farm Islands, near the town
of Wrangell, AK, by geese 12749 and 127454 (Table 5; Figure 19). Staging areas were at
times also used as wintering locations, which is the case with the Fraser River delta
(Table 5).
Wintering locations for the NPS population varied, but all geese except 127447
and 127449 split their time between the Fraser River delta and the Skagit River region
(Figure 21). The other five study geese (12746, 12748, 12750. 12752, and 127454)
overwintered in Washington in Skagit, Snohomish, and Island counties, primarily on
agricultural lands, intercostal waterways or tidelands. In Snohomish County, study geese
were tracked mainly around Fir Island and Skagit Bay. In Skagit County study geese
mainly overwintered in Port Susan Bay and at the mouth of the Stillaguamish River delta.
Island County contained the fewest LC data points, mainly out in the middle of Port
Susan Bay followed by the intertidal area near the city of Stanwood located at the
intersection of the Skagit and Island Counties at the south end of Skagit Bay.

DISCUSSION
Previous studies on WIGS migration patterns indicated that two migration routes were
used to migrate from the Fraser-Skagit region to WI (Baranyuk and Takekawa 1998;



49

Armstrong et al. 1999; Hines et al. 1999). Our results, based on continuous satellite
telemetry during migration seasons, also showed two primary migration routes were used
during the spring migration for the NPS population. Seventy-one percent (n=5) followed
a Pacific coastal route from the Fraser-Skagit region to upper Cook Inlet, AK before
cutting across the SW corner of AK and across the Bering Sea into Eastern Siberia to WI.
A second route through western Canada was used by 29% (n=2) of the snow geese during
the spring. However, we lost telemetry on goose 12752 before it completed its full
migration through BC, Canada. Taking a Pacific coastal route would provide several
advantages over taking the western Canadian route. First, a Pacific coastal route is more
direct and would be shorter than an interior route through the western providences of
Alberta, the Northern territories, and the Yukon territory of Canada. In general, shorter
distances would require less energy, if environmental stochastic events such as
precipitation, temperature, and headwinds had no effect on goose energy reserves.
Second, there would be more foraging opportunities along a Pacific coastal route then the
western Canadian route. The BC and AK coastlines provide abundant foraging
opportunities at ubiquitous river deltas formed by summer snow melt and receding
glaciers from the coastal mountain range. Different species of sedges such as Lyngbyaei
(Carex lyngbyeai) and Ramenski’s (Carex ramenskii) reside in these waterways and are
known to be part of the NPS snow geese’s main diet (Boyd 1995; Baranyuk et al. 1999;
Pacific Flyway Council 2006). NPS snow geese taking an inlet route through BC and into
central Alberta would find it more difficult to find forage sites because much of the
interior of BC is composed of forested landscapes and mountainous topography with less
access to primary food sources such as sedges or agricultural produces until you reach the



50

prairies of central Alberta. Our stopover data supports this observation because we
observed no stopover or staging area data along the interior migration route.
Fall migration results showed NPS snow geese did not follow the same migratory
route in reverse order of the spring migration route, which is different than previous
works (Baranyuk and Takekawa 1998; Armstrong et al. 1999; Pacific Flyway Council
2006). Fall migration routes would instead follow the northern coastline of Eastern
Siberia to Kolyuchinskaya Bay, Russia, cross the Bering sea then cross overland the SW
corner of AK and cross the Gulf of Alaska to make landfall near the Queen Charlotte
(Haida Gwaii) Islands, Canada before proceeding south via the Pacific coast to the
wintering grounds in the Fraser-Skagit region. This route was consistent for all the geese
we had multiple seasons of telemetry data for (n=4). Distances between data points
crossing the Gulf of Alaska (in some cases <2,000 km; goose 127450) would suggest that
our PTT duty cycles were not programmed to the optimal temporal period to maximize
migration data collection (i.e., PTT duty cycle was programmed to collect data too soon
or too late in the migration season) or this region has poor satellite coverage. However,
there was enough satellite coverage to record 3 out of the 4 snow geese (127446, 127450,
and 127454) stopping in the middle of the Bering Sea, suggesting that it could be possible
for NPS snow geese to complete extremely long segments of migration in excess of 2,000
km at a time. Completing such long flights would further suggest these geese have built
up enough energy reserves (i.e., lipids) in the breeding colonies to either not require as
many foraging stopovers as the spring migration or that their departure from WI required
them to expedite their migration to their wintering grounds and take the shortest route
possible to get there. Since, all the study geese were female an expedited departure due to



51

a late departure from the breeding grounds could explain why there is a difference
between fall and spring migration routes. Breeding female snow geese leave WI later in
the year because they have to raise their chick(s) until they are strong enough to migrate
(Baldassarre 2014). Nonbreeding snow geese don’t have to wait until so late into the year
to start their migration and tend to leave earlier then the breeding snow geese and thus
could take a different migratory route (Baldassarre 2014). Given our limited sample size
of 7 female snow geese and that we were unable to determine if any of the study geese
produced chicks on WI, further studies into WISG migration would benefit from
increased sample sizes, an increased duty cycle rate on PTT tags, and tagging both
breeding and nonbreeding snow geese of both sexes.
Stopovers were generally in the vicinity of river deltas or protected bays,
suggesting stopovers were selected for their forage availability. River deltas and
protected bays are typical environments for plant species such as Lyngbyaei (Carex
lyngbyeai) and Ramenski’s (Carex ramenskii) sedges to grow, which are known to be
part of WISG’s diet (Boyd 1995; Pacific Flyway Council 2006). If sites were selected for
other reasons such as fatigue or the need for shelter to avoid adverse weather conditions
our telemetry data would likely show a higher density of location data in areas such as the
middle of the Gulf of Alaska or the Chukchi Sea like some of the data from geese
127446, 127450, and 127454 did. However, stopover sites such as Shishmaref Inlet, Port
Clarence, and upper Cook Inlet have both topographical features to provide shelter and
habitat for foraging. Further investigation into environment stochasticity events such as
weather, tides, and temperature correlated with location data from a larger sample size
would benefit future studies of NPS stopovers.



52

MANAGEMENT IMPLICATIONS
The PTT data provided a wealth of original information about movement patterns of
WISG and NPS snow geese populations among flyway use areas. Previous studies using
banding techniques have been less effective in collecting migration data in the remote
and inaccessible regions of the western interior of Canada and AK, especially for spring
migration (Armstrong et al. 1999; Pacific Flyway Council 2006). The data showed a
small percentage (15%) of the snow geese overwintering in the Fraser-Skagit region use
an interior route through western Canada and AK to reach their nesting grounds at WI
during the spring migration. These geese were tracked to the prairies in central Alberta,
Canada, suggesting that snow goose populations from the Pacific (NPS and WISG
populations) and Midcontinent flyways could be intermingling during the spring
migration in this region (U.S. Fish and Wildlife Service 2016). For management
purposes, the mixing of different populations would require changes to existing harvest
strategies for the smaller WISG and NPS snow geese populations and increased harvest
monitoring of the larger Midcontinent populations to prevent overharvesting of WISG
and NPS snow geese.
Locating, monitoring, and documenting snow geese stopover and staging areas is
important for management to conduct due to the effects of anthropogenic influences, such
as climate change on snow goose habitat. For example, rising sea levels could
permanently flood existing salt marshes NPS snow geese depend on for forage such as
Lyngbyaei (Carex lyngbyeai) and Ramenski’s (Carex ramenskii) sedges in the Stikine
River delta staging area and upper Cook Inlet stopover (Pacific Flyway Council 2006).
Furthermore, changing precipitation patterns, such as reduced snow melt, could shrink



53

existing marshlands and river delta foraging sites causing snow geese to seek different
stopover and staging sites. These changing environmental conditions are part of a shifting
baseline of environmental changes due to climate change, and will require constant
monitoring to observe effects on NPS snow geese populations for future conservation and
management efforts (Pauly 1995).
The sample size (n=7) is too small to make any long-term predictions. However,
our stopover data would be useful for managers and researchers for future studies of
WISG. For example, using popular stopover regions, such as Kolyuchinskaya Bay,
Russia and Shishmaref Inlet, AK, to conduct habitat and local movement studies would
be an efficient use of a researcher or managers’ resources, while at the same time
increasing the probability of having a large enough sample size to make accurate
assessments and predictions. Furthermore, heavily used stopover or staging locations
would be ideal locations to test new methods of snow goose tracking technologies, such
as radio-frequency identification (RFID), to collect local movement data remotely.



54

CHAPTER FOUR: DISCUSSION AND CONCLUSIONS

Discussion

Spring Migration
Snow geese tagged from the NPS population (n=7) in Washington, USA generally took
two different migration routes from their wintering areas in the Fraser-Skagit region to
their nesting grounds on Wrangel Island, Russia. The main route used by most of the
snow geese during the spring (71%, n=5) followed a Pacific route coastline from the
Fraser-Skagit region into the upper reaches of Cook Inlet in Alaska (AK) before crossing
the SW corner of AK towards the Seward Peninsula. From the Seward Peninsula, snow
geese crossed the Being Sea to the Chukchi Peninsula, Russia, and followed the northern
coastline of Eastern Siberian to Wrangel Island. The second route used by 29% (n=2) of
the snow geese during the spring migration used an inland route through western Canada
from the Fraser-Skagit region to Wrangel Island. From the Fraser-Skagit, snow geese
flew in BC before heading east towards the prairie lands of Alberta, Canada. One goose,
12752, lost signal in BC for reason unknown and was reported deceased to Washington
Department of State Fish and Wildlife (WDFW). Goose 127450 continued to migrate
northward from Alberta into the Northern Territories and then west over the Yukon
Territories of Canada into AK (Figure 13). Goose 127450 continued over the northern
regions of AK across Kotzebue Sound, in the Bering Sea, over Kolyuchinskaya Bay,
Russia and along the East Siberian coastline to Wrangel Island. These findings support
the Pacific coastal route hypothesized by Armstrong et al. (1999) and Hines et al. (1999)
using leg and neck banding studies, but were unable to confirm due to the remoteness of


55

this region of BC and AK (Figure 4). These findings also confirmed some of the findings
by Baranyuk and Takekawa (1998), using early satellite tracking technology from the
early 1990’s.
Taking the Pacific Coastal route would offer snow geese some advantages over
taking an inland route through western Canada. First, the Coastal route in general is
shorter than the western Canadian route. Reducing the amount of distance traveled during
migration would be advantageous to snow geese because flying over shorter distances
will use less energy and saving energy increases a snow gooses’ fitness at the breeding
grounds at the end of the migration. Thereby, increasing the chances of successful
reproduction and reducing mortality of the population of snow geese on route to their
breeding grounds due to increased fitness. However, this scenario does not take into
account the effects of environmental stochasticity, such as weather and forage
availability. But if a straight line is drawn from Wrangel Island to the Fraser-Skagit
region, representing the shortest distance between these two points, the North Puget
Population (NPS) of snow geese follow a very similar route, which would suggest
distance is a strong influence on migration route regardless of any environmental
stochasticity.
A second advantage to using the Coastal route could be the availability of forage.
The temperate climate along the BC and southern AK coast often results in less
accumulation of snowfall due to the warming effects of the Pacific Ocean on atmospheric
temperatures in this region compared to the interior regions of BC and AK. Less snow
accumulation permits easier and earlier access to foraging sites during spring migrations.
Furthermore, the topography of the North American coastline from the Fraser-Skagit



56

region to the upper Cook Inlet in AK has far more river deltas containing intertidal marsh
plants, such as the bulrush (Scirpus americanus), typically consumed by NPS snow
geese, then the interior regions of BC, which consists mostly of temperate coniferous and
boreal forests and would be forage limited for snow geese (Boyd 1995). Stopover and
staging area data supports this hypothesis because there was no data for a western
Canadian migration route (Figures 19 and 21). All the stopover and staging area data was
along the Pacific coastal route. The migratory path of goose 127450 also supports the
hypothesis that the forested areas along the Pacific coastal route are resource limited.
The limited resources are due to goose 127450’s spring migratory route from the FraserSkagit region over the BC coniferous and boreal forests into the Canadian prairie lands in
Alberta, Canada, where other food sources, such as agricultural forage and grasses are
more readily available (Figure 13).
Weather and topography could both be influential on whether to take an inland
western Canadian spring migratory route or a Pacific coastal route. The ubiquitous
islands along the Pacific costal route from the Fraser-Skagit region to upper Cook Inlet
could provide some protection from the Bering Sea weather during the spring migration,
as would flying on the east side of the coastal mountains, such as goose 12447 did
(Figure 7). However, this study did not focus on the influences of migratory routes, but
rather the routes themselves. Weather data (e.g., snow accumulation, barometric pressure,
wind) is available for this data set on the movebank.org website from reputable national
weather resources, such as NOAA. The movebank.org website has the ability to cross
reference LC data points with the closed recorded weather phenomena and provide it for



57

analysis. Weather effects on migration would be a good topic for future research of NPS
migration.

Fall Migration
We collected fall migration data from 4 Platform Terminal Transmitters (PTT) from
geese 127446, 12447, 127450, and 127454. Each of these geese transmitted one season of
fall migration data before the PTT’s failed. The results from these 4 geese showed a
similar pattern of migration from Wrangel Island, Russia to the Fraser-Skagit region. The
geese started their fall migration from Wrangel Island and headed to the mainland at
Cape Billings on the northern Shores of Eastern Siberia. From Cape Billings, the geese
fallowed the coast west towards the Chukchi Peninsula to Kolyuchinskaya Bay, before
crossing the Bering Sea to the Seward Peninsula in AK and south across Norton Sound
and the SW portion of the state and cutting across the Gulf of Alaska and making landfall
near or at Queen Charlotte (Haida Gwaii) islands, Canada. After this point, the geese
followed a Pacific Coastal route to their wintering grounds in the Fraser-Skagit region.
A major difference between spring and fall migration routes was the Bering Sea
crossing during the fall migration. None of the spring migrates attempted to follow this
route. Furthermore, fall migratory data showed that geese made more use of the Eastern
Siberian coastline from Cape Billing to Kolyuchinskaya Bay during the fall then the
spring and all stops in this region were > 2 days in duration with no staging areas existing
in the region. The Eastern Siberian coastline was historically used for nesting and feeding
by WISG prior to their population being overhunted in the 19th century (Kerbes, Meeres
and Hines 1999). Its current continued use by WISG would suggest that this habitat is
still capable of supporting a recovering population of WISG. As the population of WISG



58

continues to grow at a decadal growth rate of 8%, the nesting colonies on Wrangel Island
could spread back into their historical breeding grounds as the population recovers.
Stopover and Staging Areas
Using the Select Locations method (Miller et al. 2005) and choosing only LC 1, 2 and 3
data at zero altitude reduced the number of available LC data points by 85% (n=1008) for
identifying stopover and staging areas along the migratory routes. Adding the step of
using only LC data with an elevation of zero meters was important because it was the
only way to confirm that a goose was on the ground, which would imply that the goose
was not flying and the assumption can be made that the goose was in a resting stage of
the migratory process. However, this resting assumption does not take into account the
effects of weather and other environmental stochasticity events that could force a goose
to seek shelter without the need to have to feed. Further data analysis using the
Environmental Data Automated Track Annotation System (Env-Data System) tool within
Movebank could be used to account for environmental stochasticity using weather data
and existing Argos LC data.
The results of the stopover data showed 62.5% of the stopovers were >2 days in
duration and generally located in the vicinity of river deltas or protected bays for all
geese (Figure 20). The most heavily used regions in AK were along the coast of Yakutat,
the Bering river delta, upper Cook Inlet, Port Clarence, and Shishmaref Inlet. In Russia,
the most heavily used regions were on the Chukchi Peninsula at Kolyuchinskaya Bay and
the coastal region at Cape Billings. I have no habitat data from these regions, but I would
hypothesis their close proximity to river deltas and protected bays would make these
areas ideal locations for the various marsh plants that make up the primary diet of a snow



59

goose. Furthermore, the Kolyuchinskaya Bay, Port Clarence and Shishmaref Inlet located
on either side of the Bering Sea, which is the longest leg of the spring migration route,
where forage would not be available, making it necessary to build up energy reserves to
complete this portion of the migration. My literature review sources as of June 2017 did
not describe snow geese using Kolyuchinskaya Bay, Port Clarence, Shishmaref Inlet, the
Bering River delta, or the coastal regions near Yakitat as stopovers, suggesting these
locations could be new information for either researchers or managers. However, this
information could be common knowledge to the Native Peoples or current residents
living in these regions and it has simply not been published or passed on to researchers
and managers due to the remoteness of these regions or language barriers (e.g., Russian
science literature that has not been translated to English).
Stopover data that fell into the ≤2 and ≥7 day criteria included the Arctic Lagoon
(Goose 127454; Figure 18), Bering River delta (Geese 127447; Figure 8), Klinaklini
River delta (127447), and the Serpentine River delta (Goose 127447; Table 4). These 4
locations were not described in any of the literature I reviewed, suggesting these locations
could also be new information for either researchers or managers. The only previously
described stopover location that was used by 3 of the study geese was upper Cook Inlet
located near Anchorage. Geese 127446 (Figure 6), 127448 (Figure 10), and 127454
(Figure 18) were tracked more specifically to the Knik River delta in the headwaters of
Cook Inlet during the spring migration (Table 4). Cook Inlet was not used as a stopover
or staging area during the fall.
The results of the staging areas data confirmed that WISG population in general
and the NPS population specifically used the Stikine River delta in AK as a staging area



60

only during the spring migration (geese 127454 and 12749). Geese crossing the Bering
Sea during the fall migration bypassed the Stikine River staging area and made landfall
near the Queen Charlotte (Haida Gwaii) Islands instead. After making landfall, these
geese preferred to make short (>2 days) stopovers, heading south along the Pacific coast
instead of stopping for longer periods of time at a staging area until they reached the
wintering grounds in the Fraser-Skagit region. The only exception of this observation was
goose 12746 (Figure 6), who used the Fraser River delta as a staging area prior to
heading further south into Washington to the Skagit River delta region to overwinter.
Future studies would benefit from a habitat and foraging analysis at staging and stopover
sites to further investigate why snow geese are using these sites.

Conclusions
This study has provided the first set of satellite telemetry data for NPS migration routes,
stopovers, and staging areas. The key findings from the data showed the following
results: 1) two migratory paths, the Pacific coastal route or the interior western Canadian
route, during the spring migration (north bound) and a single route during the fall
migration (south bound) over the Bering Sea and down the Pacific coast from the Queen
Charlotte (Haida Gwaii) Islands to the Fraser-Skagit region, 2) Midcontinent and NPS
snow geese populations could be intermingling during the spring migration over the
prairies in Alberta, Canada, suggesting managers should adjust their harvesting strategies
accordingly to avoid overharvesting of the NPS population, 3) most stopovers (62.5%)
were less than 2 days in duration and were located primarily at river deltas and protected
bays along the coastlines of Russia, Canada, and the U.S., 4) stopovers between 2 and 5



61

days were less common (37.5%) then rest stops that were less than 2 days, 5) new
stopover sites included the Arctic Lagoon, Serpentine River delta and Bering River delta
in AK and the Klinakline River delta in B.C., and 6) we confirmed previous finding that
WIGS use the Stikine River delta in B.C. as a staging area and specifically that the NPS
population uses this site as well. The results from these movement patterns will help
managers and researchers form a more successful conservation and management strategy
for NPS snow geese going forward.



62

Figure 1. Wrangel Island, Russia, the main breeding site of the Wrangel Island lesser
snow goose (Chen caerulescens caerulescens) population. The grey circles represent
colony sites regularly occupied during the breeding season from May to August
(Baranyuk 1999; Pacific Flyway Council 2006).



63

Figure 2. Wrangel Island, Russia lesser snow goose (Chen caerulescens caerulescens)
population trends from 1975 to 2016 (U.S. Fish and Wildlife Service 2016). No data was
collected in 2015, representing the gap in the trend line.



64

Figure 3. Known migration stopovers of North Puget Sound lesser snow geese (Chen
caerulescens caerulescens) and winter ranger in the Skagit River delta region (Kerbes et
al. 1999).



65

Figure 4. Wrangel Island, Russia lesser snow goose (Chen caerulescens caerulescens)
fall migration routes. Less than 10% of the Wrangel Island snow geese (WISG)
population migrate over the Chukchi Sea over Alaska, USA into the Canadian prairie
lands of Alberta and Saskatchewan before heading to their wintering grounds in Central
Valley, California, USA. More than 90% of the Wrangel Island population migrate down
the Pacific coast and use the Fraser-Skagit region as a staging area. Sixty percent of the
lesser snow geese using the Fraser-Skagit region will migrate further south to the Central
Valley to overwinter with the remaining 40% of the population staying within the FraserSkagit region for the winter (Armstrong et al. 1999; Hines et al. 1999; Pacific Flyway
Council 2006).



66

Table 1. Table of geographical locations with coordinates.


Stopover site
Alberta, Canada
Cape Billings, Russia
Central Valley, California
Chukochya River (Bolshaya) delta, Russia
Fir Island, Washington
Fraser River delta, BC, (Canada)
Fraser-Skagit region, Washington and BC
Freezeout Lake, Montana
Island County, Washington
Japan
Kolyma River delta, Russia
Lena River delta, Russia
Lower Columbia River, Washignton and Oregon
Lower Yukon River, Alaska
Norton Sound, Alaska
Port Susan Bay, Washington
Queen Charlotte Islands (Haida Gwaii) , BC, Canada
Saskatchewan, Canada
Skagit County, Washington
Skagit River delta, Washington
Snohomish County, Washington
St. Lawrence Island, Canada
Stikine River delta, Alaska
Summer Lake, Oregon
Tundra River Valley, Wrange Island, Russia
Upper Cook Inlet, Canada
Wrangel Island, Russia
Yukon River mouth (delta), Alaska
Yukon-Kushokwin River delta, Alaska

Coordinates
54.316°N, 144.677°E
69.850°N, 176.133°E
40.200°N, 122.240°W
48.350 N, 122.394 E
70.1 N, 159.7667 E
49.058°N, 123.177°W
48.644°N, 122.543°W
47.661°N, 112.051°W
48.224°N, 122.442°W
36.17°N, 138.237°W
69.30°N, 161.30°W
72.476°N, 126.633°W
46.225°N, 123.624°W
62.608°N, 164.884°W
48.180°N, 122.425°W
53.362°N, 132.256°W
63.579°N. 162.520°E
52.940°N, 106.451°E
48.323°N, 122.364°W
48.305°N, 122.381°W
48.210°N, 122.353°W
63.4°N, 170.167°W
56.579°N, 132.41°W
42.748°N, 120.489°W
71.433°N, 179.805°W
61.054°N, 150.129°W
71.229°N, 179.428°W
62.660°N, 165.087°W
60.969°N, 163.346°W







67

Table 2. Captured North Puget Sound lesser snow geese (Chen caerulescens
caerulescens) by individual satellite tag identifier, sex, mass, U.S. Fish & Wildlife
Service neckband number, and release site from 2013 to 2014.
Individuals
127446
127447
127448
127449
127450
127452
127454



Sex
Female
Female
Female
Female
Female
Female
Female

Mass (g)
2068
1972
1928
2042
1826
1978
2006

USFWSBAND #
1727-55979
2097-09240
2097-09252
2097-09245
2097-09256
127452
2097-09451

Release location
48.341°N, -122.386°W
48.330°N, -122.382°W
48.341°N, -122.386°W
NA
48.341°N, -122.414°W
48.545°N, -122.428°W
48.361°N, -122.417°W

68

Table 3. Information on North Puget Sound lesser snow geese (Chen caerulescens
caerulescens) implanted with Platform Transmitting Terminals from 2013 to 2014.

Deployment date

Individuals
127446
127447
127448
127449
127450
127452
127454
Total



start
(yyyy.mm.dd)
2013.02.27
2013.02.26
2013.02.27
2013.02.28
2013.02.28
2013.02.28
2013.03.01

end
(yyyy.mm.dd)
2014.05.24
2014.09.17
2013.10.05
2013.05.26
2014.03.22
2013.04.30
2014.06.07

location
data
points (n)
1016
1924
478
374
1007
221
1670
6690

Days of
satellite
tracking
342
330
98
80
201
108
292
1451

69

Table 4. Platform Transmitting Terminal program used for captured North Puget Sound
lesser snow geese (Chen caerulescens caerulescens) fitted with satellite tags from 2013
to 2014.

Season
1
2
3
4
5
6
7
8
9
10
11
12

Year
1
1
1
1
2
2
2
2
3
3
3
3

Transmit
start date
Feb-13
15-Mar-13
31-May-13
9-Aug-13
15-Oct-13
15-Mar-14
31-May-14
9-Aug-14
15-Oct-14
15-Mar-15
31-May-15

Transmit end
NonMigratory
date
migratory Migratory
season
15-Mar-13
X
31-May-13
X
Spring
9-Aug-13
X
15-Oct-13
X
Fall
15-Mar-14
X
31-May-14
X
Spring
9-Aug-14
X
15-Oct-14
X
Fall
15-Mar-15
X
31-May-15
X
Spring
9-Aug-15
X
End transmission due to low battery

Duty
cycle
(On/Off)
2/77
4/27
2/77
4/27
2/77
4/27
2/77
4/27
2/77
4/27
2/77





70

Figure 5. Migration route of female PTT-tagged lesser snow goose (Chen caerulescens
caerulescens) 127446 captured and tagged in the Skagit River delta on 27 February 2013
and tracked to 24 May 2014. Grey circles indicate previously documented rest and
staging areas (Baranyuk and Takekawa, 1998; Kerbes et al. 1999).



71

Figure 6. Stopover and staging sites (black dots) of female PTT-tagged lesser snow
goose (Chen caerulescens caerulescens) 127446 captured and tagged in the Skagit River
delta on 27 February 2013 and tracked to 24 May 2014. Grey circles indicate previously
documented stopover sites and staging areas (Baranyuk and Takekawa 1998; Kerbes et
al. 1999).



72

Figure 7. Migration route of female PTT-tagged lesser snow goose (Chen caerulescens
caerulescens) 127447 captured and tagged in the Skagit River delta on 26 February 2013
and tracked to 17 September 2014. Grey circles indicate previously documented rest and
staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



73

Figure 8. Stopover and staging sites (black diamonds) of female PTT-tagged lesser snow
goose (Chen caerulescens caerulescens) 127447 captured and tagged in the Skagit River
delta on 26 February 2013 and tracked to 17 September 2014. Grey circles indicate
previously documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et
al. 1999).



74

Figure 9. Migration route of female PTT-tagged lesser snow goose (Chen caerulescens
caerulescens) 127448 captured and tagged in the Skagit River delta on 27 February 2013
and tracked to 05 October 2013. Grey circles indicate previously documented rest and
staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



75

Figure 10. Stopover and staging sites (black crosses) of female PTT-tagged lesser snow
goose (Chen caerulescens caerulescens) 127448 captured and tagged in the Skagit River
delta on 27 February 2013 and tracked to 05 October 2013. Grey circles indicate
previously documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et
al. 1999).



76

Figure 11. Migration route of female PTT-tagged lesser snow goose (Chen caerulescens
caerulescens) 127449 captured and tagged in the Skagit River delta on 28 February 2013
and tracked to 26 May 2013. Grey circles indicate previously documented rest and
staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



77

Figure 12. Stopover and staging sites (black asterisks) of female PTT-tagged lesser
snow goose (Chen caerulescens caerulescens) 127449 captured and tagged in the Skagit
River delta on 28 February 2013 and tracked to 26 May 2013. Grey circles indicate
previously documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et
al. 1999).



78

Figure 13. Migration route of a female PTT-tagged lesser snow goose (Chen
caerulescens caerulescens) 127450 captured and tagged in the Skagit River delta on 28
February 2013 and tracked to 22 March 2014. Grey circles indicate previously
documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et al.1999).



79

Figure 14. Stopover and staging sites (black circles) of female PTT-tagged lesser snow
goose (Chen caerulescens caerulescens) 127452 captured and tagged in the Skagit River
delta on 28 February 2013 and tracked to 30 April 2013. Grey circles indicate previously
documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



80

Figure 15. Migration route of female PTT-tagged lesser snow goose (Chen caerulescens
caerulescens) 127452 captured and tagged in the Skagit River delta on 28 February 2013
and tracked to 30 April 2013. Grey circles indicate previously documented rest and
staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



81

Figure 16. Stopover and staging sites (black squares) of female PTT-tagged lesser snow
goose (Chen caerulescens caerulescens) 127452 captured and tagged in the Skagit River
delta on 28 February 2013 and tracked to 30 April 2013. Grey circles indicate previously
documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



82

Figure 17. Migration route of a female PTT-tagged lesser snow goose (Chen
caerulescens caerulescens) 127454 captured and tagged in the Skagit River delta on 1
March 2013 and tracked to 07 June 2014. Grey circles indicate previously documented
rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



83

Figure 18. Stopover and staging sites (black trianges) of female PTT-tagged lesser snow
goose (Chen caerulescens caerulescens) 127454 captured and tagged in the Skagit River
delta on 1 March 2013 and tracked to 07 June 2014. Grey circles indicate previously
documented rest and staging areas (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



84

Figure 19. Stopovers and staging sites (black dots) of all lesser snow geese (Chen
caerulescens caerulescens) tagged and released from the North Puget Sound population
in the Skagit River delta region in Washington State, USA Geese were tracked from 26
February 2013 to 5 October 2014. Grey circles represent previously documented staging
and stopover sites (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



85

Figure 20. Stopovers and staging sites (black dots) of all lesser snow geese (Chen
caerulescens caerulescens) tagged and released from the North Puget Sound population
in the Skagit River delta region in Washington State, USA Geese were tracked from 26
February 2013 to 5 October 2014. Grey circles represent previously documented staging
and stopover sites (Baranyuk and Takekawa 1998; Kerbes et al. 1999).



86

Figure 21. North Puget Sound wintering area of all 7 lesser snow geese (Chen
caerulescens caerulescens) tagged and released from the North Puget Sound population
in the Skagit River delta region in Washington State, USA Geese were tracked from 26
February 2013 to 5 October 2014.



87

Table 5. Locations of North Puget Sound stopover and staging sites by year and PTT tag
number.
Individuals by PPT tag number
Region
Coordinates
12746
British Columbia, Canada
Fraser River delta -123.134°W, 49.050°N X
Fraser River delta -123.206°W, 49.167°N
Fraser River delta -123.167°W, 49.087°N
Fraser River delta -123.167°W, 49.087°N
Fraser River delta -123.167°W, 49.087°N
Fraser River delta -123.134°W, 49.050°N
Fraser River delta -123.151°W, 49.071°N
Klinaklini River delta -125.092°W, 51.092°N
Stikine River delta -132.463°W, 56.583°N
Stikine River delta -132.463°W, 56.583°N
Stikine River delta -132.463°W, 56.583°N
Alaska, U.S.A.
Upper Cook Inlet -149.588°W, 61.483°N
Knik River delta -149.515°W, 61.48°N X
Knik River delta -149.444°W, 61.444°N
Serpentine River delta -165.553°W, 66.162°N
Arctic Lagoon
-166.492°W, 66.138°N
Bering River delta -144.353°W, 60.125°N
Bering River delta -144.284°W, 60.167°N



12747 12748 12749 127450 127452 127454

X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X

Date
(month/yy)
Oct-13
Oct-13
Mar-14
Sept-Oct-2013
Apr-14
Apr-13
Apr-13
Apr-14
Apr-May-2013
Apr-May-2013
Apr-13
Apr-14
Apr-May-2013
May-13
May-14
May-14
Apr-14
May-13

Staging Stoparea over
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X

88

Works Cited
Abraham, K. F., and R. L. Jefferies. 1997. "High goose populations: causes, impacts and
implications." In Arctic Ecosystems in Peril: Report of the Arctic Goose Habitat Working
Group., edited by B.D. J. Batt, 12-82. U.S. Fish and Wildlife Service, Washington, D.C.,
and Canadian Wildlife Service, Ottawa, Ontario: Arctic Goose Joint Venture Special
Publication.
Alisauskas, R. T., R. F. Rockwell, K. W. Dufour, E. G. Cooch, G. Zimmerman, K. L. Drake, J. O.
Leafloor, T. J. Moser, and E. T. Reed. 2011. Harvest, Survival, and Abundance of
Midcontinent Lesser Snow Geese Relative to Population Reduction Efforts. Wildlife
Monographs 179. Bethesda, Maryland: The Wildlilfe Society.
American Ornithologists' Union. 1983. Check-list of North American Birds. Edited by 7th.
Washington, D.C.: American Ornithologists' Union.
Argos, Inc. 1996. User's manual. Argos, Inc.,. Landover, Md.
Armstrong, T. W., Katherine M. Meeres, R. H. Kerbes, S. W. Boyd, J. G. Silveria, J. P. Taylor, and
B. Turner. 1999. "Routes and timing of migration of Lesser Snow Geese from the
Western Canadian Arctic and Wrangel Island, Russia, 1987-1992." In Distribution,
survival, and numbers of Lesser SNow Geese of the Western Canadian Arctic and
Wrangel Island, Russia. Occasional Paper Number 98 Canadian Wildlife Service, 75-84.
Ottawa, Ontario: Canadian Wildlife Service.
Aubry, L. M., R. F. Rockwell, E. G. Cooch, R. W. Brooks, C. P.H. Mulder, and D. N. Koons. 2013.
"Climate change, phenolgy, and habitat degradation: drivers of gosling body condition
and juvenile survival in lesser snow geese." Global Change Biology (19): 149-160.
Avis, J. C., R. T. Alisauskas, W. S. Nelson, and C. D. Ankney. 1992. "Matriarchal Population
Genetic Structure in and Avian Species with Female Natal Philopatry." Evolution 10841096.
Baldassarre, G. 2014. Ducks, Geese, and Swans of North America. Vol. 1. Baltimore, Maryland:
John Hopkins University Press.
Baranyuk, V. V. 1999. Breeding of the Lesser Snow Goose in limited nesting areas. Vol. 5, by E. E.
Syroechkovski, 161-176. Moscow: Russian Acadamy of Sciences.
—. 2005. "Warming of artic and Wrangel Island snow goose population [anstract]." 11th North
American Arctic Goose Conference and Workshop. Reno. 5-8.
Baranyuk, V. V., and E. V. Syroechkovsky. 1994. "The usage of the natural marks in Wrangel
Island Snow Geese for the population studies." Ornithologica 26: 45-49 (in Russian).



89

Baranyuk, V. V., and J. Y. Takekawa. 1998. "Migration of the Snow Geese in Chukotka and St.
Lawrence Island." Cascara 4: 169-187.
Baranyuk, V. V., J. E. Hines, and E V. Syroechkovsky. 1999. "Mineral staining of facial plumage as
an indicator of the winter ground affinities of Wrangel Island Lesser Snow Geese." In
Distribution, survival, and numbers of Lesser SNow Geese of the Western Canadian Artic
and Wrangel ISland, Russia. Occasional Paper Number 98 Canadian Wildlife Service,
edited by Richard H. Kerbes, Katherine M. Meeres and James E. Hines, 111-114. Ottawa,
Ontario: Canadian Wildlife Service Enviroment Canada.
Barlow, A. 2016. Pacific Flyway Council Migratory Bird Management. 11 11.
http://www.pacificflyway.gov/About.asp.
—. 2016. Pacific Flyway Council Migratory Bird Management. 11 11.
http://www.pacificflyway.gov/Management.asp.
Bayanyuk, V. V. 1992. "Wrangel Island Lesser Snow Geese Population Status, numbers,
Structure, and Problems of Protection [Abstract]." 7th North American Artic Goose
Conference and Workshop. Vallejo, CA. 23.
Bechet, A., A. Reed, N. Plante, J.F. Giroux, and G. Gauthier. 2004. "Estimating the size of the
Greater Snow Goose population." Journal of Wildlife Management 68 (3): 639-649.
Bellrose, F. C. 1980. Ducks, Geese and Swans of North America. 3. Washington, D.C.: Wildlife
Management Institute.
Blokpoel, H. 1974. Migration of LEsser Snow and Blue Geese in Spring across Southern Manitoba,
Part1: Distribution, Chronology, Directions, NUmbers, Heights, and Speeds. Report Series
28. Ottawa, Ontario: Canadian Wildlife Service.
Blokpoel, H., and M. C. Gauthier. 1975. Migration of Lesser Snow and Blue Geese in SPring across
Southern Manitoba, Part 2: Influence of Weather and Prediction of Major FLights. Report
Series 32. Ottawa, Ontario: Canadian Wildlife Service.
Bohrer, Gil, D. C. Douglas, R. Weinzierl, S. C. Davidson, R. Kays, M. Wikelski, and G. Bohrer. 2012.
"Moderating Argos location errors in animal tracking data." Methods in Ecology and
Evolution 3: 999-1007.
Bousfield, M. A., and E. V. Syroechkovsky. 1985. "A Review of Soviet research on the Lesser
Snow Goose on Wrangel Island, USSR." Waterfowl 36: 13-20.
Boyd, S. W. 1995. "Lesser Snow Geese (Anser c. caerulescens) and American three-aquare
bulrush (Scripus americanus) on the Frser and Skagit RIver Deltas." Ph.D. Thesis,
Department of Biological Sciences, Simon Fraser University, North Burnaby, B.C.,
Canada.



90

Boyd, W S, and F. Cooke. 2000. "Changes in the wintering distribution of Wrangel ISland SNow
Geese." Wildfowl 51: 59-66.
Britten, M. W., P. L. Kennedy, and S. Ambrose. 1999. "Performance and accuracy evaluation of
small satellite transmitters." Journal of Wildlife Management (4): 1349-1358.
Clements, J. F., T. S. Schulenberg, M. J. LLiff, D. Roberson, T. A. Fredericks, B. L. Sullivan, and C. L.
Wood. 2016. The ebird/Clements checklist of birds of the world: v2016.
http://www.birds.cornell.edu/clementschecklist.
Cooke, F., R. F. Rockwell, and B. D. Lank. 1995. The Snow Geese of La Perouse Bay: Natural
Selection in the Wild. Oxford: Oxford University Press.
Cooper, J. A. 1978. "The history and breeding biology of the Canada geese of Marshy Point,
Manitoba." Wildlife Monographs (51): 1-87.
Denny, M. 2016. Long Hops: Making Sense of Bird Migrations. Honolulu: University of Hawaii
Press.
Ely, C. R., J. Y. Takekawa, and M. L. Wege. 1993. "Distribution, abundance and age ratios of
Wrangel Island Lesser Snow Geese Anser caerulescens during autumn migration on the
Yukon-Kuskokwin Delta, Alaska." Wildfowl 44: 24-32.
Flicker, E. L. 1981. "Weather Conditions Associated with Beginning of Northward Migration
Departures of Snow Geese." The Journal of Wildlife Management 45 (2): 516-520.
Hatch, A. S., P. M. Meyers, D. M. Mulcahy, and D. C. Douglas. 2000. "Performance of Implantable
Satellite Transmitters in Diving Birds." Waterbird 23 (1): 84-94.
Hays, G. C., S. Akesson, B. J. Godley, P. Luschi, and P. Santidrian. 2001. "The impliations of
location accuracy for the interpretation of satellite-tracking data." Animal Behavior (61):
1035-1040.
Hines, J. E., M. O. Wiebe, S. J. Barry, V. V. Baranyuk, J. P. Taylor, R. McKelvey, S. R. Johnson, and
R. Kerbes. 1999. Survival rates of Lesser Snow Geese in the Pacific and Western Central
flyways, 1953-1989. Vol. 98, in Distribution, survival, and numbers of Lesser Snow Geese
of the Western Canadian Arctic and Wrangel Island, Russia, edited by Richard H. Kerbes,
Katherine M. Meeres and James E. Hines, 89-109. Canadian Wildlife Service.
Hines, J. E., V. V. Baranyuk, B. Turner, S. W. Boyd, J. G. Silveria, J. P. Taylor, S. J. Barry, K. M.
Meeres, R. H. Kerbes, and W. T. Armstrong. 1999. "Autumn and winter distributions of
Lesser Snow Geese from the Western Canadian Arctic and Wrangel Island, Russia, 19531992." In Distribution, survival, and numbers of Lesser Snow Geese of the Western
Canadian Arctic and Wrangel Island, Russia: Occasional Paper Number 98, edited by



91

Richard H. Kerbes, Katherine M. Meeres and James E. Hines, 39-74. Ottawa, Ontario:
Canadian Wildlife Service.
Hupp, J. W., A. B. Zacheis, R. M. Anthony, D. G. Robertson, W. P. Erickson, and K. C. Palacios.
2001. "Snow Cover and snow goose Anser caerulescens caerulescens distribution during
spring migration." Wildlife Biology 7: 65-76.
Johnson, M. A., and C. D. Ankney, . 2003. Direct Control and Alternative Harvest Strategies for
North American Light Geese: Report of the Direct COntrol and Alternative Harvest
Measures Working Group. Acrtic Goose Joint Venture Special Publication. U.S. Fish and
Wildlife Service, Washington, D.C., and Canadian Wildlife Service, Ottawa, Ontario.
Kennard, F. H. 1972. "The specific status of the greater snow goose." Proc. New England Zool.
Club 9: 85-93.
Kennard, F. H. 1927. "The specific status of the greater snow goose." Proc. New England Zool.
Club 9: 85-93.
Kerbes, R. H., and K. M. Meeres. 1999. "Project Overview." In Distribution, survival, and numbers
of Lesser Snow Geese of the Western Canadian Arctic and Wrangel Island, Russia.
Occasional Paper Number 98 Canadian Wildlife Service, edited by Richard H. Kerbes,
Katherine M. Meeres and James E. Hines, 15-24. Ottawa, Ontario: Canadian Wildlife
Service Environment Canada.
Kerbes, R. H., V. V. Baranyuk, and J. E. Hines. 1999. "Estimated size of the western Canadian
Arctic and Wrangel Island Lesser Snow Goose populations on their breeding and
wintering grounds." In Distribution, Survival, and Numbers of Lesser Snow Geese of the
Western Canadian Arctic and Wrangel Island, Russia. Occasional Paper 98, edited by
Richard H. Kerbes, Katherine M. Meeres and James E. Hines, 25-38. Ottawa, Ontario:
Canadian Wildlife Service.
Kerbes, R. M., K. E. Meeres, and J. E. Hines, . 1999. Distribution, survival, and numbers of Lesser
Snow Geese of the Western Canadian Artic and Wrangel Island, Russia: Occasional Paper
Number 98 Canadian Wildlife Service. Ottawa, Ontario: Canadian Wildlife Service
Enviornment Canada.
Konrad, P. M. 1993. "Satellite ornithology: researchers track snow geese from Russia to
California by satellite radio telemetry - - an interview with John Takekawa, Ph.D."
Wildlife 7 (4): 32-33.
Kraege, D. K., S. Boyd, and V. V. Baranyuk. 2008. "Monitoring and management of Wrangel
Island Lesser SNow Geese in Washington State, USA, and British Columbia, Canada."
Casarca 11 (1): 70-83.



92

Kranstauber, B., A. Cameron, R. Weinzerl, T. Fountain, S. Tilak, M. Wikelski, and R. Kays. 2011.
"The Movebank data model for animal traking." Environmental Modelling & Software 26
(6): 834-835.
Lincoln, F. C. 1979. The migration of birds. U.S. Fish and Wildlife Service Circular 16: Washington,
D.C., USA, 126.
Lok, Erika K., Daniel Esler, John Y. Takekawa, Susan W. De La Cruz, W. Sean Boyd, David R.
Nysewander, Joseph R. Evenson, and David H. Ward. 2011. "Stopover Habitats of Spring
Migrating Surf Scoters in Southeast Alaska." Jounal of Wildlife Management 75 (1): 92100.
Mehlmin, L. M., and C. W. Shaiffer. 1980. "Net-firing gun for capturing breeding waterfowl."
Journal of Wildlife Management 44: 895-896.
Miller, M. R., J. Y. Takekawa, J. P. Fleskes, D. L. Orthmeyer, M. L. Cassazza, and W. M. Perry.
2005. "Spring migration of Northern Pintails from California's Central Valley wintering
area tracked with satellite telemetry: routes, timing, and destinations." Canadian Jounal
of Zoology 83 (10): 1314-1332.
Ogilvie, Malcolm Alaxander, and Steve Young. 2002. Wildfowl of the World. London: New
Holland.
Ottenburghs, Jente, Hendrik-Jan Megens, Robert H.S. Kraus, Ole Madsen, Pim van Hooft, Sipke
E. van Wieren, Richard P.M.A. Crooijmans, Ronald C. Ydenberg, Martien A.M. Groenen,
and Herbert H.T. Prins. 2016. "A tree of geese: A phylogenomic perspective on the
evolutionary history of True Geese." Molecular Phylogenetics and Evolution 303-313.
Pacific Flyway Council. 2016. Pacific Flyway Council Recommendations, Informatoinal Notes and
Subcommittee Reports. Pacific Flyway Council.
Pacific Flyway Council. 2006. "Pacific Flyway management plan for the Wrangel Island
population of lesser snow geese." White Goose Subcommittee., Pacific Flyway Study
Commitee, Portland, OR.
Pauly, Daniel. 1995. "Anecdotes and teh shifting baseline syndrome of fisheries." Trends in
ecology and evolution 10 (10): 430.
Quinn, Tim W. 1992. "The genetic legacy of Mother Goose - phylogeographic patterns of lesser
snow goose Chen caerulescens caerulescens maternal lineages." Molecular Ecology 105117.
Raveling, D. G. 1979. "The annual energy cycle of the cackling Canada goose." In Management
and biology of Pacific Flyway geese, by R. L. Jarvis and J. C. Bartonek, 81-93. Corvalis,
Oregon: Oregon State University Bookstores.



93

Reed, E. T., G. Gauthier, and J. F. Giroux. 2004. "Effects of spring conditions on breeding
propensity of greater snow goose females." Animal Biodiversity and Conservation 27 (1):
35-46.
Ryder, J. P. 1967. "The breeding biology of the Ross' goose in tthe Perry River region, Northwest
Territories. Canadian Wildlife Service Report Series 3." Ottawa, Ontario, Canada, 56.
Schmutz, J. A., K. A. Hobson, and J. A. Morse. 2006. "An isotopic assessment of protein from diet
and endogenous stores: effects on egg production and incubation behavior of geese."
ARDEA-WAGENINGEN 94 (3): 385.
Sileo, Louis, C. E. Korschgen, P. K. Kenow, L. W. Green, M. D. Samual, and L. Sileo. 1996.
"Technique for Implanting Radio transmitters Subcutaneously in Day Old Ducklings."
Journal of Field Ornithology 67 (3): 392-397.
Syroechkovsky, E. V., and K. E. Litvin. 1986. "Investigation of the migration of the Snow Geese of
Wrangel Island by the method of individual marking." In The ringing and marking of
birds in the USSR, by V. E. Sokolov and I. N. Dobrinina, translated by M.A. Bousfield, 520. Moscow (In Russian): Nauka.
Takekawa, J. Y., D. L. Orthmeyer, M. Kurechi, Y. Sabano, E. V. Syroechkovsky, K. E. Litvin, V. V.
Baranyuk, and A. V. Andreev. 1994. "Restoration of Lesser Snow Geese to East Asia: a
North Pacific Rim conservation project." Transactions of the North American Wildlife
and Natural Resource Conference. 132=145.
U.S. Fish and Wildlife Service. 2016. Waterfowl populations status, 2016. Washington, D.C.: U.S.
Department of the Interior.
Warbock, N., and M. Bishop. 1998. "Spring stopover ecology of migrant western sanpipers." The
Condor (100): 456-467.
Washington Department of Fish and Wildlife . 2016. 2016 Game Status and Trend Report. .
Wildlife Program, Olympia, Washington, USA: Washington Department of Fish and
Wildlife.
Wege, M. L., and D. G. Raveling. 1983. "Factors influencing the timing, distance, and paths of
migrations of Canada geese." Wilson Bulletin. 2009-221.
Willams, C. K., M. D. Samuel, V. V. Baranyuk, E. G. Cooch, and D. Kraege. 2008. "Winter Fidelity
and Apparent Survival of Lesser SNow Goose Populations inthe Pacific Flyway." The
Journal of Wildlife Management 72 (1): 159-167.




94

Media

Stevick_PFMESthesis2017.pdf