External Review Draft | EPA910-R-12-004b | May 2012
United States
Environmental Protection
Agency
An Assessment of Potential Mining Impacts
on Salmon Ecosystems of Bristol Bay, Alaska
Volume 2 - Appendices A-D
U.S. Environmental Protection Agency, Seattle, WA
www.epa.gov/bristolbay
External Review Draft - Do Not Cite or Quote
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DRAFT EPA910-R-12-004b
DO NOT CITE OR QUOTE May 2012
External Review Draft
www.epa.gov/bristolbay
An Assessment of Potential Mining Impacts on
Salmon Ecosystems of Bristol Bay, Alaska
Volume 2 - Appendices A-D
NOTICE
THIS DOCUMENT IS AN EXTERNAL REVIEW DRAFT. It has not been formally released by
the U.S. Environmental Protection Agency and should not be construed to represent Agency
policy. It is being circulated for comment on its technical accuracy and policy implications.
U.S. Environmental Protection Agency
Seattle, WA
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DISCLAIMER
This document is distributed solely for the purpose of pre-dissemination peer review under
applicable information quality guidelines. It has not been formally disseminated by the U.S.
Environmental Protection Agency (USEPA). It does not represent and should not be
construed to represent any Agency determination or policy. Mention of trade names or
commercial products does not constitute endorsement or recommendation for use.
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CONTENTS
VOLUME 1
An Assessment of Potential Mining Impacts on Salmon Ecosystems of Bristol Bay, Alaska
VOLUME 2
APPENDIX A: Fishery Resources of the Bristol Bay Region
APPENDIX B: Characterizations of Selected Non-Salmon Fishes Harvested in the Fresh
Waters of Bristol Bay
APPENDIX C: Wildlife Resources of the Nushagak and Kvichak River Watersheds
APPENDIX D: Ecological Knowledge and Cultures of the Nushagak and Kvichak Watersheds,
Alaska
VOLUME 3
APPENDIX E: Bristol Bay Wild Salmon Ecosystem Baseline Levels of Economic Activity and
Values
APPENDIX F: Biological Characterization: Bristol Bay Marine Estuarine Processes, Fish, and
Marine Mammal Assemblages
APPENDIX G: Foreseeable Environmental Impact of Potential Road and Pipeline
Development on Water Quality and Freshwater Fishery Resources of Bristol Bay, Alaska
APPENDIX H: Geologic and Environmental Characteristics of Porphyry Copper Deposits
with Emphasis on Potential Future Development in the Bristol Bay Watershed, Alaska
APPENDIX I: Conventional Water Quality Mitigation Practices for Mine Design,
Construction, Operation, and Closure
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Appendix A
Fishery Resources of the Bristol Bay Region
A-l
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Fishery Resources of the Bristol Bay Region
Daniel Rinella, PhD.
Alaska Natural Heritage Program
University of Alaska Anchorage
Beatrice McDonald Hall, Suite 106
Anchorage, AK 99508
rinella@uaa.alaska.edu
907.786.4963
Rebecca Shaftel
Alaska Natural Heritage Program
University of Alaska Anchorage
Beatrice McDonald Hall, Suite 106
Anchorage, AK 99508
rsshaftel@uaa.alaska.edu
907.786.4965
Dave Athens
Environmental Protection Agency
Kenai River Center
514 Funny River Road
Soldotna, AK 99669
Athons.dave@epamail.epa.gov
907.714.2481
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Table of Contents
INTRODUCTION 1
ECOLOGY AND LIFE HISTORY OF BRISTOL BAY FISHES 3
General salmon life history 3
Species-specific life history and ecology 4
Sockeye salmon 4
Chinook salmon 6
Rainbow trout 8
Coho salmon 8
Pink salmon 9
Chum salmon 10
BRISTOL BAY FISHERIES AND FISHERIES MANAGEMENT 10
Historical perspective on commercial salmon fisheries 10
Current management of commercial salmon fisheries 12
Description of sport fisheries 15
Management of sport fisheries 16
Chinook salmon 16
Sockeye salmon 17
Rainbow trout 17
SALMON ABUNDANCE TRENDS AROUND THE NORTH PACIFIC, WITH REFERENCE TO BRISTOL BAY
POPULATIONS 18
Sockeye salmon 18
Size of Bristol Bay, Kvichak, and Nushagak sockeye salmon returns 18
Factors affecting Bristol Bay sockeye salmon abundance 25
The decline in Kvichak River sockeye salmon runs 26
Chinook salmon 28
Threatened and endangered salmon and conservation priorities 32
KEY HABITAT ELEMENTS OF BRISTOL BAY RIVER SYSTEMS (OR WHY DO BRISTOL BAY WATERSHEDS
PRODUCE SO MANY FISH?) 37
Habitat quantity 37
Habitat quality 40
Habitat diversity 43
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Tables
Table 1. Mean harvest by species and fishing district, 1990-2009 12
Table 2. Bristol Bay escapement goal ranges for sockeye salmon 13
Table 3. Bristol Bay escapement goal ranges for Chinook and chum salmon 14
Table 4. The number of businesses and guides operating in the Nushagak and Kvichak watersheds in
2005, 2008 and 2010 16
Table 5. Mean annual returns of sockeye salmon in Bristol Bay, 1956-2010, and percent of total by river
system 27
Table 6. Chinook average run sizes for 2000-2009 for rivers across the North Pacific 29
Table 7. Endangered Species Act listings for salmon ESUs in the United States 35
Table 8. Comparison of landscape features potentially important to sockeye salmon production for
watersheds across the North Pacific and across the Bristol Bay watershed 38
Table 9. A summary of life history variation within the Bristol Bay stock complex of sockeye salmon.... 44
Table 10. Variation in time spent rearing in fresh water and at sea for Bristol Bay sockeye salmon 44
Figures
Figure 1. Major river systems and fishing districts in Bristol Bay, Alaska 2
Figure 2. Sockeye salmon distribution in the Nushagak and Kvichak watersheds 5
Figure 3. Chinook salmon distribution in the Nushagak and Kvichak watersheds 7
Figure 4. Relative abundance of wild sockeye salmon stocks in the North Pacific, 1956-2005 19
Figure 5. Wild sockeye salmon abundances by region in the North Pacific, 1956-2005 20
Figure 6. Total sockeye returns by river system in Bristol Bay, 1956-2010 22
Figure 7. Sockeye salmon abundances for major rivers of the North Pacific, 1956-2010 24
Figure 8. Chinook salmon abundances by river system, 1966-2010 31
Figure 9. Map of surveyed anadromous streams in the Nushagak and Kvichak watersheds 39
Appendices
Appendix 1. Chinook and sockeye almon run sizes for Bristol Bay and other regions of the North Pacific.
56
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INTRODUCTION
Beginning each June, millions of Pacific salmon return from feeding in the open ocean
and swarm through Bristol Bay en route to their natal spawning streams. Nine major river
systems comprise the spawning grounds for Bristol Bay salmon (Figure 1), and schools navigate
toward the mouths of their respective rivers as they pass through the Bay. Each summer,
thousands of commercial fishermen use drift and set gill nets to capture millions of returning
fish, making Bristol Bay the largest sockeye salmon fishery in the world. Salmon that escape the
fishery distribute throughout the Bay's watersheds and spawn in hundreds of discreet
populations. Sport anglers target those salmon, especially sockeye, Chinook and coho, as they
migrate through the river systems toward their spawning grounds. Also prized are abundant
populations of rainbow trout and other sport fish, including Dolly Varden and Arctic grayling,
which attain trophy size by gorging on energy-rich salmon eggs, flesh from salmon carcasses,
and invertebrates dislodged by spawning salmon. The abundance of large game fish, along with
the wilderness setting, makes the Bristol Bay region a world-class destination for sport anglers.
Alongside recreationists, aboriginal people, guided by an age-old culture, harvest their share of
migrating salmon and other fish species, which provide a primary source of sustenance.
In this report we reviewed the biology, ecology, and management of the fishes of the
Bristol Bay watersheds, emphasizing those species of the greatest cultural and economic
importance - sockeye salmon, Chinook salmon, and rainbow trout. Rather than to imply that
other fishes are not important, this focus reflected the disproportionate amount of research on
these species (especially sockeye salmon) and was necessary to keep the amount of material
manageable. In contrast, there is relatively little information available for the region's
freshwater species, despite the importance of some in subsistence and sport fisheries. Our
objectives were to describe the commercial and sport fishery resources of the region and to
discuss the importance of Bristol Bay salmon populations in the context of the greater North
Pacific Ocean. The literature reviewed consisted primarily of agency reports and peer-reviewed
scientific papers, although unpublished data and personal communications were used where no
pertinent published literature existed and popular sources were consulted to characterize the
more subjective attributes of the sport fisheries. Our geographic focus was the Kvichak River
watershed (including the Alagnak River) and the Nushagak River watershed (including the
Wood River). Since the Kvichak and Nushagak sockeye salmon populations are components of
the Bristol Bay-wide stock complex, however, we typically discuss their abundance trends at
both the Bristol Bay scale and at the scale of the individual river systems. The economics of
Bristol Bay's fisheries and the role offish in the region's aboriginal cultures are each covered in
separate sections of the Bristol Bay Watershed Analysis.
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Bristol Bay Fishing Districts
Bristol Bay Watershed
Major Lakes
Major Rivers
Figure 1. Major river systems and fishing districts in Bristol Bay, Alaska.
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ECOLOGY AND LIFE HISTORY OF BRISTOL BAY FISHES
General salmon life history
Five species of Pacific salmon are native to North American waters - pink (Oncorhynchus
gorbuscha), chum (0. keta), sockeye (0. nerka), coho (0. kisutch), and Chinook (0. tshawytscha)
salmon - and all have spawning populations in the Bristol Bay region. These species share a
rare combination of life history traits that contribute to their biological success, as well as their
status as cultural icons around the North Pacific rim. These traits - anadromy, homing, and
semelparity - are described briefly in the following paragraphs.
All Pacific salmon hatch in fresh water, migrate to sea for a period of relatively rapid
growth, and return to fresh water to spawn. This strategy, termed anadromy, allows salmon to
capitalize on the resource-rich marine environment, where growth rates are much faster than
in fresh water. Thus, anadromy allows salmon to attain larger body size, mature more quickly,
and maintain larger spawning populations than would be possible with a non-migratory life
history (McDowall 2001). A prevailing theory is that anadromy evolves where a disparity in
productivity exists between adjacent freshwater and marine environments (Gross et al. 1988).
Freshwater productivity generally declines with latitude, and in the spawning range of Pacific
salmon is half (or less) of that in lower latitudes. Conversely, ocean productivity generally
increases with latitude, peaking within the range of Pacific salmon (Gross et al. 1988).
When salmon enter fresh water to spawn, the vast majority return to the location
where they were spawned. By this means, termed homing, salmon increase juvenile survival by
returning to spawn in an environment with proven suitability (Cury 1994). Another adaptive
advantage of homing is that it fosters reproductive isolation that enables populations to adapt
to their particular environment (Blair et al. 1993, Dittman and Quinn 1996, Eliason et al. 2011).
For instance, populations that travel long distances to reach inland spawning sites develop large
lipid reserves to fuel the migration (Quinn 2005, pgs. 77-78 and figures 4-6), since adult salmon
generally do not feed after entering fresh water. As another example, sockeye fry from
populations that spawn downstream of nursery lakes are genetically programmed to migrate
upstream after emergence, while fry from populations that spawn upstream of nursery lakes
are programmed to migrate downstream (Burgner 1991, pgs. 33-35). Examples of adaptations
are many, and include heritable anatomical, physiological, and behavioral traits. Without
homing, gene flow would occur throughout the species, making adaptation to specific
freshwater conditions impossible; in this sense, homing counteracts the dispersal effects of
anadromy (McDowall 2001). Homing is not absolute, however, and a small amount of straying
ensures that amenable habitats are colonized by salmon (e.g., Milner and Bailey 1989).
Pacific salmon, quite famously, die after spawning only once. This trait, termed
semelparity, serves to maximize the investment in one reproductive effort at the expense of
any future reproductive effort. In salmon, it may have evolved as a response to the high cost of
migration to natal streams and the associated reduction in adult survival (Roff 1988). The
evolution of semelparity in Pacific salmon was accompanied by increased egg size so, while long
migrations may have been a prerequisite, the driving force behind the evolution of semelparity
was likely the increase in egg mass and associated increase in juvenile survival (Crespi and Teo
2002).
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As salmon approach sexual maturity, the countershading and silvery sheen that hide
them at sea give way to characteristic spawning colors, often with hues of red. Males develop
hooked snouts (the generic name Oncorhynchus refers to this trait) and protruding teeth, and
their previously bullet-shaped bodies become laterally flattened. These spawning colors and
secondary sexual characteristics, which develop to varying degrees among species and even
among populations, probably serve multiple purposes on the spawning grounds, including
species recognition, sex recognition, and territorial displays.
With few exceptions, preferred spawning habitat consists of gravel-bedded stream
reaches with moderate depth and current (30-60 cm deep and 30-100 cm per second,
respectively; Quinn 2005, pg. 108). Females excavate a nest (redd) in the gravel to receive the
eggs, which are fertilized by one or more competing males as they are released and
subsequently buried by the female. The seasonality of spawning and incubation is roughly the
same for all species of Pacific salmon, although the timing can vary somewhat by species,
population, and region. In general, salmon spawn during summer or early fall and the fry
emerge from the spawning gravel the following spring. While in the gravel, the embryos
develop within their eggs and then hatch into fry that continue to subsist on yolk sacs. After
emerging from the gravel, basic life history patterns of the five species differ in notable ways.
Species-specific life history and ecology
Sockeye salmon
Sockeye salmon originate from river systems along the North American and Asian shores
of the North Pacific and Bering Sea, roughly from the latitude of the Sacramento River to that of
Kotzebue Sound. The largest North American populations occur between the Columbia and
Kuskokwim rivers (Burgner 1991, pg. 5). Spawning sockeye are readily identified by their
striking red bodies with green heads and tails; males additionally develop a large hump in front
of the dorsal fin.
Sockeye are unique among salmon in that most stocks rely on lakes as the primary
freshwater rearing habitat. Some sockeye spawn within the nursery lake where their young will
rear. Others spawn in nearby stream reaches, and their fry migrate to the nursery lake after
emerging from spawning redds. Sockeye are by far the most abundant salmon species in the
Bristol Bay region (Salomone et al. 2011, pg. 1), undoubtedly due to the abundance of
accessible lakes in this landscape (Figure 1; also see discussion of habitat quantity). Tributaries
to Iliamna Lake, Lake Clark, and the Wood Tikchik Lakes are major spawning areas, and
juveniles rear in each of these lakes (Figure 2). On average, Iliamna Lake produces more
sockeye than any other lake in the Bristol Bay region (see data for Kvichak River in Appendix 1).
Riverine sockeye populations spawn and rear throughout the Nushagak River basin (Figure 2).
Juveniles in Bristol Bay systems rear for one or two years in their nursery lakes (West et al.
2009, pg. 235), feeding primarily on zooplankton in the limnetic zone (Burgner 1991, pg. 37).
Fish then typically spend two or three years at sea (West et al. 2009), returning at an
average weight of 5.9 Ib (2.7 kg, based on recent commercial catches; Salomone et al. 2011, pg.
105). At sea, sockeye salmon feed on a range of invertebrates, small fish, and squid (Burgner
1991, pg 83).
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Sockeye streams intneAWC
Spawning
Present or rearing
Figure 2. Sockeye salmon distribution in the Nushagak and Kvichak watersheds. Data are from the Alaska Department of Fish and
Game's Anadromous Waters Catalog for 2011.
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Chinook salmon
Chinook salmon spawn in streams on both shores of the North Pacific and Bering Sea,
roughly from the latitude of central California to that of Point Hope. There are more than a
thousand North American spawning populations and a much smaller number in Asia. These
populations tend to be relatively small, however, making Chinook the rarest of North America's
Pacific salmon species (Healey 1991, pg. 316). They are also the largest of the Pacific salmon; at
least one specimen over 60 kg has been reported, but most weigh less than 23 kg (Mecklenburg
etal.2002, pg. 207).
Chinook salmon have two different behavioral forms. The "stream type" form is
predominant in Bristol Bay, as well as other areas of northern North America, Asia, and the
headwaters of Pacific Northwest rivers (Healey 1991, pg. 314). These fish spend one or more
years as juveniles in fresh water, range widely at sea, and return to spawning streams during
spring or summer. "Ocean type" Chinook, by contrast, migrate to sea soon after hatching,
forage primarily in coastal marine waters, and return to spawning streams in the fall (Healey
1991, pg. 314). In fresh water, juvenile Chinook tend to occupy flowing water and feed on
aquatic insects. At sea, Chinook are generally pisciverous (Brodeur 1990) and feed higher on
the food chain than other salmon species (Satterfield and Finney 2002).
Chinook spawn and rear throughout the Nushagak River basin and in many tributaries of
the Kvichak River (Figure 3). Some life history data are available from adults returning to the
Nushagak River, Bristol Bay's largest Chinook salmon run. Essentially all Chinook spend one
year rearing in fresh water, and the vast majority (typically >90% of a given brood year) spend
two to four years at sea (Gregory Buck, ADF&G, unpublished data). Fish that spend four years
at sea are the dominant age class and comprise approximately 43% of the average return,
followed by those that spend 3 years (35%) and two years (17%) at sea. Chinook salmon
individuals in recent Bristol Bay commercial catches have averaged 16.6 Ib (7.5 kg; Salomone et
al. 2011, pg. 105).
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Chinook streams in the AWC
Spawning
Present or rearing
Figure 3. Chinook salmon distribution in the Nushagak and Kvichak watersheds. Data are from the Alaska Department of Fish and
Game's Anadromous Waters Catalog for 2011.
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Rainbow trout
Rainbow trout (Oncorhynchus mykiss) are native to western North America and the
eastern coast of Asia, although their popularity as a sport fish has led to introduced populations
around the world. Bristol Bay's rainbow trout are of the coastal variety (sensu Behnke 1992,
pg. 193), which ranges from the Kuskokwim River to southern California. While classified in the
same genus as the Pacific salmon, there are some key differences. Foremost, rainbow trout are
not genetically programmed to die after spawning, making iteroparity (i.e., repeat spawning) a
feature of most populations. Also, most coastal drainages support populations of both resident
and anadromous (i.e., steelhead) forms, although only the resident form occurs near the
northern and southern limits of rainbow trout distribution (Behnke 1992, pg. 197), including the
rivers of Bristol Bay. Finally, rainbow trout spawn in the spring, as opposed to summer or early
fall, although their spawning habitat and behavior is otherwise generally similar to that of
salmon.
Bristol Bay rainbow trout tend to mature slowly and grow to relatively large size. For
example, 90% of spawners in Lower Talarik Creek were more than seven years old; the vast
majority of these were longer than 500 mm and a few exceeded 800 mm (years 1971-1976;
Russell 1977, pgs. 30-31). Growth (mm/year) was fastest for fish between four and six years of
age and winter growth appeared to be minimal (Russell 1977, pgs. 44-45).
Bristol Bay trout utilize complex and varying migratory patterns that allow them to
capitalize on different stream and lake habitats for feeding, spawning, and wintering. Fish from
Lower Talarik Creek migrate downstream to Iliamna Lake after spawning. From there, they
appear to utilize a variety of habitats, as some tagged individuals have been recovered in other
Iliamna Lake tributaries and in the Newhalen and Kvichak Rivers (Russell 1977, pg. 23). In the
Alagnak River watershed, a number of rainbow trout life history types have been identified,
each with their own habitat use and seasonal migratory patterns (Meka et al. 2003). These
consist of lake, lake-river, and river residents, the latter of which range from non-migratory to
highly migratory (Meka et al. 2003). Individuals comprising each of these life history types
migrate in order to spend the summer in areas with abundant spawning salmon (Meka et al.
2003).
Eggs from spawning salmon are a major food item for Bristol Bay trout and are likely
responsible for much of the growth attained by these fish. Upon the arrival of spawning
salmon in the Wood River basin, rainbow trout shifted from consuming aquatic insects to
primarily salmon eggs for a 5-fold increase in ration and energy intake (Scheuerell et al. 2007).
With this rate of intake, a bioenergetics model predicts a 100-g trout to gain 83 g in 76 days;
without the salmon-derived subsidy, the same fish was predicted to lose five g (Scheuerell et al.
2007). Rainbow trout in Lower Talarik Creek were significantly fatter (i.e., higher condition
factor) in years with high spawner abundance than in years with low abundance (Russell 1977,
pg. 35).
Coho salmon
Coho salmon are native to coastal drainages in western North America and eastern Asia,
approximately from the latitude of the Sacramento River to that of Point Hope (Sandercock
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1991, pg. 398). Coho salmon occur in relatively small populations, and are second only to
Chinook salmon in rarity.
Most Alaskan coho salmon populations tend to spend two years in fresh water and one
year at sea (Sandercock 1991, pg. 405). Few age data exist for Bristol Bay, but samples from
two years on the Nushagak River indicated that approximately 90% of escaped coho salmon
shared this age structure, while the remaining fish had spent either one year or three years in
fresh water (West et al. 2009, pg. 84). Coho salmon individuals in recent Bristol Bay commercial
catches have averaged 6.7 Ib (3.0 kg; Salomone et al. 2011, pg. 105).
At sea, coho salmon consume a mix of fish and invertebrates (Brodeur 1990, pg. 15).
Their trophic position is intermediate for Pacific salmon; Chinook salmon consume more fish
while sockeye, pink, and chum salmon eat more zooplankton and squid (Satterfield and Finney
2002).
In fresh water, coho salmon feed primarily on aquatic insects, although salmon eggs and
flesh can be important nutritional subsidies (Heintz et al. 2010, Rinella et al. In press). They
utilize a wide range of lotic and lentic freshwater habitats, including stream channels, off-
channel sloughs and alcoves, beaver ponds, and lakes. Coho distribute widely into headwater
streams, where they are often the only salmon species present (Woody and O'Neal 2010, King
et al. 2012, ADF&G Anadromous Waters Catalog). Production of juvenile coho is often limited
by the extent and quality of available wintering habitats (Nickelson et al. 1992, Solazzi et al.
2000), and preliminary work in southcentral Alaska suggests that upwelling groundwater is an
important feature (D.J. Rinella, unpublished data).
Pink salmon
Pink salmon spawning populations occur on both sides of the North Pacific and Bering
Sea, as far south as the Sacramento River and northern Japan. Northward, small spawning
populations are scattered along the North American and Asian shores of the Arctic Ocean. The
most abundant Pacific salmon overall (Irvine et al. 2009, pg. 2), pink salmon have a simplified
life history that relies little on freshwater rearing habitat. They typically spawn in shallow,
rocky stream reaches relatively low in the watershed and their young migrate to sea soon after
emerging (Heard 1991, pg. 144).
Essentially all pink salmon breed at two years of age, and this strict two-year life cycle
results in genetic isolation of odd- and even-year spawning runs, even within the same river
system. For reasons not entirely clear, large disparities between odd- and even-year run sizes
occur across geographic regions and extend over many generations. An extreme example is the
Fraser River, in southern British Columbia, where millions of pink salmon return during odd-
numbered years, yet no fish return during even-numbered years (Riddell and Beamish 2003, pg.
4). In Bristol Bay rivers, even-year runs dominate the returns (Salomone et al. 2011, pg. 5).
Pink salmon are the smallest of the Pacific salmon species; individuals in recent Bristol
Bay commercial catches have averaged 3.6 Ib (1.6 kg; Salomone et al. 2011, pg. 105). Sexually
mature males become highly laterally compressed and develop a massive dorsal hump, hence
the common name "humpy."
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Chum salmon
Chum salmon spawn on both shores of the Bering Sea and North Pacific, extending
south to the latitude of Japan and California (Salo 1991, pg. 234). Scattered spawning
populations also occur on the Asian and North American shores of the Arctic Ocean.
Populations tend to be relatively large, and chum salmon are the third most abundant species,
behind pink and sockeye salmon.
Chum salmon, like pink salmon, migrate to sea soon after emerging from spawning
gravel. Across their range, the vast majority spends two to four years at sea (Salo 1991, pg.
272), and one year's run in the Nushagak River showed similar age structure (West et al. 2009,
pgs. 82-83). At sea, chum salmon consume a range of invertebrates and fishes, and gelatinous
material is commonly found in stomachs leading to speculation that jellyfish may be a common
prey item (Brodeur 1990, pg. 8, Azuma 1992). Individuals in recent Bristol Bay commercial
catches have averaged 6.8 Ib (3.1 kg, Salomone et al. 2011, pg. 105).
BRISTOL BAY FISHERIES AND FISHERIES MANAGEMENT
Historical perspective on commercial salmon fisheries
Salmon have long been an important economic driver in Alaska's economy and have
played an important role in the state's history. Commercial fishing interests were among the
original supporters of the purchase of Alaska from Russia in 1867 (King 2009, pg. 1). The first
canneries were established eleven years later, and by the 1920s salmon surpassed mining as
Alaska's major industry as Alaska became the world's principal salmon producer (Ringsmuth
2005, pg. 21).
In the early years, fish packing companies essentially had a monopoly on the harvest of
salmon. Packers in Bristol Bay and elsewhere built industrial fish traps, constructed of wood
pilings and wire fencing with long arms that guided schools of migrating salmon into holding
pens (King 2009, pg. 4). In Bristol Bay, packing interests also upheld a federal ban on fishing
with power boats until 1951. Ostensibly a conservation measure, this law served to protect
obsolete cannery-owned sailboat fleets by excluding independent Alaska-based fishermen who
largely used power boats by this time (Troll 2011, pg. 39).
Salmon harvest peaked in 1936 then declined steadily for many years, leading to a
federal disaster declaration in the 1950s (King 2009, pg. 1). A lack of scientific management,
poor federal oversight, excessive harvest during World War II, and natural changes in ocean
conditions contributed to the decline.
Declining salmon runs, along with Alaskans' desire for more control over their fisheries,
was a significant factor in the drive toward Statehood (Augerot 2005, King 2009, pg. 2). In
1955, Alaskans began to develop a state constitution that included provisions intended to
preserve Alaska's fisheries and, unique among state constitutions, to guarantee equal access to
fish and game for all residents. Alaska became a state in 1959, the year that marked the lowest
salmon harvest since 1900 (King 2009, pg. 3). Statehood was a turning point for Alaska's
salmon fisheries, with the end of federal management, fish traps, and undue control of the
resource by the canning industry. With the mandate for equal access came decentralization of
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the fishing industry, and thousands of individual fishermen began harvesting salmon for market
to the canneries (Ringsmuth 2005, pg. 65).
When the Alaska Department of Fish and Game (ADF&G) assumed management of the
fisheries in 1960, restoring salmon runs to their former abundance became a primary objective.
Inventorying fish stocks, understanding basic ecology, and improving run strength forecasting
were central research goals. Of particular importance was the development and application of
methods for counting salmon runs in spawning streams, which allowed the establishment of
escapement goals and management based on scientific principles of sustained yield. Bristol Bay
salmon research has been conducted primarily by ADF&G staff and researchers at the
University of Washington's Alaska Salmon Program (see
http://fish.washington.edu/research/alaska/). The latter, funded largely by the salmon
processing industry, began researching factors controlling sockeye salmon production in 1947.
While the scope of their investigations has expanded over the years, sockeye monitoring is still
a focus and represents the world's longest-running program for monitoring salmon and their
habitats.
Over time, a number of state and federal policy changes have affected Bristol Bay
salmon fisheries. A 1972 constitutional amendment set the stage for a bill that limited
participation in Alaska commercial salmon fisheries. This legislation, designed to curb the
expanding commercial fishery, set an optimum number of permits for each fishery, which were
then issued by the State based on an individual's fishing history. Permits are owned by the
individual fisherman and are transferable, making them a limited and valuable asset (King 2009,
pg. 22). The Fishery Conservation and Management Act of 1976, commonly known as the
Magnuson-Stevens Act, was introduced to Congress by the late senator Ted Stevens as a means
to curtail high seas salmon fishing. In response to intensive Japanese gill netting in the western
Aleutians and Bering Sea since 1952, this legislation extended America's jurisdiction from 12 to
200 miles (19 to 322 km) offshore. This ensured that salmon produced in Alaskan rivers would
be harvested and processed locally and gave Alaska's fishery managers much more control in
deciding when and where salmon are harvested. Both the Policy for the Management of
Sustainable Salmon Fisheries and the Policy for Statewide Salmon Escapement Goals were
adopted in the winter of 2000-2001 (Baker et al. 2009, pg. 2). The former established a
comprehensive policy for the regulation and management of sustainable fisheries and the latter
defined procedures for establishing and updating salmon escapement, including a process for
public review of allocation disputes associated with escapement goals
The Alaska Department of Fish and Game is responsible for managing fisheries under
the sustained yield principle. Fishing regulations, policies, and management plans are enacted
by the Board of Fisheries, which it does in consultation with ADF&G, advisory committees, the
public, and other state agencies. The Board of Fisheries consists of seven citizens, appointed by
the governor and confirmed by the legislature, that serve three-year terms. Eighty-one
advisory committees, whose members are elected in local communities around the state,
provide local input. While regulations and management plans provide the framework for
fisheries regulation, local fisheries managers are ultimately responsible for their execution.
They are delegated authority to make "emergency orders," in-season changes to fishing
regulations, which allow rapid adjustments to changing conditions, often with very short notice.
Managers use them to provide additional protection to fish stocks when conservation concerns
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arise and to liberalize harvest when surplus fish are available. Management plans directed at
specific fish stocks are often based on anticipated scenarios and give specific directions to
managers, making the in-season management process predictable to ADF&G, commercial
fishermen, and the public. Alaska's management of its salmon fishery has proven successful; it
was the second fishery in the world to be certified as well managed by the Marine Stewardship
Council (Hilborn 2006) and is regarded as a model of sustainability (Hilborn et al. 2003a, King
2009).
Current management of commercial salmon fisheries
While all five species of Pacific Salmon are harvested in Bristol Bay, sockeye salmon
dominate the runs and harvest by a huge margin (Table 1). Salmon return predominately to
nine major river systems, located on the eastern and northern sides of the Bay, and are
harvested in five fishing districts in close proximity to the river mouths that allow managers to
regulate harvest individually for the various river systems (Figure 1). The Naknek-Kvichak
district includes those two rivers as well as the Alagnak. The Nushagak district includes the
Nushagak, Wood, and Igushik Rivers. The Egegik, Ugashik, and Togiak districts include the rivers
for which they are named.
Table 1. Mean harvest by species and fishing district, 1990-2009. Unpublished data, Paul
Salomone, ADF&G Area Management Biologist.
Naknek-
Egegik Ugashik Nushagak Togiak Total
Kvichak
Sockeye
Chinook
Chum
Pink*
Coho
8,238,895
2,816
184,399
73,661
4,436
8,835,094
849
78,183
1,489
27,433
2,664,738
1,402
70,240
138
10,425
5,478,820
52,624
493,574
50,448
27,754
514,970
8,803
158,879
43,446
14,234
25,732,517
66,494
985,275
169,182
84,282
*Pink salmon data are from even-numbered years only since harvest is negligible during the
smaller odd-year runs.
Fishing is conducted with drift or set gillnets. Set gillnets have a maximum length of 150
fathoms (274 m) and are fished from boats no longer than of 32 ft. (9.8 m) in length. Set
gillnets are fished from beaches, often with the aid of an open skiff, and have a maximum
length of 50 fathoms (91 m). There are approximately 1900 drift gillnet permits and 1000 set
gillnet permits in the Bristol Bay salmon fishery, of which around 90% are fished on a given year
(1990-2010 average; Salomone et al. 2011, pg. 84).
The management of the Bristol Bay sockeye salmon fishery is focused on allowing an
adequate number of spawners to reach each river system while maximizing harvest in the
commercial fishery (Salomone et al. 2011, pg. 2). This balancing act is achieved through the
establishment of escapement goals which represent the optimum range of spawners for a given
river system. Escapement goals are established using a time series of spawner counts where a
spawning run of a given size (i.e., stock) can be linked to the number of its offspring returning in
subsequent years (i.e., recruits). Established stock-recruit models (Ricker 1954, Beverton and
12
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Holt 1957) are then used to estimate the stock size that results in the largest number of
recruits, or the maximum sustained yield (Baker et al. 2009, pg. 4). In theory, spawning runs
that are too small or large can result in reduced recruitment. With the former, too few eggs are
deposited. With the latter, superimposition of spawning redds can diminish egg viability and
competition in nursery lakes can reduce growth and survival. Once escapement goals are set,
the timing and duration of commercial fishery openings are then adjusted during the fishing
season (i.e., in-season management) to ensure that escapement goals are met and any
additional fish are harvested. Escapement goals are periodically reviewed and updated based
on regulatory policies, specifically, the Policy for the Management of Sustainable Salmon
Fisheries and the Policy for Statewide Salmon Escapement Goals.
Each of Bristol Bay's nine major river systems has an escapement goal for sockeye
salmon (Table 2), and in-season management of the commercial fishery is used to keep
escapement in line with the goals. Management responsibility is divided among three
managers: one for the Naknek, Kvichak, and Alagnak rivers; one for the Nushagak, Wood,
Igushik, and Togiak rivers; and one for the Ugashik and Egegik rivers. Fishery openings are
based on information from a number of sources, including preseason forecasts, the test fishery
at Port Moller, the early performance of the commercial fishery, and in-river escapement
monitoring.
Table 2. Bristol Bay escapement goal ranges for sockeye salmon.
River
Kvichak
Alagnak
Naknek
Egegik
Ugashik
Wood River
Igushik
Nushagak-Mulchatna
Togiak
Escapement range
(thousands)
2,000-10,000
320 minimum
800-1,400
800-1,400
500-1,200
700-1,500
150-300
340-760
120-170
Preseason forecasts are the expected returns of the dominant age classes in a given
river system, and they are based on the number of spawning adults that produced each age
class. In the Port Moller test fishery, gill netting at standardized locations provides a daily index
of the overall number of fish entering Bristol Bay (Flynn and Hilborn 2004), with approximately
seven days' lead before they enter the commercial fishing districts. Genetic samples from the
test fishery are analyzed within four days (Dann et al. 2009, pg. 3) to give managers an advance
estimate of run strength for each of the nine major river systems. As salmon move upstream,
escapement is monitored with counting towers on each of the major rivers, except the
Nushagak where a sonar system is used. Counting towers are elevated platforms along small to
medium-sized (10-130 m wide), clear rivers from which migrating salmon are visually counted
(Woody 2007). The Nushagak River's DIDSON sonar uses sound waves to detect and
enumerate migrating salmon. Since tower and sonar monitoring occurs well upstream of the
13
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commercial fishery, all information regarding the performance of the fishery must be analyzed
on a continual basis to ensure escapement levels will be met (Clark 2005, pg. 4, Salomone et al.
2011).
The fishery is typically opened on a schedule during the early part of the season, during
which time the frequency and duration of openings are primarily based on preseason forecasts
and management is conservative. As the fishing season progresses and more information
becomes available, managers make constant adjustments to fishing time and area. If the
escapement goal is exceeded at a given monitoring station, the fishery is opened longer and
more frequently. If the escapement goal is not reached, the fishery is closed. Fishing time is
opened and closed using emergency orders, and fishermen often learn of changes only a few
hours before they go into effect. Since the bulk of the sockeye salmon harvest occurs during a
short timeframe - from the last week of June until the middle of July - this short warning system
is needed to maximize fishing time while ensuring that escapement levels are met. Migrating
fish move quickly through the fishing districts, and delaying an opener by one day during the
peak of the migration can forego the harvest of a million salmon. This is a significant loss of
revenue to individual fishermen, and compounded by the missed revenue of workers,
processors, and marketers (Clark 2005, pg. 5). The fishery will periodically close de facto during
the peak of the season when catch rates exceed processing capacity and processors stop buying
fish. This lack of buyers can also curtail salmon harvest early and late in the season when
numbers offish do not warrant keeping processing facilities operational.
In-season management is also used to help meet an escapement goal for Chinook
salmon on the Nushagak River (Table 3), where escapement is monitored by sonar. There are
also Chinook salmon goals for the Togiak, Alagnak, Naknek, and Egegik rivers and a chum
salmon goal for the Nushagak River (Table 3), but in-season management is not used to help
attain these goals.
Bristol Bay salmon fisheries are regarded as a management success (Hilborn et al.
2003a, Hilborn 2006), and Hilborn (2006) lists four contributing factors: "(1) a clear objective of
maximum sustainable yield, (2) the escapement-goal system, which assures maintenance of the
biological productive capacity; (3) management by a single agency with clear objectives and
direct line responsibility; and (4) good luck in the form of lack of habitat loss and good ocean
conditions since the late 1970s."
Table 3. Bristol Bay escapement goal ranges for Chinook and chum salmon.
River
Togiak
Nushagak
Nushagak
Alagnak
Naknek
Egegik
Species
Chinook
Chinook
chum
Chinook
Chinook
Chinook
Escapement goal
9,300 minimum
40,000-80,000
190,000 minimum
2,700 minimum
5,000 minimum
450 minimum
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Description of sport fisheries
The sport fisheries in Bristol Bay's river systems are regarded as world class. A recent
ADF&G report (Dye and Schwanke 2009) notes that "The BBMA [Bristol Bay Management Area]
contains some of the most productive Pacific salmon, rainbow trout, Arctic grayling, Arctic char
and Dolly Varden waters in the world. The area has been acclaimed for its sport fisheries since
the 1930s." Similar views prevail in the popular sport fishing literature, where articles praising
Bristol Bay as a destination are common. For example, Fly Rod and Reel (Williams 2006) says
"No place on earth is wilder or more beautiful or offers finer salmonid fishing." Over the years,
many other articles in Field and Stream, Fly Fisherman, Fish Alaska, Fly Rod and Reel, Salmon
Trout Steelheader, World Angler, and other magazines have touted the high quality fishing and
wilderness ambiance.
Large numbers of salmon and trout are caught in Bristol Bay's sport fisheries each year
(see below), but the area is best known for its rainbow trout fishing. ADF&G (1990) notes that
"Wild rainbow trout stocks of the region are world famous and are the cornerstone to a
multimillion dollar sport fishing industry." Articles in the sport fishing press laud the trout
fisheries, especially those of the Kvichak River drainage. Fish Alaska magazine calls the Iliamna
system "One of the greatest trophy trout fisheries in the world...the crown of Alaska's sport
fishing" (Weiner 2006) and names seven Bristol Bay drainages, five of which are in the
Nushagak or Kvichak river basins, in a rundown of Alaska's top ten spots for trophy rainbow
trout (Letherman 2003). Thirty-inch (76 cm) rainbow trout can be caught in many areas of the
Kvichak River and other drainages (Randolph 2006) and 43% of clients at remote Bristol Bay
sport fishing lodges reported catching a rainbow trout longer than 26 inches (66 cm) on their
most recent trip (Duffield et al. 2006, pg. 48).
Unlike commercial fisheries, whose salient features tend to be readily quantifiable (e.g.,
economics, sustainability), the quality of a sport fishery can hinge on personal and subjective
attributes. Despite the potential to catch high numbers of sizeable fish, Bristol Bay anglers rate
aesthetic qualities as most important in selecting fishing locations. Of 11 attributes that
capture different motivations and aesthetic preferences, including "catching and releasing large
numbers offish" and "chance to catch large or trophy-sized fish," Alaska resident and
nonresident anglers picked the same top five: "natural beauty of the area", "being in an area
with few other anglers", "being in a wilderness setting", "chance to catch wild fish", and
"opportunities to view wildlife" (Duffield et al. 2006, pg. 45). The same priorities apply for
nonresident anglers across Alaska (Romberg 1999, pg. 85).
The Bristol Bay region is not linked to the State's highway system and roads connected
to the major communities provide very limited access. Small aircraft with floats are the primary
source of access followed by boats based out of communities and remote lodges (Dye and
Schwanke 2009, pg. 1). A range of services are available for recreational anglers. Anglers
willing to pay $7,500 to $9,500 a week can stay in a plush remote lodge and fly to different
streams each day with a fishing guide (Purnell 2011). Modest river camps, with cabins or wall
tents, are a lower-budget option. Many self-guided expeditions center on multi-day raft trips
that use chartered aircraft for transport to and from access points along a river.
Site-specific data regarding participation, effort and harvest have been collected from
sport fishing guides and businesses since 2005 (Sigurdsson and Powers 2011). In 2010, the
most recent year for which data are available, 72 businesses and 319 guides operated in the
15
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Kvichak and Nushagak watersheds (Table 4; Dora Sigurdsson, ADF&G, unpublished data). In
addition, Table 4 shows figures for 2005, the first year of data collection, and 2008, a peak year.
Table 4. The number of businesses and guides operating in the Nushagak and Kvichak
watersheds in 2005, 2008 and 2010.
2005
2008
2010
Wateished . _ . , . ., .
Businesses Guides BusmessesGuides Businesses
Kvichak River (including Alagnak River)
Nushagak River (including Wood River)
Kvichak and Nushagak combined1
53
67
91
204
199
336
59
60
92
274
245
426
46
47
72
Guides
211
162
319
1 Business and guide totals are not additive because a business and/or guide can operate in
multiple watersheds.
Management of sport fisheries
The Alaska Department of Fish and Game's Division of Sport Fish manages recreational
fisheries in the Bristol Bay Management Area (BBMA), which includes all fresh waters flowing
into Bristol Bay between Cape Menshikof, on the Bay's southeast shore, and Cape Newenham
in the northwest. Three local management plans guide sport fishing regulations in the Bristol
Bay region (in addition to several statewide plans). The Nushagak-Mulchatna King Salmon
Management Plan and the Kvichak River Drainage Sockeye Salmon Management Plan call for
sport fishing bag limit reductions or closures by emergency order during poor runs. The
Southwest Alaska Rainbow Trout Management Plan instated conservative trout management
uniformly throughout the region, replacing the fragmentary restrictions that had been
established over the previous decades. Sport fishing regulations are updated annually and can
be accessed on ADF&G's website:
http://www.adfg. a laska.gov/index.cf m?adfg=fishregulations. sport.
The Division of Sport Fish uses the annual Statewide Harvest Survey, mailed to
randomly-selected licensed anglers, to monitor effort, catch, and harvest. Between 1997 and
2008, angler-days of effort within the BBMA ranged from 83,994 to 111,838 (Dye and Schwanke
2009, pg. 4). Total annual sport harvest for the same period ranged from 39,362 to 71,539 fish,
of which sockeye, Chinook and coho salmon comprise the majority (Dye and Schwanke 2009,
pg. 8). Resident fish species, including rainbow trout, Dolly Varden, Arctic char, Arctic grayling,
northern pike and whitefish, are also harvested in the BBMA (Dye and Schwanke 2009, pg. 8).
Harvest rates are lower for these species than for salmon, likely due to restrictive bag limits and
the popularity of catch-and-release fishing (Dye and Schwanke 2009, pgs. 6 and 8).
Chinook salmon
In the Nushagak drainage, the general season runs from May 1 to July 31 for Chinook
salmon, although some areas close on July 24 in order to protect spawners. The daily limit is
two per day, only one of which can be over 28 inches (71 cm). The annual limit is four fish. The
Nushagak-Mulchatna King Salmon Management Plan calls for an in-river return of 75,000 fish
with a spawning escapement of 65,000 fish. The guideline harvest for the sport fishery is 5,000
fish, although restrictions are triggered if the in-river return falls below 55,000 fish. In other
16
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Bristol Bay drainages, the daily limit for Chinook salmon is three and the annual limit is five,
although there are additional restrictions in the Wood and Naknek river drainages.
The major Chinook salmon sport fisheries in the BBMA include the Nushagak, Naknek,
Togiak and Alagnak rivers and the average annual harvest is 11,100 fish for the period from
1997 to 2008. The largest individual fishery takes place in the Nushagak River, where harvest
from 2003 to 2007 averaged 7,281, approximately 58% of the total Bristol Bay sport harvest for
that period (Dye and Schwanke 2009, pg. 13).
Sockeye salmon
Sockeye salmon fishing is open year round with a daily limit of five fish. Runs enter
rivers starting in late June, peak in early July, and continue into late July or early August. The
Kvichak River Drainage Sockeye Salmon Management Plan places restrictions on the sport
fishery to avoid conflicts with subsistence users when the escapement falls below the minimum
sustainable escapement goal of two million fish. Restrictions include actions such as reducing
the daily limit for sockeye and closure of areas for sport fishing that are used by both
subsistence and recreational anglers.
Sockeye are the most abundant salmon species in the BBMA. Recent annual sport
harvest ranged from 8,444 to 23,002 fish (Dye and Schwanke 2009, pg. 22). The two locations
that support the largest sport harvest are the Kvichak River, near the outlet of Iliamna Lake, and
the Newhalen River, just above Iliamna Lake (Dye and Schwanke 2009, pg. 24). Other drainages
that support moderate harvests of sockeye salmon include the Naknek and Alagnak rivers and
the Wood River lake system (Dye and Schwanke 2009, pg. 22).
Rainbow trout
Due to their relatively small spawning populations and their popularity as a game fish,
fishing regulations for rainbow trout are more restrictive than those for any other species. The
Southwest Alaska Rainbow Trout Management Plan (ADF&G 1990) calls for conservative
management, allows limited harvest in specific areas, and bans stocking of hatchery trout
(although stocking had not been practiced previously). Special management areas were
created to preserve a diversity of sport fishing opportunities: eight catch-and-release areas, six
fly-fishing catch-and-release areas, and eleven areas where only single-hook artificial lures can
be used (Dye and Schwanke 2009, pgs. 34-36).
Only single-hook artificial lures can be used in the Kvichak River drainage, and all sport
fishing is banned from April 10 through June 7 to provide protection for spawning rainbow
trout. From June 8 through October 31 anglers are allowed to keep one trout per day, with the
exception of a number of streams where no harvest is allowed. From November 1 through
April 9, when anglers are few, the daily limit increases to five fish although only one may be
longer than 20 inches (51 cm). Rainbow trout fishing regulations are similarly restrictive in
other drainages across the BBMA.
The most popular rainbow trout fisheries are found in the Kvichak drainage, the Naknek
drainage, portions of the Nushagak and Mulchatna drainages, and streams of the Wood River
Lakes system (Dye and Schwanke 2009, pg. 26). Field surveys and the Statewide Harvest Survey
show that harvest has decreased over the past decade but that total catch and effort have
remained stable or increased (Dye and Schwanke 2009, pg. 26). The annual BBMA-wide harvest
17
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between 1997 and 2008 averaged 1900 fish, but the catch estimate over this period was nearly
100 times greater (183,000 fish; (Dye and Schwanke 2009, pgs. 29 and 31). Although the
fishery is widespread, approximately eighty percent of the total catch (144,400 fish) was from
the eastern portion of the BBMA, where the Naknek and Kvichak systems are located. Eastern
BBMA streams with estimated sport catches greater than 10,000 fish in 2008 included the
Naknek, Brooks, Kvichak, Copper, and Alagnak rivers (Dye and Schwanke 2009, pg. 31).
SALMON ABUNDANCE TRENDS AROUND THE NORTH PACIFIC, WITH REFERENCE TO
BRISTOL BAY POPULATIONS
North Pacific salmon species, from most to least abundant overall, are pink, sockeye,
chum, coho, and Chinook (Ruggerone et al. 2010). The relative abundance of Pacific salmon
species relates to their life histories, as those species that are not constrained by the availability
of stream rearing habitat (i.e., pink, sockeye, and chum salmon) are able to spawn and rear in
greater numbers than those that are (i.e. coho and Chinook; Quinn 2005, pg. 319). The highest
Pacific-wide salmon harvest occurred in 2007 and totaled 513 million fish, over 300 million of
which were pink salmon (Irvine et al. 2009, pg. 2). Approximately five billion juvenile salmon
are released annually from hatcheries around the North Pacific (Irvine et al. 2009, pg. 6),
although none are reared or released in the Bristol Bay region.
Sockeye salmon
Size of Bristol Bay, Kvichak, and Nushagak sockeye salmon returns
Escapement monitoring within the Bristol Bay watershed has been conducted since the
1950s, when ADF&G established counting towers on the nine major river systems. When
combined with commercial, subsistence and sport harvest, data from escapement monitoring
allows estimates of total run sizes. A recent synthesis of salmon returns for 12 regions around
the North Pacific also extends back to the 1950s, allowing comparisons of wild sockeye salmon
returns between Bristol Bay and other regions for the period 1956 to 2005 (Ruggerone et al.
2010). The average global abundance of wild sockeye salmon over that period was 65.3 million
(M) fish, and Bristol Bay constituted the largest proportion of that total at 46% (Figure 4). Total
returns to Bristol Bay ranged from a low of 3.5 M in 1973 to a high of 67.3 M in 1980 (Figure 5),
with an annual average of 29.8 M. The region with the second largest returns is southern
British Columbia/Washington, which averaged 14% of the total (Figure 4), or 8.9 M salmon.
Other regions that produce high abundances of wild sockeye salmon include the Kamchatka
Peninsula, northern British Columbia, Cook Inlet and Kodiak Island (Ruggerone et al. 2010).
18
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Figure 4. Relative abundance of wild sockeye salmon stocks in the North Pacific, 1956-2005. See
Bristol.Bay
S AKPen
Kodiak
CookJnlet
PWS
SE.AK
Other.W..AK
E..Kamchatka
W.. Kamchatka
Russia
N..BC
S..BC.and.WA
Appendix 1 for data and sources. Stocks are ordered from west to east across the North Pacific.
19
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\ }
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1960 1980 2000
Year
Figure 5. Wild sockeye salmon abundances by region in the North Pacific, 1956-2005. See
Appendix 1 for data and sources. Each graph shows three regions organized from west to east
across the North Pacific.
20
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Hatchery production of sockeye salmon started in 1977 and accounted for an annual
average of 3 M fish, or 4% of the world total, during the 10-year period from 1995 to 2005
(Ruggerone et al. 2010). No hatchery production has occurred in the Bristol Bay region.
Regions with major hatchery production include Prince William Sound, Cook Inlet, and Kodiak
Island, which produced a respective 1.0, 0.9 and 0.6 M hatchery fish, on average, from 1995-
2005 (Ruggerone et al. 2010).
Although the Alagnak River is part of the Kvichak watershed and the Wood River is part
of the Nushagak watershed, we report sockeye salmon data separately for these systems
(unless noted otherwise) because ADF&G monitors returns on each. On average, the Kvichak
River has the largest sockeye salmon run in Bristol Bay, with an average annual return of 10.4 M
fish between 1956 and 2010 (Figure 6). Iliamna Lake provides the majority of the rearing
habitat for sockeye in the Kvichak watershed, followed by Lake Clark where the estimated
proportion of the escapement ranges from 7 to 30% (Young 2005, pg. 2). Runs exceeding 30 M
fish have occurred three times in the Kvichak River: 47.7 M, 34.6 M and 37.7 M fish returned in
1965,1970 and 1980, respectively (Tim Baker, ADF&G, unpublished data). Those runs
accounted for 57%, 49% and 40% of world production of sockeye salmon during those years
(Ruggerone et al. 2010). The Egegik River supports Bristol Bay's second largest run, with a
mean annual return of 6.3 M fish from 1956 to 2010 (Figure 6). The Nushagak and Wood rivers
are smaller runs and average returns from 1956 to 2010 were 1.3 and 3.3 M fish, respectively.
21
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50 -
40 -
30 -
20 -
10 -
0
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° 40 -
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i 30 -
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1960 1970 1980 1990 2000 2010
Year
Figure 6. Total sockeye returns by river system in Bristol Bay, 1956-2010. See Appendix 1 for
data and sources. Each graph shows three river systems listed from west to east across Bristol
Bay.
22
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The Kvichak River sockeye salmon runs are not only the largest in Bristol Bay, but also
the largest in the world (Figures 5 through 7). As noted above, returns to the Kvichak River
have averaged 10.4 M fish, and this number climbs to 11.9 M fish when returns to the Alagnak
River are included (Tim Baker, ADF&G, unpublished data). The Fraser River system supports the
world's second largest run, with an average of 8.1 M fish for the same period (Catherine
Michielsens, Pacific Salmon Commission, unpublished data). Other major producers outside of
Bristol Bay include the Copper, Kenai, Karluk, and Chignik rivers in Alaska and the Skeena River
in British Columbia (Figure 7). The Kamchatka Peninsula in Russia also has rivers with large
sockeye runs, but abundances for individual rivers were not readily available. The combined
runs for the western and eastern Kamchatka Peninsula averaged less than 5 M sockeye during
the period from 1952 to 2005 (Ruggerone et al. 2010).
23
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o
en
50 -
40
30 -
20 -
10 -
o H
50 -
o 40 H
30 -
20 -
10 -
o -
Nushagak-Wood
Kvichak-Alagnak
Chignik
Karluk
i i
- Kenai (late run)
-- Copper (wild)
Skeena
Fraser
1960
1970
1
1980
Year
1990
2000
2010
Figure 7. Sockeye salmon abundances for major rivers of the North Pacific, 1956-2010. See
Appendix 1 for data and sources. The top graph includes time series for the Nushagak-Wood
and Kvichak-Alagnak systems from 1956 to 2010, the Chignik River from 1970 to 2010, and the
Karluk River from 1985 to 2010. The bottom graph shows the Kenai River late run from 1972 to
2010, the Copper River wild run from 1961 to 2010, the Skeena River from 1985 to 2010, and
the Fraser River from 1956 to 2010. Rivers are listed in the graphs as they occur from west to
east across the North Pacific.
24
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Factors affecting Bristol Bay sockeye salmon abundance
Changes in the ocean and freshwater environments that affect sockeye salmon
abundances and trends across the North Pacific are many. A major driver is the Pacific decadal
oscillation (PDO), an inter-decadal pattern of correlated changes in sea-level pressures and sea-
surface temperatures (Mantua et al. 1997). The warm phase of the PDO is characterized by
warmer than average winter sea surface temperatures along the western coastline of North
America and increased stream flows around the Gulf of Alaska, both of which are linked to
increased salmon survival (Mantua et al. 1997, Ruggerone et al. 2007). There are three regime
shifts documented in the recent climate record that correlate with salmon productivity: 1947,
1977 and 1989. From 1947 to 1977, the PDO was in a cool phase marked by low productivity
for Alaskan and British Columbia sockeye salmon. The PDO shifted to a warm phase in 1977,
after which most North American stocks increased (Figure 5). For Bristol Bay stocks, this warm
phase corresponded with increased marine growth and, in turn, increased abundances and
numbers of recruits (returning adults) generated per spawner (Ruggerone et al. 2007). Bristol
Bay stocks more than doubled during this warm phase and remained high until the mid-90s,
when declines in the Kvichak and other rivers reduced the overall abundance (Figure 4,
Ruggerone et al. 2010). Biological indicators suggest that decreased productivity associated
with a cool phase began in 1989, while climate indices point to a short-lived reversal from 1989
to 1991, followed by a return to a warm phase (Hare and Mantua 2000). Late marine growth
and adult length-at-age of Bristol Bay sockeye decreased after the 1989 regime shift, potentially
reducing stock productivity (Ruggerone et al. 2007).
Another factor affecting sockeye salmon productivity is competition with increasing
numbers of hatchery smolts released into the North Pacific. Alaska produces the most hatchery
pink salmon in the world, averaging 42 M fish for the period 1995 to 2005, followed next by
Russia, with 12.6 M for the same period (Ruggerone et al. 2010). Approximately 75% of the
pink salmon hatchery production in Alaska occurs in Prince William Sound, with other facilities
located in Kodiak, Cook Inlet, and Southeast Alaska. Japan dominates the production of
hatchery chum salmon, with 67.3 M fish returning on average for 1995 to 2005 (Ruggerone et
al. 2010). Coming in a distant second behind Japan, Southeast Alaska averaged 9.7 M hatchery
chum salmon for the same period (Ruggerone et al. 2010). Bristol Bay sockeye smolts that
migrated to sea during even-numbered years and interacted with dominant odd-year Asian
pink salmon experienced decreased growth, survival and adult abundance compared to the
smolts that migrated during odd-numbered years (Ruggerone et al. 2003). Additionally, Kvichak
sockeye salmon productivity was negatively correlated with a running three-year mean of
Kamchatka pink salmon abundances (Ruggerone and Link 2006).
In the freshwater environment, spawning and rearing habitats can limit sockeye salmon
populations through negative density dependence. The amount of suitable spawning habitat is
fixed within a given system, so when spawning densities are high and suitable spawning sites
are occupied, females will dig nests on top of existing nests, dislodging many of the previously
laid eggs, or die without spawning (Semenchenko 1988, Essington et al. 2000). As such, the
amount of available spawning habitat can impose an upper limit on potential fry production. In
nursery lakes, juvenile growth rates decrease with rearing densities (Kyle et al. 1988, Schindler
et al. 2005a), leading to decreased survival for small individuals in the subsequent marine stage
25
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(Koenings et al. 1992). Together, these processes limit the number of recruits potentially
produced by a large spawning run.
Kvichak sockeye abundances follow five-year cycles that are unique amongst the nine
major systems of Bristol Bay. Previous hypotheses for the cycle included natural depensatory
mechanisms, such as predation, and fishing-related depensation. Since the first escapement
goal was established for the Kvichak River in 1962 until the most recent change in 2010, the
escapement goals were managed to match the cycle year. Most recently, off-cycle years had an
escapement goal range of 2 to 10 M spawners, while pre-peak and peak cycle years were
managed for escapement of 6 to 10 M spawners (Baker et al. 2009, pg. 6). In 2010, the
escapement goal was changed to one goal for all years of 2 to 10 M spawners. Ruggerone and
Link (2006) recently analyzed the population characteristics of Kvichak sockeye and found that
the cycle is likely perpetuated by three factors: density dependence during pre-peak and peak
cycle years reducing productivity in off-cycle years, higher percentage interceptions in off-cycle
years biasing productivity low, and the dominance of age 2.2 salmon (2 years in fresh water and
two years in the ocean), which return after five years. Kvichak salmon were shown to have high
interception rates in the Egegik and Ugashik fisheries in years when the Egegik and Ugashik
returns were more than double the Kvichak return, which biased the number of returning
recruits during off-cycle years. They did not find any evidence of natural depensatory
mechanisms, nor did they find reason to believe that the change in the escapement goal in
1984 could have had any effect on the decline in the 1990s.
In recent years, ADF&G has developed genetic stock identification methods, which are
being used to reanalyze past interceptions of Kvichak salmon from the mixed stock fisheries on
the east side of the Bay (Dann et al. 2009, pg. 37). It is anticipated that current brood tables
from which total runs by system are reconstructed will change as this analysis progresses (Tim
Baker, ADF&G, personal communication) giving researchers a more accurate understanding of
the dynamics of Bristol Bay stock composition and return dynamics.
The decline in Kvichak River sockeye salmon runs
From 1977 through 1995, during the warm PDO phase, Bristol Bay runs averaged almost
41 M fish annually, while runs to the Kvichak River averaged nearly 15 M, comprising about 36%
of the entire Bristol Bay run (Table 5). Beginning in 1996, with the spawning return of the 1991
brood year, Kvichak runs dropped to an average of 4.7 M fish, comprising less than 14% of the
total Bristol Bay run (Table 5). This decline was accompanied by a decline in stock productivity,
as expressed by the number of recruits generated per spawner (R/S). Bristol Bay systems
averaged approximately two recruits for every spawner prior to the 1977 regime shift, and R/S
increased substantially for many systems, such as the Egegik and Ugashik rivers, during the
subsequent warm phase (Hilborn 2006). R/S for the Kvichak averaged 3.2 for the 1972 to 1990
broods, but five of the nine broods from 1991 onward failed to replace themselves (i.e., R/S
<1). Productivity also decreased during this time in two other systems on the east side of
Bristol Bay, the Egegik and Ugashik rivers (Ruggerone and Link 2006). The decline in the Kvichak
River run led ADF&G to classify it as a stock of yield concern in 2001 (Morstad and Baker 2009,
pg. 1), indicating an inability to maintain a harvestable surplus. The Kvichak run was further
downgraded to a stock of management concern in 2003, based on failure to meet escapement
goals.
26
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Table 5. Mean annual returns of sockeye salmon in Bristol Bay, 1956-2010, and percent of total
by river system. See Appendix 1 for data and sources. Rivers are listed from east to west across
Bristol Bay.
Rivers
Ugashik
Egegik
Naknek
Alagnak
Kvichak
Nushagak
Igushik
Wood
Togiak
Total
1956-1976
882,458
2,320,059
2,200,534
514,544
10,482,754
392,574
516,021
1,707,120
305,069
19,321,134
%
4.6
12.0
11.4
2.7
54.3
2.0
2.7
8.8
1.6
1977-1995
4,123,115
9,100,953
4,454,164
1,360,651
14,784,340
1,919,420
1,349,775
3,150,620
661,011
40,904,050
%
10.1
22.2
10.9
3.3
36.1
4.7
3.3
7.7
1.6
1996-2010
3,522,697
8,402,365
5,251,810
3,008,922
4,757,008
1,933,461
1,341,581
5,834,787
742,696
34,795,327
%
10.1
24.1
15.1
8.6
13.7
5.6
3.9
16.8
2.1
1956-2010
2,722,023
6,321,361
3,811,227
1,487,121
10,407,190
1,340,272
1,029,198
3,331,511
547,384
30,997,285
%
8.8
20.4
12.3
4.8
33.6
4.3
3.3
10.7
1.8
Ruggerone and Link (2006) analyzed the decline in the Kvichak run starting with the
1991 brood year and identified a number of potential factors. The number of smolts per
spawner declined by 48% and smolt-to-adult survival declined by 46%, suggesting that factors
in both freshwater and marine habitats were involved. The average number of smolts out-
migrating from the Kvichak River during the years 1982 to 1993 was approximately 150 M,
which declined to an approximate average of 50 M from 1994 to 2001 (Ruggerone and Link
2006). The declines were accompanied by a shift in the dominant age structure of Kvichak
spawners from 2.2 (i.e., two years in fresh water followed by two years at sea), which
represented an average of 84% of the return, to 1.3, indicating that salmon were spending less
time in fresh water and more time at sea. Across the nine monitored Bristol Bay watersheds,
the decrease in the percentage of 2.2 salmon in the total return correlated strongly with
decreases in R/S and run size. The decrease in spawner length at age starting in 1991 and
higher than normal sea surface temperatures in June from 1990-1998 both may have
contributed to lower reproductive potential, since smaller females produce fewer eggs.
Competition with Asian pink salmon also may have played a role. Abundances of Asian pink
salmon have been linked to decreased size at age of returning Bristol Bay sockeye salmon in
addition to decreased abundance during even-year migrations when interactions are highest
(Ruggerone et al. 2003). Abundances of Kamchatka pink salmon were high from 1994 to 2000,
the beginning of which correlates to age-1 smolts from the 1991 brood year. The three eastern
Bristol Bay stocks that experienced the largest declines during the 1990s (Kvichak, Egegik and
Ugashik rivers) have greater overlap with Asian pink salmon stocks in their marine distribution
than other stocks that did not decline significantly (Ruggerone and Link 2006, pg. 31).
Ultimately, conditions outside of the freshwater environment likely led to the decline of
Kvichak sockeye salmon. Warmer summer temperatures in both fresh water (Schindler et al.
2005a) and the ocean (Hare and Mantua 2000) and interactions with Asian pink salmon
27
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affected Kvichak sockeye salmon disproportionately to other systems due to the dominance of
ocean-age-two salmon in the Kvichak watershed (Ruggerone and Link 2006, pg. 12). Because
ocean-age-two salmon interact with only one Asian pink salmon population at sea, the effects
on growth and abundance are greater than for ocean-age-three salmon, which interact with
both large (even) and small (odd) Asian pink salmon populations at sea and thus, have the
opportunity for higher growth rates during odd years (Ruggerone et al. 2003). The decrease in
spawner to smolt survival may also be related to marine conditions causing smaller length at
age of returning adults and reduced reproductive success (Ruggerone and Link 2006, pg. 15).
In 2009, following several years of improvement, ADF&G upgraded the Kvichak's
classification to a stock of yield concern (Morstad and Baker 2009). Since 2004, Bristol Bay
returns have again totaled more than 40 million fish annually and in 2010 the Kvichak run
increased to over 9.5 million fish, equating to 23% of the total for the Bay.
Chinook salmon
The total commercial harvest of Chinook salmon in the North Pacific ranged between
three and four million fish until the early 90s; recent total catches have decreased to one to two
million fish (Eggers et al. 2005). Lacking escapement data for many runs, commercial harvest is
a good surrogate for salmon abundance, and suggests a decline in Chinook salmon abundance
in recent decades. The U.S. makes over half of the total commercial catch, followed by Canada,
Russia, and Japan (Heard et al. 2007). Recreational, subsistence, and aboriginal catch is
significant for this salmon species and totaled approximately one million annually in 2003-2004
(Heard et al. 2007). Washington dominates hatchery production of Chinook salmon, with over
one billion juveniles released annually from 1993-2001 (Heard et al. 2007).
The Columbia River historically produced the largest Chinook salmon run in the world,
with peak runs (spring, summer, and fall combined) estimated at 3.2 M fish during the late
1800s (Chapman 1986). Peak catches for the Columbia River summer-run Chinook salmon
occurred at this time, until overfishing decimated the run. Fishing effort then shifted to the fall
run, which suffered a similar demise in the early 1900s. There are currently five stocks of
Chinook salmon in the Columbia River watershed listed under the Endangered Species Act and
the majority of the current returns are hatchery fish (70%, 80% and 50% of the spring, summer
and fall runs, respectively; Heard et al. 2007).
Currently, the largest runs of Chinook salmon in the world originate from three of the
largest watersheds that drain to the North Pacific: the Yukon, Kuskokwim and Fraser rivers
(Table 6). Total Chinook escapements to the Kuskokwim and Yukon rivers have not been
quantified directly due to their large watershed area, but recent total run estimates based on
mark-recapture studies put them at 217,000 and 265,000 fish, respectively (Molyneaux and
Brannian 2006, pg. 102, Spencer et al. 2009, pg. 28). On the Fraser River, the average size of
the spring, summer, and fall Chinook runs combined (including the Harrison River) for the most
recent ten-year period (2000-2009) was 287,000 fish (PSC 2011, pg. 87).
28
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Table 6. Nushagak River Chinook average run sizes for 2000-2009, in comparison to other rivers
across the North Pacific. Other rivers are sorted in order of decreasing run size.
Watershed
Nushagak R.
Region
Bristol Bay, Western Alaska
Average run size
(2000-2009)
151,348 1
Area15
(km2)
31,383
Fraser R., total run
Kuskokwim R., total run
Yukon R., total run
Harrison R. (trib. of Fraser R.)
Taku R.
Copper R.
Kenai R. (early and late runs)
Skeena R.
Yukon R., Canadian mainstem
Nass R.
Grays Harbor (Chehalis R. + 5 others)
Skagit R.
Nehalem R.
Bristish Columbia, Canada
Western Alaska
Western Alaska
Bristish Columbia, Canada
Southeast Alaska
Southcentral Alaska
Southcentral Alaska
Bristish Columbia, Canada
Yukon Territory, Canada
Bristish Columbia, Canada
Washington
Washington
Oregon
287,475 2
284,000 3
217,405 4
98,257 5
78,081 6
75,081 7
70,976 8
63,356 9
59,346 10
31,738 n
23,964 12
18,286 13
12,267 14
233,156
118,019
857,996
7,870
17,639
64,529
5,537
51,383
323,800
20,669
6,993
8,234
2,193
Unpublished data, Gregory Buck, ADF&G
2 Pacific Salmon Commission 2011, pg. 88
3 Unpublished data, Kevin Schaberg, ADF&G
4 Average from 2000-2004, Spencer et al. 2009, pg. 28
5 Pacific Salmon Commission 2011, pg. 88
6 McPherson et al. 2010, pg. 14
7 Unpublished data, Steve Moffitt, ADF&G
8 Begich and Pawluk 2010, pg. 69
9 Pacific Salmon Commission 2011, pg. 87
10 Howard etal. 2009, pg. 35
11 Pacific Salmon Commission 2011, pg. 87
12 Pacific Salmon Commission 2011, pg. 90
13 Pacific Salmon Commission 2011, pg. 89
14 Pacific Salmon Commission 2011, pg. 93
15 Watershed area from the Riverscape Analysis Project 2010 (http://rap.ntsg.umt.edu).
29
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Chinook sport and commercial harvests in the Nushagak River are larger than all of the
other systems in Bristol Bay combined (Dye and Schwanke 2009, pg. 13, Salomone et al. 2011,
pg. 86). The Nushagak produces runs that are periodically at or near the world's largest (Figure
8), which is remarkable considering its relatively small watershed area (Table 6). Returns
consistently number over 100,000 fish, while returns greater than 200,000 fish have occurred
eleven times between 1966 and 2010 (Figure 8). An especially productive six-year period from
1978-1983 produced three returns greater than 300,000 fish (Figure 8). Other rivers that
produce large returns of Chinook salmon include the Copper, Kenai, and Taku rivers in Alaska
and the Skeena and Harrison rivers in British Columbia (Table 6). The Harrison River is the
dominant fall run stock for the Fraser River.
30
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tn
o
500 -
400 -
300 -
200 -
100 -
0 -
500 -
Yukon (Canadian stock)
Kuskokwim
Nushagak
Kenai j
^ U/ '? s.
%AW-V .,xt
To 400 H
"o
h-
300 -
200 -
100 -
0 -
Copper
Taku
Skeena
Fraser
M
f \
\ ,
J ' \'X
- r
1 s
1970
1980
2000
2010
1990
Year
Figure 8. Chinook salmon abundances by river system, 1966-2010. See Appendix 1 for data and
sources. The top graph shows total runs for the Yukon River (Canadian stock) from 1982 to
2009, the Kuskokwim River from 1976 to 2010, the Nushagak River from 1966 to 2010, and the
Kenai River from 1986 to 2010. The bottom graph shows total runs for the Copper River from
1980 to 2010, the Taku River from 1973 to 2010, the Skeena River from 1977 to 2009, and the
Fraser River from 1984 to 2009. Rivers are organized from west to east across the North
Pacific.
31
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A sustainable escapement goal (SEG) was implemented for Nushagak Chinook salmon in
2007 with a target of 40,000 to 80,000 fish. Sonar counts used to estimate escapement were
initiated in 1989 and since that time, the Nushagak run has consistently met the minimum
escapement for the current SEG and was over the SEG 12 times (Gregory Buck, ADF&G,
unpublished data). The Nushagak Chinook stock is considered stable (Heard et al. 2007, Dye
and Schwanke 2009, pg. 17), in contrast to Chinook stocks on the Kuskokwim and Yukon rivers,
which experienced declines starting in the late 1990s. Both the Yukon and Kuskokwim Chinook
were listed as stocks of yield concern in 2000 (Estensen et al. 2009, pg. 2, Howard et al. 2009,
pg. 1). The Yukon River stock is still listed but the Kuskokwim River Chinook stock was delisted
as a stock of concern in 2007, based on higher than normal returns starting in 2004 (Estensen et
al. 2009, pg. 2).
The decline in Yukon and Kuskokwim Chinook stocks that began in the late 1990s may
have resulted from the 1997-1998 El Nino (Kruse 1998, Myers et al. 2010 pg. 199). That event
was characterized by sea surface temperatures at least 2° C higher than normal in the Bering
Sea, along with weak winds and high solar radiation that led to two anomalous phytoplankton
blooms, typically associated with nutrient-limited waters (Kruse 1998). The decline in Chinook
stocks that persisted after the 1997-1998 El Nino indicate that multiple ocean age classes were
affected by this event (Ruggerone et al. 2009).
Chinook salmon hatchery production contributes to harvests in both southeast and
southcentral Alaska. The average number of returning hatchery Chinook salmon in Alaska for
2000 to 2009 was 118,000 fish annually and, in 2009, hatchery Chinook salmon contributed
16% of the total commercial harvest for the State (White 2010). There are no salmon
hatcheries located in western Alaska and none of the total runs for the Alaskan rivers listed in
Figure 8 or Table 6 include contributions from hatcheries (Yukon, Kuskokwim, Nushagak, Kenai,
Copper, and Taku rivers). Salmon enhancement programs for Chinook salmon in British
Columbia are significant; for the period 1990 to 2000, hatchery releases averaged
approximately 50 million fish annually and hatcheries contributed approximately 30% to the
total Canadian catch (MacKinlay et al. Undated). The Chehalis River hatchery in the Harrison
River watershed and the Chilliwack River, Inch Creek, and Spius Creek hatcheries in the Fraser
River watershed all contribute to the Chinook salmon runs on those systems (FOC 2011).
Threatened and endangered salmon and conservation priorities
Although it is difficult to quantify the true number of extinct salmon populations around
the North Pacific, estimates for the Western United States (California, Oregon, Washington and
Idaho) have ranged from 106 to 406 populations (Nehlsen et al. 1991, Augerot 2005, pg. 65,
Gustafson et al. 2007). Chinook had the largest number of extinctions followed by coho and
then either chum or sockeye (Nehlsen et al. 1991, Augerot 2005, pg. 67). Many of the patterns
of population extinction are related to time spent in fresh water: interior populations have
been lost at a higher rate than coastal populations, stream-maturing Chinook and steelhead
(which may spend up to nine months in fresh water before spawning) had higher losses than
their ocean-maturing counterparts, and species that relied on fresh water for rearing (Chinook,
coho, and sockeye) had higher rates of extinction than pink or chum salmon, which go to sea
soon after emergence (Gustafson et al. 2007). No populations from Alaska are known to have
gone extinct. Salmon populations in the southern extent of their range have suffered higher
32
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extinction rates and are considered at higher risk than populations further to the north (Brown
et al. 1994, Kope and Wainwright 1998, Rand 2008).
In addition to the large number of populations now extinct, there are many that are
considered at risk due to declining population trends. The Columbia River basin dominated the
list of at risk stocks identified by Nehlson et al. (1991), contributing 76 stocks to the total of 214
for California, Oregon, Washington, and Idaho. Approximately half of the 214 stocks evaluated
were listed as high risk because they failed to replace themselves (fewer than one recruit per
spawner) or had recent escapements below 200 individuals. More recent analyses of the status
of salmon populations in the North Pacific continue to highlight the declines in the Pacific
Northwest. A detailed assessment of salmon populations in the Columbia River basin from
1980 to 2000 showed that many are declining and this trend is heightened when hatchery fish
are excluded (McClure et al. 2003). A comparison between time periods reflecting both good
and bad ocean productivity for Columbia River salmon populations further indicates that the
declining trends are not due to the regime shift of 1977 (McClure et al. 2003). An analysis of
over 7,000 stocks across the North Pacific found that over 30% of sockeye, Chinook, and coho
stocks were at moderate or high risk and that the Western U.S. (Washington, Oregon,
California, and Idaho) had the highest concentrations of high-risk stocks (Augerot 2005, pgs. 66-
67).
A detailed assessment of sockeye salmon populations across the North Pacific highlights
threats for this species in British Columbia (Rand 2008). At the global population level, sockeye
salmon are considered a species of least concern. Eighty subpopulations were identified for
assessment, five of which are extinct and 26 did not have the necessary data with which to
conduct a status assessment. Of the remaining 49 subpopulations, 17 were identified as
threatened and two as nearly threatened. British Columbia has 12 threatened (vulnerable,
endangered, or critically endangered) subpopulations, 70% of the worldwide total. Three key
threats to sockeye salmon were identified: mixed stock fisheries that lead to high harvests of
small, less productive populations; poor marine survival rates and high rates of disease in adults
due to changing climatic conditions; and negative effects of enhancement activities such as
hatcheries and spawning channels (Rand 2008). Twenty-five subpopulations were assessed for
Alaska: 10 were data deficient, 12 were of least concern (including the one subpopulation
identified for Bristol Bay), one subpopulation in the eastern Gulf of Alaska was listed as
vulnerable (four of six sites had declining trends: Bering, East Alsek, Italic, and Situk rivers), and
two subpopulations in Southeast Alaska (McDonald and Hugh Smith Lakes) were listed as
endangered. Both the Hugh Smith and McDonald Lake populations were listed as stocks of
management concern by ADF&G in 2003 and 2009, respectively (Piston 2008, pg. 1, Eggers et
al. 2009, pg. 1). Both were de-listed within four years after runs met escapement goals for
several consecutive years following implementation of successful fishing restrictions (Piston
2008, pg. 1, Regnart and Swanton 2011).
Government agencies in the United States and Canada are tasked with identifying and
protecting salmon populations at risk. In the U.S., the National Marine Fisheries Service (NMFS)
manages listings of salmon species under the Endangered Species Act (ESA). Salmon stocks
considered for listing under ESA must meet the definition of an Evolutionary Significant Unit
(ESU): it must be substantially reproductively isolated from other nonspecific population units
and it must represent an important component of the evolutionary legacy of the species
33
-------
(Federal Register 58612, November 20, 1991). Current determinations for the U.S. include one
endangered and one threatened ESU for sockeye; two threatened ESUs for chum; one
endangered, three threatened, and one ESU of concern for coho; two endangered, seven
threatened, and one ESU of concern for Chinook; and one endangered, ten threatened, and
one ESU of concern for steelhead (Table 7, NMFS 2010). All listed ESUs occur in the western
contiguous U.S. (California, Oregon, Washington, and Idaho). In Canada, the Committee on the
Status of Endangered Wildlife in Canada (COSEWIC) conducts status assessments to determine
if a species is at risk nationally. The Minister of the Environment and the federal cabinet then
decide whether to list the species under the Species at Risk Act (SARA). Currently, COSEWIC
status assessments have recommended listing two endangered sockeye salmon populations,
one endangered coho salmon population, and one threatened Chinook salmon population, but
none of these assessments have resulted in legal listings under SARA (COSEWIC 2009). On the
Asian side of the Pacific, no information was found regarding listings of threatened or
endangered salmon populations under a legal framework. Other assessments of Asian salmon
distribution and status have relied on interviews with fishery biologists due to the scarcity of
data and the dominance of hatcheries in Japanese fisheries (Augerot 2005, pg. 66, Rand 2008).
34
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Table 7. Endangered Species Act listings for salmon ESUs in the United States.
Species
Chinook
Chinook
Chinook
Chinook
Chinook
Chinook
Chinook
Chinook
Chinook
Chinook
chum
chum
coho
coho
coho
coho
coho
sockeye
sockeye
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
steelhead
ESU Name
Sacramento River Winter-run
Upper Columbia River Spring-run
California Coastal
Central Valley Spring-run
Lower Columbia River
Puget Sound
Snake River Fall-run
Snake River Spring/Summer-run
Upper Willamette River
Central Valley Fall- and Late Fall-run
Hood Canal Summer-run
Columbia River
Central California Coast
Southern OR/Northern CA Coasts
Lower Columbia River
Oregon Coast
Puget Sound/Strait of Georgia
Snake River
Ozette Lake
Southern California
California Central Valley
Central California Coast
Lower Columbia River
Middle Columbia River
Northern California
Puget Sound
Snake River Basin
Southcentral California Coast
Upper Columbia River
Upper Willamette River
Oregon Coast
ESA Listing Status
endangered
endangered
threatened
threatened
threatened
threatened
threatened
threatened
threatened
species of concern
threatened
threatened
endangered
threatened
threatened
threatened
species of concern
endangered
threatened
endangered
threatened
threatened
threatened
threatened
threatened
threatened
threatened
threatened
threatened
threatened
species of concern
Date of Most
Recent Review
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
4/15/2004
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
8/15/2011
4/15/2004
8/15/2011
8/15/2011
1/5/2006
8/15/2011
1/5/2006
8/15/2011
8/15/2011
1/5/2006
8/15/2011
8/15/2011
1/5/2006
8/15/2011
8/15/2011
4/15/2004
The causes leading to extinction and continued population declines are numerous and
analyses are confounded by the effects of interacting factors within watersheds. In California,
both the building of dams that eliminated access to upstream spawning and rearing areas and
destruction of coastal habitat from extensive logging were major contributors to the decline of
coho salmon populations in the southern extent of their range (Brown et al. 1994). Heavy
fishing pressure at the end of the 19th century followed by extensive impacts to river habitats
35
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from agriculture, logging, mining, irrigation and hydroelectric dams all led to the extensive
decline of Columbia River salmon by the mid 20th century (Chapman 1986, McConnaha et al.
2006).
Restoration activities to help restore salmon habitat and populations in the Pacific
Northwest require huge expenditures with results that are often difficult to measure due to
annual variation, the time lapse between restoration action and effect on the population, and
changing climate and ocean conditions (GAO 2002, pg. 4). Approximately $1.5 billion was spent
on Columbia River salmon and steelhead for the period 1997 through 2001 (GAO 2002, pg. 2).
Predicted outcomes from restoration rarely take into account climate change scenarios.
Models developed to predict the outcome of restoration on Snohomish basin Chinook salmon
habitat showed that increased temperatures resulting from climate change changed snow to
rain in high elevation watersheds and affected three hydrologic parameters that decreased fish
populations: higher flows during egg incubation, lower flows during spawning, and increased
temperatures during pre-spawning (Battin et al. 2007). Often used as mitigation for lost habitat,
salmon hatcheries have resulted in decreased survival of the wild populations they are intended
to support (NRC 1996, pg. 319, Naish et al. 2008). Impacts of hatchery fish include overfishing of
wild populations in mixed-stock fisheries (Hilborn and Eggers 2000), competition with wild
salmon in both fresh water and the ocean (Ruggerone and Nielsen 2009), and a reduction in life
history diversity making populations more susceptible to climate variability (Moore et al. 2010).
Due to the high costs of restoration and the difficulty in predicting or measuring
outcomes, some have argued that the best way to protect salmon for future generations is to
create salmon sanctuaries that maintain intact and connected habitats throughout the
watershed from headwaters to the ocean (Rahr et al. 1998, Lichatowich et al. 2000, Rahr and
Augerot 2006). Protecting entire watersheds is especially important to sockeye, Chinook, and
coho salmon, which spend 1-2 years rearing in fresh water prior to entering the ocean. These
sanctuaries would provide habitat for salmon populations with heightened resilience to factors
outside of management control, such as climate change and changes in the ocean environment.
The salmon populations in Bristol Bay meet all the criteria for selecting sanctuaries across the
North Pacific by having intact habitats, abundant populations, and a high diversity of life history
patterns (Schindler et al. 2010). In addition, several studies have targeted Bristol Bay as a high
priority for salmon conservation. The Kvichak, Nushagak, and Wood watersheds were ranked
third, 44th, and fourth, respectively, in an analysis of physical complexity of 1574 watersheds
from California to the Kamchatka Peninsula (Luck et al. 2010, FLBS 2011). Pinsky et al. (2009)
characterized high conservation value salmon catchments across the North Pacific as the top
20% (out of 1046 total) based on abundance and run timing diversity. Bristol Bay, the
Kamchatka Peninsula, and coastal British Columbia all had clusters of high conservation value
catchments. Fewer than 9% of the high conservation value watersheds had greater than half of
their area under protected status.
36
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KEY HABITAT ELEMENTS OF BRISTOL BAY RIVER SYSTEMS (OR WHY DO BRISTOL
BAY WATERSHEDS PRODUCE SO MANY FISH?)
No published materials specifically address the question "Why do Bristol Bay watersheds
support so many salmon?" While this isn't particularly surprising given the complexity and
scope of the question, it does require us to draw on experts and a diverse body of literature to
posit an answer. Obviously, the simplest answer is "Habitat." But what is it about the habitat
in Bristol Bay watersheds that allows them to sustain such prolific fisheries? Our inquiry led us
to the conclusion that interplay between the quantity, quality, and diversity of habitats in these
river systems accounts for their productivity. The major habitat attributes discussed here were
identified in personal communications with Dr. Tom Quinn (University of Washington) and Dr.
Jack Stanford (University of Montana).
Habitat quantity
An obvious feature of the Bristol Bay watershed is the abundance of large lakes (Figure
9). The Kvichak River drains Iliamna Lake, Alaska's largest, in addition to Lake Clark, Nonvianuk
Lake, Kukaklek Lake, and an array of smaller lakes. The Nuyakuk River, a major tributary to the
upper Nushagak River, drains Nuyakuk, Tikchik, Chauekuktuli, Chikuminuk, Upnuk, Nishlik, and
a number of smaller lakes. The Wood River, a major tributary to the lower Nushagak River,
drains an interconnected chain of four major lakes- lakes Kulik, Beverly, Nerka, and Aleknagik-
and several smaller lakes. Lakes cover 7.9% of the Bristol Bay region, which is substantially
higher than the other major salmon-producing regions analyzed (Table 8). Lakes cover 13.7% of
the Kvichak River basin (Table 8). Within the Nushagak River basin, lakes cover 11.3% of the
Wood River drainage and a much smaller percentage of the remainder (1.7%; Table 8). With
the exception of Chikuminuk Lake, all of the major lakes named above are accessible to
anadromous salmon.
Since watershed elevations in the Bristol Bay region are relatively low (Table 8), barriers
to fish migration are few and a large proportion of the watershed can be accessed by salmon.
The Nushagak and Kvichak watersheds have over 58,000 km of streams (National Hydrography
Dataset), of which 7,671 km (13%) have been documented as anadromous fish streams (ADF&G
2011 Anadromous Waters Catalog; Figure 9). Since fish use must be documented firsthand by
field biologists, a large proportion of anadromous fish habitat undoubtedly remains
undocumented. For example, a recent survey targeted 135 undocumented headwater (i.e., 1st-
and 2nd-order) stream reaches with low to moderate gradient (i.e., <10% channel slope) north
of Iliamna Lake (Woody and O'Neal 2010, pgs. 11-12). Of these stream reaches, 16% were dry
or nonexistent, 53% had juvenile salmon, 66% had resident fish, and 3% contained no fish at
the time of sampling (Woody and O'Neal 2010, pg. 22).
37
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Table 8. Comparison of landscape features potentially important to sockeye salmon production for watersheds across the North
Pacific (top portion of table) and across the Bristol Bay watershed (bottom portion of table). All landscape data are from the
Riverscape Analysis Project (Luck et al. 2010).
Watershed
Kamchatka
Kenai
Copper
Fraser
Columbia
Bristol Bay
Location
Russia
Central Alaska
Prince William Sound
British Columbia
Washington
Western Alaska
Watershed
area (km2)
53,598
5,537
64,529
233,156
669,608
88,233
Mean
watershed
elevation
(m)
549
522
1,194
1,188
1,328
269*
Number
of lakes
>lkm2
82
2
9
119
68
69
Average
elevation of
lakes (m)
15
97
448
763
1,212
219*
% Lake
coverage in
watershed
0.4
2.9
0.5
1.6
0.2
7.9*
Mean annual
sockeye run
(millions offish,
1990-2005)+
3.2
5.2
3.0
10.7
42.8
Togiak
Igushik
Nushagak (inc. Wood)
Kvichak (inc. Alagnak)
Naknek
Egegik
Ugashik
Bristol Bay
Bristol Bay
Bristol Bay
Bristol Bay
Bristol Bay
Bristol Bay
Bristol Bay
4,600
2,126
35,237
25,328
9,624
7,117
4,201
322
74
250
340
312
168
104
6
2
20
29
8
1
3
160
15
325
193
230
4
4
1.4
3.3
2.7
13.7
8.3
16.5
9.9
0.7
1.3
6.0
10.9
5.2
11.0
3.8
Some figures for Bristol Bay represent the weighted average of individual Bristol Bay watersheds.
+Salmon abundance sources: Kamchatka, Fraser, and Columbia are from Ruggerone et al. 2010 (Fraser and Columbia rivers were
combined into one region "Southern B.C. and Washington."); Kenai is from sockeye brood tables for Kenai River (pers. comm. Pat
Shields, 2011); Copper is from sockeye brood tables for Copper River (pers. comm. Jeremy Botz, 2011); Bristol Bay and individual
watersheds within Bristol Bay are from sockeye brood tables for Bristol Bay (pers. comm. Tim Baker, 2011).
38
-------
Kvichak*
Total length ofanadromous
streams =7,671 km
Figure 9. Map of surveyed anadromous streams in the Nushagak and Kvichak watersheds. Data are from ADF&G 2011 Anadromous
Waters Catalog.
39
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Habitat quality
In addition to the overall abundance of salmon habitat, there are a number of habitat
attributes that likely contribute to the productivity of Bristol Bay's river systems. First of all,
Bristol Bay streambeds tend to have abundant gravel, which is essential substrate for salmon
spawning and egg incubation (Bjornn and Reiser 1991, pgs. 95-97 , Quinn 2005, pg. 108).
Several Pleistocene glacial advances have left behind a complex landscape of gravel-rich
moraines, melt-water deposits, and outwash plains (Stilwell and Kaufman 1996, Hamilton and
Kleiforth 2010). As stream channels meander and cut through these deposits, gravel and other
sediments are captured and formed into riffles, bars and other habitat features. In a survey of
76 wadeable stream reaches across the Kvichak and Nushagak watersheds, gravel (2-64 mm)
was the dominant substrate, covering 56% (±15%) of each streambed (D.J. Rinella, unpublished
data).
Groundwater inputs to streams and lakes are also an important feature of salmon
habitat in the Kvichak and Nushagak watersheds. Rainwater and melting snow infiltrate the
extensive glacial deposits and saturate pore spaces below the water table, thus recharging the
groundwater aquifer (Power et al. 1999, pg. 402). Ponds are common on the Bristol Bay
landscape and contribute disproportionately to groundwater recharge (Rains 2011). Once in
the aquifer, groundwater flows slowly downhill and eventually surfaces in areas of relatively
low elevation, like stream channels or lake basins. Areas of groundwater upwelling are heavily
used by spawning sockeye salmon because they provide circulation, stable flows, and stable
temperatures (Burgner 1991, pgs. 16-19). These habitats include lake beaches and spring-fed
ponds, creeks, and side channels (Burgner 1991, pgs. 16-19). Studies in the Wood River system
of Bristol Bay demonstrate the importance of groundwater upwelling to spawning sockeye
salmon. In lakes, densities of beach spawners were highest at sites with the strongest
upwelling, while spawners were absent at beach sites with no upwelling (Burgner 1991, pg. 19).
Beach spawners comprise substantial portions of the spawning populations in three of the four
main Wood River lakes: 47% in Nerka, 87% in Beverly, 59% in Kulik, but only 3% in Aleknagik
(1955-1962; Burgner et al. 1969, pg. 420). In a spring-fed tributary to Lake Nerka, the
distribution of sockeye salmon spawners also corresponded with areas of groundwater
upwelling (Mathisen 1962, pgs. 145-146). Large numbers of sockeye salmon in the Kvichak
River system also spawn in lake beaches, spring-fed ponds, and other groundwater-associated
habitats (Morstad 2003, pgs. 2-17). In addition to spawning sockeye, groundwater is an
important habitat feature for other salmon species and life history stages. Chum salmon have
been shown to preferentially spawn in areas of groundwater upwelling (Salo 1991, pg. 240,
Leman 1993). Groundwater also maintains ice-free habitats used extensively by wintering
fishes, helps to maintain streamflow during dry weather, and provides thermal refuge during
periods of warm water (Reynolds 1997, Power et al. 1999).
Salmon themselves are another important habitat feature of Bristol Bay watersheds.
Each year, the region's spawning salmon populations convey massive amounts of energy and
nutrients from the North Pacific to fresh waters. These marine-derived nutrients (MDN),
released as excreta, carcasses, and energy-rich eggs, greatly enhance the productivity of
freshwater ecosystems, making Pacific salmon classic examples of keystone species that have
40
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large effects on the ecosystems where they spawn (Willson and Halupka 1995, Power et al.
1996).
Salmon contain limiting nutrients (i.e, nitrogen and phosphorus) and energy (i.e.,
carbon) in the same relative proportions as needed for growth by rearing fishes, making MDN
an ideal fertilizer for salmon ecosystems (Wipfli et al. 2004). Given the high densities of
spawning salmon in some streams, MDN subsidies can be large. On average, spawning sockeye
salmon import 50,200 kg of phosphorus and 397,000 kg of nitrogen to the Kvichak River system
and 12,700 kg of phosphorus and 101,000 kg of nitrogen to the Wood River system each year
(Moore and Schindler 2004). In high latitudes, the importance of marine nutrients is magnified
by the low ambient nutrient levels in freshwater systems (Gross et al. 1988, Perrin and
Richardson 1997). In Iliamna Lake, for example, nitrogen inputs from spawning salmon greatly
exceed inputs from the watershed (Kline et al. 1993).
Resident fishes (e.g., rainbow trout, Dolly Varden, Arctic grayling) and juvenile salmon of
species that rear for extended periods in streams (i.e., coho and Chinook) derive clear and
substantial nutritional benefits through the consumption of salmon eggs and flesh and other
food sources related to spawning salmon (Bilby et al. 1996). In streams in the Nushagak River
basin, for example, ration size and energy consumption among rainbow trout and Arctic graying
increased by 480 to 620% after the arrival of spawning salmon (Scheuerell et al. 2007). The
increase in rainbow trout diet was attributable to salmon eggs, salmon flesh, and maggots that
colonized salmon carcasses, while the increase in Arctic grayling diet was attributable to
consumption of benthic invertebrates dislodged by spawning salmon (Scheuerell et al. 2007). A
bioenergetics model suggested that these subsidies were responsible for a large majority of the
annual growth of these fish populations (Scheuerell et al. 2007). In a stream in the Kvichak
River basin, Dolly Varden moved into ponds where sockeye salmon spawned and fed almost
entirely on salmon eggs (Denton et al. 2009). The growth rate of these Dolly Varden increased
three-fold while salmon eggs were available (Denton et al. 2009). On the Kenai Peninsula,
Alaska, recent work has shown that the number of salmon spawning in a given stream is an
important predictor of the growth rate and energy storage among coho salmon and Dolly
Varden rearing there (Rinella et al. in press). These and other studies indicate that the
availability of MDN enhances growth rates (Bilby et al. 1996, Wipfli et al. 2003, Giannico and
Hinch 2007), body condition (Bilby et al. 1998), and energy storage (Heintz et al. 2004) of
stream-dwelling fishes, likely leading to increased chances of survival to adulthood (Gardiner
and Geddes 1980, Wipfli et al. 2003, Heintz et al. 2004).
MDN is also linked with bottom-up effects on aquatic food webs, but any resulting
effects on fish are less clear. In streams, increased standing stocks of biofilm (Wipfli et al. 1998,
Wipfli et al. 1999, Johnston et al. 2004, Mitchell and Lamberti 2005) and macroinvertebrates
(Claeson et al. 2006, Lessard and Merritt 2006, Walter et al. 2006) have been associated with
MDN inputs. Stream-dwelling fishes may benefit indirectly through increased
macroinvertebrate production, but this has yet to be established. Likewise, MDN can comprise
a major proportion of the annual nutrient budget in Bristol Bay lakes (Mathisen 1972, Koenings
and Burkett 1987, Schmidt et al. 1998), and salmon-derived nitrogen is ultimately taken up by
juvenile sockeye salmon (Kline et al. 1993). However, it is not clear if these nitrogen inputs
have measurable effects on sockeye salmon populations (Schindler et al. 2005b, Uchiyama et al.
2008).
41
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The importance of MDN to fish populations is perhaps most clearly demonstrated in
cases where MDN supplies are disrupted by depletion of salmon populations. The prolonged
depression of salmon stocks in the Columbia River basin is a prime example, where a chronic
nutrient deficiency hinders the recovery of endangered and threatened Pacific salmon stocks
(Gresh et al. 2000, Petrosky et al. 2001, Achord et al. 2003, Peery et al. 2003, Scheuerell et al.
2005, Zabel et al. 2006) and diminishes the potential of expensive habitat improvement
projects (Gresh et al. 2000). Density-dependent mortality has been documented among
juvenile Chinook, despite the fact that populations have been reduced to a fraction of historic
levels, suggesting that nutrient deficits have reduced the carrying capacity of spawning streams
in the Columbia River basin (Achord et al. 2003, Scheuerell et al. 2005). A population viability
analysis has indicated that declines in MDN have very likely contributed to low productivity of
juvenile salmon and that increasing the productivity could lead to large increases in the salmon
population (Zabel et al. 2006). Diminished salmon runs, thus, present a negative feedback loop
where the decline in spawner abundance reduces the capacity of streams to produce new
spawners (Levy 1997). Fisheries managers recognize the importance of MDN in sustaining the
productivity of salmon systems and are now attempting to supplement nutrient stores by
planting hatchery salmon carcasses and analogous fertilizers in waters throughout the Pacific
Northwest (Stockner 2003, Shaff and Compton 2009).
In addition to their inherent natural productivity, Bristol Bay watersheds have not been
subjected to anthropogenic watershed disturbances that have contributed to declining salmon
populations elsewhere. For example, Nehlsen et al. (1991) reviewed the status of native
salmon and steelhead stocks in California, Oregon, Washington, and Idaho. They found that
214 stocks appeared to face a risk of extinction; of these, habitat loss or modification was a
contributing factor for 194. These cases were in addition to at least 106 stocks that had already
gone extinct (Nehlsen et al. 1991). A National Research Council committee (NRC 1996),
convened to review the population status of Pacific Northwest salmon, summarized that:
The ecological fabric that once sustained enormous salmon populations has been
dramatically modified through heavy human exploitation - trapping, fishing,
grazing, logging, mining, damming of rivers, channelization of streams, ditching
and draining of wetlands, withdrawals of water for irrigation, conversions of
estuaries, modification of riparian systems and instream habitats, alterations to
water quality and flow regimes, urbanization, and other effects.
Thus, it is generally agreed that a complex and poorly understood combination of factors - with
direct and indirect effects of habitat degradation at the fore - are responsible for declining
Pacific Northwest salmon stocks (NRC 1996, Gregory and Bisson 1997, Lackey 2003).
In watersheds of the Bristol Bay region, including the Nushagak and Kvichak rivers,
human habitation is confined to a few small towns and villages, roads are few, and large-scale
habitat modifications are absent. The Riverscape Analysis Project, using spatial data from the
Socioeconomic Data and Applications Center (Sanderson et al. 2002), ranked 1574 salmon-
producing watersheds around the North Pacific based on an index of human footprint
(http://rap.ntsg.umt.edu/humanfootprintrank; accessed 9/1/11). Of these, the Kvichak River
ranked 197, the Nushagak (exclusive of the Wood River) ranked 131, and the Wood River
42
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ranked 332. Additionally, invasive fishes and riparian plants, which can negatively impact
native fish populations, have not been introduced to Bristol Bay's watersheds.
Habitat diversity
A diverse assemblage of spawning and rearing habitats is an exceedingly important
feature of Bristol Bay's riverine ecosystems. Since salmon adapt in predictable ways to
conditions within their specific environments, a high level of habitat diversity fosters a
correspondingly high level of population and life history diversity. The utilization of different
types of spawning habitat is an easily observable example. Suitable lotic habitats range from
small gravel-bed creeks to large cobble-bed rivers (Hilborn et al. 2003b), and even silt-laden
glacial streams (Ramstad et al. 2010). Spring-fed ponds are also used, as are areas of
groundwater upwelling on mainland lake beaches, and rocky beaches of low-lying islands
(Hilborn et al. 2003b). Sockeye salmon have adapted to each of these environments in
predictable ways, optimizing behavioral and physiological traits like timing of spawning, egg
size, and the size and shape of spawning adults (Table 9; Hilborn et al. 2003b). The result is a
stock complex comprised of hundreds of distinct spawning populations, each adapted to its
own spawning and rearing environment.
This complexity is compounded by variation within each spawning population, likely in
ways that are not yet fully understood (Hilborn et al. 2003b). One clear example is variation in
the amount of time spent rearing in fresh water and at sea (Table 10). Within a given cohort,
most individuals rear for either one or two years in fresh water, although a small number may
spend three years or go to sea shortly after hatching (i.e., zero years in fresh water). The latter
life history is relatively common among Nushagak River sockeye, many of which rear in rivers as
opposed to lakes. Once at sea, most fish will rear for an additional two or three years, although
a few will rear for as little as one year or as many as five years. This life history complexity
superimposed on localized adaptations results in a high degree of biological complexity within
the stock complex.
43
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Table 9. A summary of life history variation within the Bristol Bay stock complex of sockeye salmon (from Hilborn et al. 2003).
Element of biocomplexity
Range of traits or options found
Watershed location within Bristol Bay complex
Seven different major watersheds, ranging from maritime-influenced systems on the Alaska
Peninsula to more continental systems
Time of adult return to fresh water
June -September
Time of spawning
July- November
Spawning habitat
Major rivers, small streams, spring fed ponds, mainland beaches, island beaches
Body size and shape of adults
130 - 190 mm body depth at 450 mm male length: sleek, fusiform to very deep bodied, with
exaggerated humps and jaws
Egg size
88 - 166 mg at 450 mm female length
Energetic allocation within spawning period
Time spent rearing in fresh water
Time between entry into spawning habitat and death ranges from 1-3 days to several weeks
0-3 years
Time spent at sea
1-4 years
Table 10. Variation in time spent rearing in fresh water and at sea for Bristol Bay sockeye salmon. Numbers represent percentage of
fish returning to the respective river systems after a given combination of freshwater and sea rearing periods. + indicate
combinations that were represented in the data but comprised <1% of returns to the respective river system. Data are from ADF&G
and cover 1956 to 2005 brood years, except for Nushagak River data which cover 1979 to 2003 brood years.
Number of years spent in fresh water
Number of years spend at sea
Kvichak
Alagnak
Nushagak
Wood
Naknek
Egegik
Ugashik
Igushik
Togiak
0
3 4
1
17 2
1 1
25
42
11
48
16
9
27
20
21
10
40
60
43
44
17
28
68
63
+
+
5
+
+
+
+
+
+
58
12
1
5
17
44
30
5
6
7
5
2
3
21
29
15
5
7
3
2
44
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These layers of biocomplexity result in a situation where different stocks within the
complex show asynchronous patterns of productivity (Rogers and Schindler 2008). This is
because differences in habitat and life history lead to different population responses despite
exposure to the same prevailing environmental conditions. For example, a year with low
stream flows might negatively impact populations that spawn in small streams but not those
that spawn in lakes (Hilborn et al. 2003b). Asynchrony in population dynamics of Bristol Bay
sockeye has been demonstrated at both the local scale (i.e., individual tributaries) and the
regional scale (i.e., major river systems; Rogers and Schindler 2008). The latter is demonstrated
nicely by the relative productivity of Bristol Bay's major rivers during different climatic regimes
(Hilborn et al. 2003b), where small runs in the Egegik River were offset by large runs in the
Kvichak prior to 1977, but declining runs in the Kvichak River in the 2000s were in turn offset by
large runs in the Egegik River (see Figure 6).
Population and life history diversity within Bristol Bay sockeye populations can be
equated to spreading risk with a diversified portfolio of financial investments (Schindler et al.
2010). Under any given set of conditions, some assets perform well while others perform
poorly, but maintenance of a diversified portfolio stabilizes returns over time. Within the
sockeye stock complex, the portfolio of population and life history diversity greatly reduces
year-to-year variability in run size, making the commercial salmon fishery much more reliable
than it would be otherwise. With the current level of biocomplexity in Bristol Bay sockeye,
salmon runs are large enough to meet bay-wide escapement goals of ~10 M fish nearly every
year and fishery closures are rare (i.e., less than four closures per 100 years; Schindler et al.
2010). If Bristol Bay sockeye lacked biocomplexity and the associated stabilizing effects, run
sizes would fluctuate widely and complete fishery closures would happen every two to three
years (Schindler et al. 2010).
While the analyses described here apply to the Bristol Bay commercial sockeye fishery,
portfolio effects certainly stabilize populations of other fish species and increase the reliability
of sport and subsistence fisheries. In addition, portfolio effects stabilize and extend the
availability of salmon to consumers in the watershed food webs. Poor runs in some habitats
will be offset by large runs in others, allowing mobile predators and scavengers (e.g., bears,
eagles, rainbow trout) to access areas of relatively high spawner density each year (Schindler et
al. 2010). Different populations vary in the timing of spawning, which substantially extends the
period when salmon are occupying spawning habitats (Schindler et al. 2010).
Since a diversified salmon stock complex is contingent upon a complex suite of habitats,
an important question becomes: How does habitat diversity in Bristol Bay watersheds compare
to that in other salmon-producing regions? The Riverscape Analysis Project calculated
remotely-sensed indices of physical habitat complexity, allowing comparisons among salmon
producing watersheds at the North Pacific Rim scale (Luck et al. 2010, Whited et al. in press).
Rankings of overall physical complexity were based on 10 attributes: variation in elevation;
floodplain elevation; density of floodplains and stream junctions; human footprint; the
proportion of watershed covered by glaciers, floodplains, and lakes; and the elevation and
density of lakes. While the characterization of habitat complexity at this broad spatial scale is
necessarily imprecise and certainly fails to detect nuanced habitat features, it does seem to
quantify attributes that are important to salmon as it explained general patterns in salmon
abundance in validation watersheds (Luck et al. 2010). Overall physical complexity was
45
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relatively high for the watersheds considered in this assessment; of the 1574 Pacific Rim
watersheds characterized, the Kvichak River ranked the 3rd highest, the Nushagak River
(exclusive of the Wood River) ranked 44th, and the Wood River ranked 4th
(http://rap.ntsg.umt.edu/overallrank; accessed 9/1/11).
The studies reviewed here demonstrate how biocomplexity in salmon populations
provides resilience to environmental change. This resilience can break down when habitats are
degraded or when the genetic diversity that allows salmon to utilize the full complement of
available habitats is diminished. The loss of habitat diversity and associated loss of population
diversity has contributed to declines of once prolific salmon fisheries, including those in the
Sacramento (Lindley et al. 2009) and Columbia rivers (Bottom et al. 2005, Moore et al. 2010).
Lindley et al. (2009), summarizing causes for the recent crash in Sacramento River fall Chinook,
highlighted the importance of life history diversity:
In conclusion, the development of the Sacramento-San Joaquin watershed has greatly
simplified and truncated the once-diverse habitats that historically supported a highly
diverse assemblage of populations. The life history diversity of this historical assemblage
would have buffered the overall abundance of Chinook salmon in the Central Valley
under varying climate conditions.
46
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Whited, D. C., J. A. Lucotch, N. K. Maumenee, J. S. Kimball, and J. A. Stanford. In press. A riverscape
analysis tool developed to assist wild salmon conservation. Fisheries.
Williams, T. 2006. Pits in crown jewels. Fly Rod and Reel.
Willson, M. F. and K. C. Halupka. 1995. Anadromousfish as keystone species in vertebrate
communities Conservation Biology 9:489-497.
Wipfli, M. S., J. Hudson, and J. Caouette. 1998. Influence of salmon carcasses on stream productivity:
response of biofilm and benthic macroinvertebrates in southeastern Alaska, USA. Canadian
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Wipfli, M. S., J. P. Hudson, and J. P. Caouette. 2004. Restoring productivity of salmon-based food webs:
Contrasting effects of salmon carcass and salmon carcass analog additions on stream-resident
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Wipfli, M. S., J. P. Hudson, J. P. Caouette, and D. T. Chaloner. 2003. Marine subsidies in freshwater
ecosystems: Salmon carcasses increase the growth rates of stream-resident salmonids.
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54
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AK.
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55
-------
Appendix A. Chinook and sockeye salmon run sizes for Bristol Bay and other regions
of the North Pacific
Table Al. Chinook total run sizes by river system, 1966-2010
Table A2. Sockeye total run sizes by river system, 1956-2010
Table A3. Sockeye total run sizes by region, 1956-2005
56
-------
Table Al. Chinook total run sizes by river system, 1966-2010
Year
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
Nushagak
101,951
164,960
154,714
123,007
143,743
127,231
74,983
73,113
109,185
98,279
167,584
156,196
255,590
261,875
218,515
356,181
356,190
312,771
154,396
193,138
117,720
139,485
80,845
102,076
92,317
134,191
140,511
Yukon,
Kenai Canadian
mainstem
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
106,917
100,123
89,462
59,409
50,751
52,810
54,302
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
60,746
63,427
66,800
59,736
61,789
58,921
61,126
78,243
78,439
63,335
57,058
Copper
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
29,659
41,047
84,098
82,730
86,373
55,997
103,024
69,910
55,801
73,423
52,899
68,175
64,172
Taku
NA
NA
NA
NA
NA
NA
NA
38,307
35,442
46,870
44,555
41,856
56,386
60,190
64,247
75,280
37,042
19,943
41,850
71,814
51,190
41,474
66,601
57,086
66,517
80,066
84,882
Skeena
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
39,606
35,055
28,166
38,626
42,018
35,185
39,510
53,516
76,544
87,566
76,349
102,563
83,439
89,447
79,343
92,184
Nass
NA
NA
NA
NA
NA
NA
NA
NA
NA
17,874
16,583
18,410
21,807
16,229
18,744
17,606
13,287
20,516
31,408
24,768
47,967
26,568
21,094
36,594
33,384
13,136
25,405
Nehalem
NA
NA
NA
NA
NA
NA
NA
NA
NA
5,060
9,446
11,552
11,676
12,058
5,645
10,577
5,111
4,376
20,939
18,845
11,570
15,268
16,684
11,650
6,617
7,498
11,558
Skagit
NA
NA
NA
NA
NA
NA
NA
NA
NA
22,252
23,939
18,514
20,962
22,261
30,346
20,720
21,475
15,225
15,701
27,709
23,507
14,782
16,390
14,596
20,717
9,696
10,211
57
-------
Table Al. Chinook total run sizes by river system, 1966-2010
Year
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
Nushagak
175,376
228,739
177,509
136,663
155,562
242,184
79,168
75,074
113,367
133,552
136,008
227,994
244,082
218,005
123,139
126,700
115,559
95,592
Yukon,
Kenai Canadian
mainstem
89,748
90,552
81,563
77,228
69,773
55,540
86,553
63,373
60,320
61,878
73,210
99,765
91,309
76,186
76,472
61,152
46,095
NA
52,855
77,647
71,557
93,672
70,349
41,434
49,652
30,749
62,703
51,616
90,213
59,707
79,625
72,005
39,997
37,434
69,418
NA
Copper
65,301
90,073
96,710
113,868
107,760
112,365
95,951
70,746
81,155
72,972
94,505
80,559
66,341
99,877
87,770
53,880
43,007
32,999
Taku
98,073
70,253
74,564
98,184
130,091
51,706
33,500
51,055
59,449
71,902
62,436
113,923
81,173
68,842
29,766
126,700
115,559
NA
Skeena
96,018
68,127
48,351
96,453
65,350
65,167
70,993
77,320
112,346
63,069
82,410
61,065
39,278
43,689
44,185
54,279
55,921
NA
Nass
36,678
32,864
16,187
30,889
27,658
34,922
22,310
31,159
44,595
21,528
36,503
25,137
24,067
37,098
34,221
26,202
36,865
NA
Nehalem
9,137
9,194
8,671
12,975
12,732
10,591
10,361
10,817
14,293
20,552
23,569
14,456
8,222
13,129
6,648
5,651
5,332
NA
Skagit
7,691
7,082
10,096
13,364
7,198
16,067
5,725
18,231
15,947
20,979
11,933
25,863
24,701
23,115
13,003
15,942
13,144
NA
58
-------
Table Al. Chinook total run sizes by river system, 1966-2010
Year
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
Gray's
Harbor
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
6,852
10,086
7,919
10,869
17,067
10,581
9,886
8,473
23,888
14,225
25,139
35,114
42,811
57,787
40,606
34,569
34,813
Harrison
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
131,740
181,367
177,662
81,799
38,285
76,294
180,837
93,363
132,042
Fraser Yukon Kuskokwim
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
227,421
303,308
322,279
210,498
167,872
183,137
315,961
209,918
262,291
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
200,000
210,000
250,000
230,000
220,000
310,000
210,000
160,000
180,000
180,000
160,000
250,000
250,000
280,000
300,000
240,000
280,000
59
-------
Table Al. Chinook total run sizes by river system, 1966-2010
Year
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
Gray's
Harbor
31,513
32,468
34,067
39,102
35,927
23,390
14,865
18,595
22,405
19,787
24,945
48,690
26,365
27,230
17,976
19,149
14,493
NA
Harrison
120,600
100,839
29,840
38,568
72,061
189,103
107,884
78,098
74,419
91,122
251,453
138,890
92,993
52,798
83,445
43,798
75,550
NA
Fraser
230,837
246,142
164,318
224,127
274,856
358,436
248,823
233,307
251,427
312,142
483,142
333,330
265,274
295,676
220,651
231,389
248,408
NA
Yukon
NA
NA
NA
NA
NA
NA
NA
144,173
392,000
243,443
372,697
311,377
NA
NA
NA
NA
NA
NA
Kuskokwim
340,000
470,000
420,000
330,000
370,000
260,000
190,000
180,000
260,000
240,000
260,000
430,000
370,000
380,000
270,000
240,000
210,000
140,000
Data Sources
Nushagak: Pers. comm. Gregory Buck, ADF&G
Kenai: Begich and Pawluk 2010, pg. 69
Yukon, Canadian mainstem: Howard et al. 2009, pg. 35
Copper: Pers. comm. Steve Moffitt, ADF&G
Taku: McPherson et al. 2010, pg. 14; 2008/2009 data are preliminary pers. comm. Ed Jones, ADF&G
Skeena: PSC 2011, pg. 87
60
-------
Table Al. Chinook total run sizes by river system, 1966-2010
Nass: PSC 2011, pg. 87
Nehalem: PSC 2011, pg. 93
Skagit: PSC 2011, pg. 89
Gray's Harbor: PSC 2011, pg. 90
Harrison: PSC 2011, pg. 88
Fraser: PSC 2011, pg. 88
Yukon: Spencer et al. 2009, pg. 28
Kuskokwim: Pers. comm. Kevin Schaberg, ADF&G
61
-------
Table A2. Sockeye total run sizes by river system, 1956-2010
Year
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
Ugashik
779,000
940,000
776,702
678,064
3,377,000
960,000
559,409
673,000
1,101,179
2,236,533
1,315,949
449,557
179,413
372,879
1,030,000
1,790,000
129,031
60,108
65,801
464,000
594,000
325,175
95,380
2,158,312
4,469,800
3,705,000
2,603,342
4,565,269
Egegik
2,324,000
2,044,000
812,799
1,827,157
3,600,000
4,600,000
1,878,432
1,981,649
2,056,111
5,344,000
3,331,241
1,908,340
1,195,917
2,273,888
2,660,244
2,282,819
1,884,000
788,940
1,530,000
2,365,792
2,031,920
2,714,435
2,230,099
3,385,860
3,921,579
5,430,399
3,919,251
8,024,339
Naknek
3,155,000
2,588,000
603,781
3,403,474
2,095,000
1,865,815
1,277,933
1,786,728
2,685,504
2,270,357
2,418,111
1,372,352
2,119,324
2,623,702
2,011,095
3,247,238
1,810,000
724,941
1,728,781
3,804,529
2,619,548
2,744,790
2,005,239
2,292,995
5,027,516
7,913,237
4,226,271
5,754,315
Alagnak
1,282,000
474,000
206,930
1,295,000
2,289,000
509,000
150,000
368,227
554,998
506,729
354,000
298,956
302,531
329,748
479,019
599,080
235,000
53,833
236,681
128,700
152,000
177,471
1,178,690
1,562,870
1,594,128
862,018
2,173,398
1,531,412
Kvichak
13,800,000
10,711,000
1,180,705
1,004,118
24,942,000
14,279,000
4,961,330
657,349
1,801,221
47,657,000
9,064,868
5,577,403
3,471,140
13,472,862
34,599,600
6,948,068
1,763,000
336,241
4,761,892
15,359,808
3,789,238
2,266,442
8,266,273
25,297,982
37,695,437
7,489,183
3,328,986
20,983,178
Nushagak
106,788
262,805
543,003
113,107
237,544
185,798
114,209
452,272
244,344
513,460
402,292
114,332
290,366
197,135
885,640
662,007
99,603
428,733
240,197
1,071,353
1,079,065
946,903
1,482,163
930,285
5,343,159
3,764,287
2,889,822
2,073,502
Wood
1,494,000
945,000
1,744,000
3,668,000
2,124,466
957,144
2,438,322
1,460,090
2,263,164
1,468,609
2,310,435
1,017,456
1,357,407
1,218,238
2,169,211
1,912,659
935,000
716,226
2,211,000
1,836,317
1,602,770
928,878
4,294,726
3,775,140
4,760,312
4,926,000
3,864,630
4,484,000
Igushik
903,000
440,000
276,000
995,000
1,177,000
632,000
107,024
212,000
338,000
410,000
470,000
563,134
398,190
1,114,000
754,083
529,000
161,000
133,000
471,000
365,000
388,000
164,000
1,145,339
1,910,000
3,276,190
2,410,000
2,029,000
853,000
62
-------
Table A2. Sockeye total run sizes by river system, 1956-2010
Year
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
Ugashik
4,093,955
7,874,523
6,216,732
2,925,832
2,256,139
5,049,283
2,982,276
5,628,282
5,831,999
5,912,214
5,605,405
6,040,271
5,237,819
2,239,051
1,794,126
4,058,177
2,301,000
1,356,716
2,563,977
2,584,062
4,160,179
3,093,169
3,507,652
7,897,526
3,053,322
4,033,383
4,960,291
Egegik
6,623,390
9,093,576
6,173,448
6,884,561
8,369,057
10,983,145
12,931,258
9,938,166
18,614,125
24,481,560
12,998,886
16,200,980
12,253,942
9,362,876
5,060,215
9,407,420
8,403,612
4,323,287
5,839,236
3,503,084
12,865,161
9,553,946
9,066,558
8,209,756
9,027,266
13,039,645
6,119,472
Naknek
3,056,116
3,912,742
4,069,000
2,485,316
1,796,819
3,303,641
8,678,358
10,285,831
5,327,022
4,905,051
3,144,067
3,700,788
7,076,342
1,515,318
2,784,308
3,970,846
4,935,000
6,682,794
2,775,032
5,182,926
3,948,000
8,059,330
5,503,654
9,047,000
6,518,196
4,870,271
5,908,135
Alagnak
1,522,640
733,068
1,086,130
811,320
872,367
1,456,693
1,517,000
1,652,944
1,349,052
2,257,321
1,733,796
1,780,054
1,916,634
680,123
1,072,721
2,841,755
2,014,897
1,106,728
793,470
3,790,173
6,667,385
5,436,640
2,866,000
4,430,633
6,157,000
2,699,010
2,660,659
Kvichak
23,907,123
14,061,000
2,025,616
9,839,116
6,940,540
20,548,328
17,988,530
8,329,970
10,969,638
9,901,170
22,734,248
28,329,704
3,538,945
1,826,856
3,550,243
13,309,000
3,031,000
1,436,000
727,186
1,750,361
7,902,000
2,924,275
5,882,074
4,381,000
5,869,320
5,723,862
9,503,000
Nushagak
1,421,706
963,888
2,267,373
1,794,967
1,093,735
1,260,160
1,797,229
1,800,480
1,898,491
2,330,448
1,618,150
792,229
1,804,324
929,880
1,022,443
991,826
1,528,923
2,126,175
663,000
2,273,000
2,227,000
3,567,000
3,308,000
2,670,000
1,713,315
1,983,000
2,194,032
Wood
2,076,000
1,693,723
1,822,225
2,917,462
1,793,902
2,601,691
2,687,000
3,424,694
2,570,505
3,937,623
3,111,885
4,191,376
5,160,000
3,629,898
4,101,957
6,160,000
5,545,000
4,013,792
3,841,698
5,743,906
5,948,000
4,607,385
11,304,221
6,755,813
5,456,186
7,402,102
7,851,845
Igushik
455,000
489,000
908,000
644,000
414,000
1,253,000
1,317,000
2,515,000
830,000
1,663,194
1,379,000
1,991,000
1,514,000
314,000
602,074
1,626,000
1,812,000
1,325,000
213,000
1,036,071
523,000
2,089,000
1,466,000
1,826,000
3,433,000
953,000
1,391,576
63
-------
Table A2. Sockeye total run sizes by river system, 1956-2010
Year
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
Togiak
331,000
108,066
118,000
310,000
338,000
421,520
174,191
352,000
367,058
391,000
338,000
171,109
135,086
306,027
425,000
484,000
175,000
270,000
238,000
407,392
546,000
401,000
770,000
614,000
1,173,000
999,000
972,230
784,000
Kenai
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
831,241
920,826
435,344
485,352
1,374,607
2,268,568
2,096,341
797,838
1,495,962
1,184,445
2,766,912
3,982,112
Copper,
wild fish
NA
NA
NA
NA
NA
860,258
1,112,218
664,596
949,861
1,208,709
1,402,430
850,993
829,329
1,258,136
1,492,530
1,250,648
1,168,448
668,670
869,756
538,743
1,161,149
1,047,326
502,359
618,538
651,014
1,297,758
1,883,434
1,395,556
Fraser
2,866,977
5,401,219
18,778,820
4,769,576
3,421,281
4,713,837
3,512,304
3,985,486
1,824,500
3,166,871
5,459,849
6,803,585
2,955,662
4,941,025
6,163,676
7,696,359
3,708,113
6,878,291
8,616,165
3,683,576
4,340,815
5,887,114
9,420,144
6,358,912
3,133,187
7,741,247
13,985,095
5,240,936
64
-------
Table A2. Sockeye total run sizes by river system, 1956-2010
Year
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
Togiak
383,000
306,198
405,215
574,000
1,001,000
178,117
342,000
805,000
863,250
697,000
520,207
771,000
585,349
264,239
312,646
565,258
1,127,000
1,436,000
406,000
897,000
508,000
580,171
905,450
1,066,000
891,541
854,568
741,211
Kenai
1,287,187
2,498,144
2,955,276
9,425,518
6,094,157
6,662,137
3,290,388
2,226,730
8,273,968
4,451,954
3,908,776
2,658,341
3,743,751
4,650,889
1,953,963
3,018,164
1,842,904
2,214,605
3,511,797
4,447,000
5,716,924
6,117,166
2,835,742
3,592,167
2,065,205
2,440,138
3,595,867
Copper,
wild fish
1,821,370
1,600,390
1,329,070
1,721,153
985,913
1,435,481
1,459,380
1,766,134
1,537,006
2,039,851
1,839,406
1,778,450
2,888,442
3,820,171
1,661,543
1,568,335
1,206,275
2,000,609
1,774,724
1,839,605
1,739,197
2,060,867
2,305,355
2,828,457
1,051,154
1,583,006
1,248,019
Fraser
5,919,324
13,878,493
15,927,438
7,680,095
3,773,551
18,594,484
21,985,937
12,390,664
6,442,239
23,630,664
17,284,640
4,020,414
4,520,445
16,351,769
10,873,000
3,643,000
5,217,000
7,213,000
15,137,000
4,873,502
4,184,200
7,077,100
12,981,200
1,510,600
1,755,355
1,505,096
29,005,410
65
-------
Table A2. Sockeye total run sizes by river system, 1956-2010
Data Sources: Ugashik, Egegik, Naknek, Alagnak, Kvichak, Nushagak, Wood, Igushik, and Togiak rivers, pers. comm. Tim Baker, ADF&G; Kenai
River, pers. comm. Pat Shields, ADF&G; Copper River, pers. comm. Jeremy Botz, ADF&G; Fraser River, pers. comm. Catherine Michielsens, PSC.
66
-------
Table A3. Sockeye total run sizes by region, 1956-2005
Year
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
Bristol Bay
24,174,788
18,512,871
6,261,920
13,293,920
40,180,010
24,410,277
11,660,850
7,943,315
11,411,579
60,797,688
20,004,896
11,472,639
9,449,374
21,908,479
45,013,892
18,454,871
7,191,634
3,512,022
11,483,352
25,802,891
12,802,541
10,669,094
21,467,909
41,927,444
67,261,121
37,499,124
Russia
Mainland
and Islands
312,723
1,212,664
442,975
391,364
439,229
441,422
402,798
343,339
238,866
293,827
279,251
362,571
297,307
249,157
245,200
221,785
201,509
202,599
538,427
185,335
180,082
177,717
188,339
256,120
192,795
175,829
West
Kamchatka
5,568,959
10,172,076
6,286,252
5,046,656
5,520,707
8,884,293
8,304,347
5,294,022
1,681,381
3,616,954
2,496,149
3,438,364
952,912
705,033
1,051,653
1,908,446
1,708,238
1,266,604
2,914,942
1,315,733
1,556,672
412,752
936,931
835,766
1,353,186
1,641,425
East
Kamchatka
3,508,292
4,146,156
6,080,691
5,879,205
6,741,619
2,865,949
2,940,810
4,291,282
5,400,484
4,299,788
5,651,091
7,534,661
7,347,250
6,672,415
6,377,430
4,283,328
3,917,303
4,389,459
1,096,312
3,858,358
3,470,759
2,648,024
3,596,414
3,328,120
3,221,802
2,910,208
Western
Alaska
(excluding
Bristol Bay)
2,921,799
1,651,132
1,477,590
1,713,792
1,649,156
1,284,695
1,236,964
1,080,004
1,281,320
879,413
1,100,324
1,197,823
1,017,865
1,459,903
1,028,643
1,224,259
1,025,402
877,777
1,184,430
1,171,178
1,587,266
1,469,757
2,695,103
4,264,190
3,261,091
3,764,080
South
Alaska
Peninsula
1,439,813
823,438
654,585
837,418
1,301,201
728,145
856,552
936,188
918,361
1,136,937
816,878
1,022,036
1,771,470
997,774
2,477,613
2,224,301
996,272
1,745,569
1,515,481
1,048,430
2,219,569
3,082,269
2,547,058
1,855,669
1,534,564
3,009,576
Kodiak
1,036,251
976,164
1,064,076
1,134,597
1,189,167
1,265,417
1,870,103
1,263,847
1,415,449
1,161,768
1,630,675
1,098,764
1,832,648
1,566,384
2,071,227
1,382,529
957,567
880,634
1,283,380
854,537
1,586,702
1,645,986
1,925,502
1,745,390
2,235,004
1,977,914
Cook Inlet
2,107,703
1,272,942
1,026,900
1,227,947
1,663,849
1,982,278
1,962,984
1,690,524
1,727,099
2,304,205
2,849,643
2,263,184
1,906,856
1,341,961
1,399,803
1,262,215
1,604,503
1,310,905
1,056,869
1,331,877
2,619,311
3,194,737
3,250,421
1,626,406
2,485,427
2,266,861
Prince
William
Sound
1,357,869
1,219,564
795,032
767,304
921,272
1,246,740
1,446,375
965,103
1,413,881
1,631,195
1,867,747
1,119,440
1,334,651
1,728,312
2,007,971
1,362,728
1,671,399
986,426
1,361,911
1,092,387
1,713,575
1,629,798
1,026,705
798,885
553,557
1,396,065
67
-------
Table A3. Sockeye total run sizes by region, 1956-2005
Year
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
Russia
Bristol Bay Mainland
and Islands
26,006,930
49,053,015
43,538,930
39,127,718
24,973,739
28,876,574
24,537,559
46,634,058
50,240,651
44,380,367
48,254,082
56,085,581
52,845,644
63,797,402
39,087,355
20,762,241
20,300,733
42,930,282
30,698,432
23,806,492
17,822,599
26,760,583
44,748,725
39,910,916
256,135
272,271
188,414
129,556
177,623
173,853
134,865
162,907
131,959
278,341
290,791
414,830
330,884
547,226
578,622
273,153
186,020
314,421
402,372
458,915
254,755
189,284
92,408
681,161
West
Kamchatka
1,317,999
1,363,540
1,853,895
3,456,410
2,993,349
4,388,792
2,961,712
3,929,794
6,533,656
6,654,665
5,946,498
6,867,277
6,052,779
5,142,880
5,416,529
3,623,111
4,216,452
4,198,803
5,731,743
4,698,927
11,373,958
6,430,409
6,655,869
9,281,680
Western
East Alaska
Kamchatka (excluding
Bristol Bay)
2,495,343
3,255,333
2,869,830
2,266,824
2,088,398
2,244,085
1,735,950
1,614,359
683,440
716,325
2,171,680
3,721,809
3,184,687
5,342,393
5,181,509
4,525,486
3,350,431
4,688,991
3,228,330
3,295,161
1,969,758
3,111,533
2,370,070
3,082,258
1,960,326
2,962,209
2,854,259
5,074,028
3,648,527
1,881,441
2,428,248
2,984,749
4,066,861
4,709,511
4,550,924
5,252,589
4,707,327
5,231,199
3,904,663
3,327,626
2,342,865
3,551,763
3,417,071
2,741,406
2,750,691
2,998,568
3,968,890
5,282,123
South
Alaska
Peninsula
2,647,192
3,289,732
4,463,088
1,879,199
2,750,217
3,234,737
1,577,614
2,239,029
3,209,313
3,506,006
2,376,718
2,946,843
3,067,554
2,921,709
3,148,403
1,613,997
1,928,313
4,462,260
3,054,013
3,234,246
2,357,095
2,108,670
1,724,633
2,045,602
Kodiak
2,304,607
1,994,142
3,164,169
4,325,529
4,020,270
1,573,040
5,179,735
2,465,794
7,291,759
8,376,886
3,727,396
1,977,835
2,732,833
6,683,435
6,366,442
4,081,554
4,297,254
6,441,216
4,468,203
4,042,683
2,842,606
6,492,011
5,735,821
4,370,163
Cook Inlet
4,058,186
5,983,442
3,023,601
4,911,883
5,195,708
10,612,907
7,981,926
6,653,855
3,791,787
2,341,570
9,803,503
5,525,342
4,823,347
3,916,052
4,828,498
5,623,149
2,240,231
3,448,544
2,071,076
2,035,309
3,058,610
4,147,632
5,507,777
6,028,983
Prince
William
Sound
3,298,288
1,544,252
2,058,228
2,224,415
1,999,005
2,503,899
591,622
1,196,514
672,793
1,737,506
2,109,967
2,269,986
1,925,999
1,917,252
3,031,366
3,734,337
1,653,216
2,340,818
1,640,060
2,118,769
1,877,644
2,104,632
2,039,862
2,162,713
68
-------
Table A3. Sockeye total run sizes by region, 1956-2005
Year
1956
1957
1958
1959
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
Southeast
Alaska
1,223,955
1,433,321
1,348,999
1,191,656
787,118
996,105
1,033,237
907,045
1,236,191
1,452,134
1,410,391
1,299,903
1,111,561
1,085,977
893,721
833,222
714,626
907,999
1,010,069
924,210
1,638,128
2,040,197
1,480,429
1,927,777
1,506,153
North
British
Columbia
2,874,454
1,785,678
3,563,691
2,827,063
1,505,791
3,161,029
3,567,790
3,841,872
4,200,152
2,214,164
1,954,638
3,624,937
6,486,401
2,737,311
1,270,879
2,565,992
2,187,271
6,614,542
2,691,442
2,341,434
2,592,622
3,045,063
2,612,221
2,414,113
5,903,153
South
British
Columbia,
Washington,
and Oregon
3,724,473
5,923,358
22,137,627
5,976,277
4,497,613
5,430,221
4,092,561
4,991,161
2,315,203
3,698,689
6,316,328
8,400,670
3,609,851
5,809,127
7,194,502
9,733,215
4,565,063
8,336,516
10,137,727
4,472,874
5,296,487
8,025,282
10,353,993
8,310,609
5,106,260
69
-------
Table A3. Sockeye total run sizes by region, 1956-2005
Year
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
Southeast
Alaska
1,484,281
1,951,773
1,803,879
1,641,315
2,133,525
1,596,155
1,755,611
1,332,203
2,022,589
2,041,318
2,001,214
2,493,953
3,183,080
2,052,188
1,625,062
3,066,710
2,232,489
1,351,217
1,569,562
1,255,042
1,827,078
1,537,801
1,670,133
1,915,752
1,693,703
North
British
Columbia
9,878,197
7,676,011
4,742,841
4,030,945
8,899,568
5,738,111
5,591,872
7,076,794
4,706,414
5,204,017
7,068,326
8,841,375
8,529,952
4,533,119
7,471,188
9,353,278
5,836,899
2,339,626
2,145,620
5,784,376
5,418,729
3,512,452
4,119,532
2,661,373
1,709,492
South
British
Columbia,
Washington,
and Oregon
9,518,792
15,580,715
7,330,812
8,240,361
15,583,867
16,389,443
9,113,405
5,538,086
19,501,105
22,849,561
14,639,516
8,320,825
25,605,669
18,058,968
4,253,526
5,386,660
17,469,309
11,600,660
4,283,929
6,008,081
8,409,348
12,222,016
5,028,196
3,501,674
3,827,344
70
-------
Table A3. Sockeye total run sizes by region, 1956-2005
Data Sources: Bristol Bay, pers. comm. Tim Baker, ADF&G; Other regions are from Ruggerone et al. 2010
71
-------
Appendix B
Characterizations of Selected
Non-Salmon Fishes Harvested in the
Fresh Waters of Bristol Bay
B-l
-------
Characterizations of selected non-salmon fishes
harvested in the fresh waters of Bristol Bay
Michael Wiedmer
Malma Consulting
2500 Susitna Drive
Anchorage, AK 99517
mikewiedmer@acsalaska.net
907.243.7005
- 1 -
-------
CHARACTERIZATIONS OF SELECTED NON-SALMON FISHES
HARVESTED IN THE FRESH WATERS OF BRISTOL BAY
INTRODUCTION
The fresh waters of the EPA Bristol Bay Watershed Assessment Area (BBWAA) in southwest
Alaska support diverse and robust populations of at least 11 families, 22 genera, and 35 species
offish (Table 1). This appendix provides biological, ecological, and human use information for
selected species which are, or have been, targeted by sport, subsistence, and/or commercial
fisheries within the BBWAA. This appendix does not review Pacific herring, Pacific cod, saffron
cod, Pacific staghorn sculpin, Arctic flounder, and starry flounder that are primarily marine
species that only venture intermittently into the lower reaches of some of the drainages
(Mecklenburg et al. 2002; Morrow 1980b). This appendix also does not describe species
harvested in fresh water for which there is limited BBWAA distributional or biological
information (e.g., boreal smelt), nor the five species of North American Pacific salmon, which
are reviewed in Appendix A of this assessment.
Each of the species described in this appendix: northern pike, humpback whitefish, rainbow
trout, Arctic char, Dolly Varden, lake trout, and Arctic grayling, are well distributed within the
BBWAA. Unlike the obligate anadromous BBWAA Pacific salmon populations, in which
essentially all individuals migrate from lakes and streams to the sea to feed and grow, individual
fish in these seven species do not need to journey to marine waters to successfully complete their
life cycle, although some individuals of certain species (e.g., Dolly Varden and humpback
whitefish) may. Anadromous fish spawn in fresh waters but feed for part of their lives in marine
waters (Myers 1949). Nonanadromous (resident) fish remain throughout their lives in fresh
water, but may move seasonally between habitats within a given drainage (see species
descriptions below). Also unlike the North American Pacific salmon, individuals in each of these
seven species can survive to spawn more than once (they are iteroparous, Stearns 1992, p. 180)
and, compared to salmon, have longer potential life spans (see species descriptions below).
Northern pike Esoxlucius
Freshwater distribution and habitats
Northern pike have a circumpolar distribution across the northern hemisphere and is the only
species in the family Esocidae that has colonized arctic waters (Grossman 1978). In North
America they inhabit lakes and low gradient rivers from the Arctic Ocean south to the Missouri
and Mississippi river drainages, and from the North Atlantic Ocean west to the Rocky Mountains
(Scott and Grossman 1998, p. 357). In Alaska, northern pike are native primarily north of the
Alaska Range, including waters of the BBWAA (Mecklenburg et al. 2002, p. 144; Morrow
1980b, p. 168). In Bristol Bay, northern pike occur in coastal plain lakes (Hildreth 2008, p. 9),
inland lakes (Burgner et al. 1965, p. 4; Dye et al. 2002, p. 1; Russell 1980, p. 87), and river
systems (ADF&G 2011) providing suitable habitat. The Nushagak and Nuyakuk river
mainstems, Lake Aleknagik, and the Lake Clark drainage support the largest sport fisheries
within the BBWAA (Jennings et al. 2011, p. 126, 128).
Northern pike primarily spawn in sections of lakes, wetlands, or very low gradient streams
providing shallow (less than 1 m), slow or still waters with soft substrates and aquatic vegetation
-2-
-------
(Cheney 1971d, p. 13; Chihuly 1979, p. 48, 57; Dye et al. 2002, p. 5, 6-7; Rutz 1999, p. 15).
Summer habitat is in slightly deeper, but still warm water with dense aquatic vegetation (Chihuly
1979, p. 46, 58; Dye et al. 2002, p. 5; Joy and Burr 2004, p. 22; Roach 1998, p. 3; Rutz 1999, p.
9). In southcentral Alaska's Susitna River drainage, river-dwelling northern pike are often found
in side sloughs where water temperatures are several degrees warmer than the adjacent main
channel (Rutz 1999, p. 19). Among the large, deep, cold, glacially-formed lakes of the BBWAA,
shallow, vegetated habitats are scarce, making those found in Lake Clark's Chulitna Bay and the
shallow bays of Lake Aleknagik particularly important northern pike concentration areas
(Chihuly 1979, p. 48; Dye et al. 2002, p. 6-7; Russell 1980, p. 91).
Northern pike overwinter in lakes, spring-fed rivers, or larger deep rivers where there is likely to
be sufficient water and oxygen to survive until spring (Dye et al. 2002, p. 5; Roach 1998, p. 18-
21; Scanlon 2009, p. 17; Taube and Lubinski 1996, p. 5-8). Water depth beneath winter ice may
be 0.8 m or less (Taube and Lubinski 1996, p. 8). In winter, local residents ice fish for northern
pike along the BBWAA's large rivers (Krieg et al. 2009, p. 135, 220, 215, 344).
Life cycle
At spring ice-out in the BBWAA's Lake Aleknagik, large fish are in water 1 to 1.5 m deep and
within 10 m of shore. In late May to mid-June, as water temperatures rise to about 6 °C, mature
fish move inshore to spawn in brush and aquatic vegetation (Dye et al. 2002. p. 5). Female
northern pike can produce over 100,000 adhesive 3-mm diameter ova, which they scatter in
small batches among aquatic vegetation or rocks, while an attending male fertilizes them.
Neither females nor males construct redds (Morrow 1980b, p. 166-167; Scott and Grossman
1998, p. 359). After spawning, as Lake Aleknagik water temperatures rise above 8 °C, fish move
slightly offshore, to 1 to 3 m of water, but remain in the bays where they spawned, moving little
for the remainder of the summer (Dye et al. 2002, p. 5). As water levels and temperatures drop in
mid-September through October, fish move out of shallow bays to depths of 3 to 5 m in the main
lake and then move little until the following spring (Dye et al. 2002, p. 5).
Mature northern pike living in Alaskan river systems and river-lake complexes ascend tributaries
in spring, beneath the ice. Spawning occurs from mid-May to early July as ice melts in side-
channel slack waters or lake margins. After spawning, mature pike move to deeper water to feed,
where they remain until moving in September and October to lakes, spring-fed streams, and
larger, deeper rivers where they overwinter (Cheney 1971d, p. 13-14; Cheney 1972, p. 5;
Chythlook and Burr 2002, p. 13; Kepler 1973, p. 75; Russell 1980, p. 91; Taube and Lubinski
1996, p. 6-8).
Northern pike eggs hatch in less than a month. At hatching, fry are 6 to 9 mm long, and have a
yolk sac, but no mouth. Before they start actively feeding, fry cling to the substrate, debris, or
vegetation for around 10 days, absorbing their yolk sacs while their mouths develop (Morrow
1980b, p. 167; Scott and Crossman 1998, p. 359). In BBWAA lakes, young-of-the-year northern
pike are actively swimming by at least late June to early July and grow rapidly through the
summer (Chihuly 1979, p. 32, 34; Russell 1980, p. 91, 93). In river systems, fry remain near or
downstream of spawning areas (Cheney 197Id, p. 13). In interior Alaska, age-0 fish reach a
mean length of 140 mm by September (Cheney 1972, p. 15). In Lake Aleknagik, northern pike
grow rapidly to about age 4 and a total length of around 419 mm, then growth slows to about an
average of 25 mm per year (Chihuly 1979, p. 27-28, 33). Some male northern pike in Lake
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Aleknagik mature at age 3, and by around age 5 and lengths of approximately 438 to 469 mm, all
fish are mature (Chihuly 1979, p. 34).
Many mature northern pike do not travel far (Chihuly 1979, p. 64; Dye et al. 2002, p. 5; Joy and
Burr 2004, p. 25; Rutz 1999, p. 8), but some river-system individuals make extensive seasonal
migrations between spawning, feeding, and overwintering areas (Scanlon 2009, p. 11),
sometimes moving at least 290 km per year (180 mi per year, Cheney 1971a, p. 7). Mature
northern pike may disperse through the summer and then aggregate prior to moving to
overwintering locations and while overwintering (Roach 1998, p. 14). Mature northern pike
show high fidelity to spawning (Joy and Burr 2004, p. 29; Roach 1998, p. 13) and winter areas
(Scanlon 2009, p. 20; Taube and Lubinski 1996, p. 8) and moderate fidelity to summer feeding
areas (Taube and Lubinski 1996, p. 8). Because fish must exceed a minimum size before they
can be successfully tracked with standard telemetry methods, most movement studies are limited
to bigger individuals and seasonal movements of immature Alaskan northern pike are largely
unknown.
Mature females often tend to be larger than males of the same age (Clark et al. 1988, p. 22, 25;
Pearse 1991, p. 36; Rutz 1999, p. 9), but males appear to have a greater mortality rate (Cheney
1971c, p. 17; Chihuly 1979, p. 26; Pearse 1991, p. 36). In the BBWAA, northern pike can reach
total lengths of at least 1.04 m, weights in excess of 7 kg, and ages of 18 years (Chihuly 1979, p.
33, 37; Dye et al. 2002, p. 6; Russell 1980, p. 92, 93). In the Yukon River drainage, fish can
reach 1.2 m in length (Scanlon 2009, p. 20), and 26 years in age (Cheney 197lc, p. 15).
Predator-prey relationships
Northern pike are highly adaptable predators able to consume a wide range of invertebrates and
vertebrates, but they are particularly efficient consumers of fish (Craig 2008). Where they are
available, a wide variety of fish dominate the diet of larger BBWAA northern pike, including
Alaska blackfish, round whitefish, least cisco, smaller northern pike, ninespine and threespine
stickleback, juvenile sockeye salmon, Arctic char, pygmy whitefish, sculpins, longnose suckers,
and lake trout (Chihuly 1979, p. 79-86; Russell 1980, p. 95-97). The diet of larger northern pike
illegally introduced into southcentral Alaska's Susitna River drainage was dominated by coho
and sockeye salmon, whitefish species, stickleback species, and rainbow trout (Rutz 1999, p. 17).
Immediately after hatching, young-of-the-year fry eat zooplankton and immature aquatic insects,
but quickly transition to small sticklebacks and other small fish (Chihuly 1979, p. 85-88; Morrow
1980b, p. 167). Northern pike smaller than 200 mm feed substantially on invertebrates; fish over
400 mm eat invertebrates (e.g., crustaceans, leeches, beetle larvae, and mollusks, Russell 1980,
p. 95-97) only incidentally (Cheney 1972, p. 29; Chihuly 1979, p. 79-88). Northern pike diets are
adaptable and can include a wide variety of foods in the absence of fish prey, although growth
rates are then lower (Cheney 1971b, p. 23). Northern pike are a keystone predator and often the
greatest predator of northern pike are larger northern pike (Cheney 1972, p. 27; Chihuly 1979, p.
82; Craig 2008).
Abundance and harvest
Total abundance of northern pike in the BBWAA is unknown. Dye et al. (2002, p. 6) estimated
that in 1998 and 1999, the abundance of northern pike longer than 299 mm in Lake Aleknagik
was more than 11,580. Chulitna Bay on Lake Clark has supported a large subsistence fishery; in
June 1978 an estimated 350 to 500 large northern pike were harvested from Turner Bay at the
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head of Chulitna Bay (Russell 1980, p. 91). In the mid-2000s, residents in ten of the BBWAA
villages annually harvested an estimated 4,385 northern pike (Fall et al. 2006, p. 45, 80, 113,
150, 194; Krieg et al. 2009, p. 40, 78, 118, 162, 202), and they were the most important non-
salmon fish in four of those villages (Fall et al. 2006, p. 152; Krieg et al. 2009, p. 46, 124, 171).
From the mid-1970s to the mid-2000s, northern pike were estimated to represent between 9.9
and 14.1% of the total weight of the Kvichak River drainage non-salmon freshwater fish
subsistence harvest (Krieg et al. 2005, p. 214). In 2009, sport anglers caught an estimated 8,217
northern pike in the BBWAA and the adjacent Togiak River drainage (10% of the statewide
total) and harvested (kept) an estimated 1,177 (6% of the statewide total; Jennings et al. 2011, p.
75). Annual sport harvests have declined, due at least in part to both lower bag limits and the
increasing popularity of catch-and-release fishing (Dye and Schwanke 2009, p. 6). In 1966 and
1967, an experimental freshwater commercial fishery on Tikchik Lake harvested 316 northern
pike, the third-most commonly harvested fish (6% of total number offish harvested; Yanagawa
1967, p. 10).
Stressors
Because northern pike are long-lived, have a piscivorous diet, and prefer relatively warm water,
they bioaccumulate and biomagnify atmospherically deposited mercury, and tissue mercury
concentrations correlate strongly with length and age (Headlee 1996; Mueller et al. 1996, p. 36).
Lindesjoo and Thulin (1992) reported that wild northern pike exposed to pulp mill effluents
developed severe jaw deformities. They did not determine if the deformities were directly caused
by constituents of the effluents, if the deformities resulted from a secondary reduction of
dissolved oxygen (DO) levels, or through some other mechanism. Northern pike are highly
tolerant of low DO levels. In laboratory experiments, juvenile northern pike survived DO levels
down to at least 0.25 mgT1 (Petrosky and Magnuson 1973).
Casselman (1978) found that, for a Canadian stock of northern pike, maximum summer growth
occurred at 19 °C, growth stopped at 28 °C, and 29.4 °C was the upper incipient lethal
temperature. For an Ohio stock, Bevelhimer et al. (1985) reported maximum summer growth
occurred at 25 °C and that northern pike continued to grow at 30 °C. Combined, these results
suggest a possible latitudinal cline in temperature tolerances and optimal and lethal temperatures
for BBWAA northern pike may be lower than those reported by Casselman (1978).
Humpback whitefish C Oregon us pidschian
The taxonomic status of humpback whitefish remains unsettled. Some sources (e.g.,
Mecklenburg et al. 2002, p. 180; Morrow 1980b, p. 24) distinguish three separate Alaskan
whitefish species (lake C. clupectformis, Alaska C. nelsonii, and humpback C. pidschian) based
on gill raker counts; other authors (e.g., Alt 1979; Brown 2006, p. 2; McDermid et al. 2007)
consider them a single variable species (the C. clupeaformis complex). This appendix treats the
three forms synonymously. In addition, Bernatchez and Dodson (1994) suggest that this species
should be considered synonymous with the European whitefish C. lavaretus.
Freshwater distribution and habitats
In combination with the European whitefish, the humpback whitefish has a circumpolar
distribution across the northern hemisphere (Bernatchez and Dodson 1994). In North America,
the humpback whitefish freshwater range extends from the Arctic Ocean coastal plain south to
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approximately the southern border of Canada, and from the Atlantic seaboard to the Bering Strait
(Scott and Grossman 1998, p. 271). Humpback whitefish are found in lakes, streams, and
brackish water across much of Alaska, primarily north of the Alaska Range (Alt 1979;
Mecklenburg et al. 2002, p. 186-188). In the BBWAA, humpback whitefish are reported in
deeper lakes, mainstem rivers, and slow-flowing tributaries (ADF&G 2011; Burgner et al. 1965,
p. 4, 5; Fall et al. 2006, p. 321, 337, 354, 381; Krieg et al. 2009, p. 301, 318, 339, 365, 370;
Metsker 1967, p. 6; Russell 1980, p. 72-76; Woody and Young 2007, p. 8; Yanagawa 1967, p.
12).
In northwest Ontario, lake spawning sites were found in nearshore areas at average depths of 2.7
to 3.5 m; primarily over boulders, cobbles, and detritus (Anras et al. 1999). In interior Alaska,
stream spawning sites are in spatially discrete reaches, often glacially-fed, with moderate to high
gradients, moderate to swift currents, and gravel substrates (Alt 1979; Brown 2006, p. 25-26;
Kepler 1973, p. 71). In interior Alaska's Chatanika River, fish spawn in water 1.3 to 2.6 m deep,
flowing at approximately 0.5 nvs"1 (Kepler 1973, p . 71).
After spawning, adults migrate downstream to more slowly flowing waters with fine substrates
(Brown 2006, p. 26). In Canada's Mackenzie River system, overwintering locations are in deep
mainstem channels or delta areas (Reist and Bond 1988). Lakes supporting summer feeding
aggregations in interior Alaska are well connected to mainstem channels, ensuring that feeding
fish can reliably enter in spring and exit in late summer to migrate to spawning and
overwintering areas (Brown 2006, p. 31).
In early August, apparently mature fish were collected in the lower Swan River, about 2 km
upstream of the confluence with the Koktuli River (ADF&G 2011, sites FSN0604A02,
FSN0604A04), and mature fish were collected at the mouth of Koggiling Creek, at its confluence
with the lower Nushagak River (ADF&G 2011, sites FSN0607C08, FSN0607C10). The
stomachs of most of the Koggiling Creek fish were empty (Wiedmer unpublished). These fish
may have recently left summer feeding lakes in the Swan River and Koggiling Creek drainages
and were staging before beginning their upstream spawning migration (see Life cycle and
Predator-prey discussions below).
In late August, apparently mature and perhaps larger immature fish were collected in small
upland lakes draining to the upper North Fork Koktuli River (ADF&G 2011, sites
PEB91NK011, PEB91NK019). Whether these fish overwinter in these lakes is not known. In
fall, BBWAA residents harvest humpback whitefish in mainstem rivers, as the whitefish move
upstream to spawn. In winter, residents also harvest humpback whitefish in Sixmile and Iliamna
lakes, Lake Clark, and mainstem rivers (Fall et al. 2006, p. 39, 200, 289, 321, 337, 354, 381;
Krieg et al. 2009, p. 55, 135, 159, 178, 220, 301, 339, 365).
In Alaska, the habitat preferences of juvenile humpback whitefish have been particularly difficult
to define (Brown 2004, p. 19; Brown 2006, p. 25, 30; Brown et al. 2002, p. 18). In the lower
Mackenzie River, nursery habitats and foraging areas for young-of-the-year are in delta lakes and
main delta channels (Chang-Kue and Jessop 1992, p. 27). No young-of-the-year were found in
main-channel rivers and streams in the Nushagak River drainage in August 2006 (ADF&G
2011), suggesting either a year-class failure (Bogdanov et al. 1992) or that they were occupying
off-channel habitats. In Lake Clark and adjacent lakes, juveniles were captured mostly in shallow
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(less than 3 m) nearshore areas, while larger fish were more broadly distributed (Woody and
Young 2007, p. 8).
Life cycle
North of the BBWAA, some humpback whitefish populations include anadromous individuals,
but the proportion of anadromy within stocks appears to decrease with increasing distance from
marine waters (Brown 2004, p. 17; Brown 2006, p. 14; Sundet and Pechek 1985, p. 34). Within
the BBWAA, limited otolith isotope analyses have yet to reveal evidence for anadromy in fish
collected in Lake Clark or the lower Nushagak River (Randy Brown, U. S. Fish and Wildlife
Service, Fairbanks, personal communication; Woody and Young 2007, p. 12).
In interior Alaska, large fish feed in lakes until late summer. They then move into mainstem
rivers and stay near lake outlets for up to 3 weeks before beginning to migrate upstream to
spawning areas in late August to early September. Most spawners arrive in the spawning areas
by mid-September, and spawning extends from late September to mid-October (Brown 2006, p.
26). Russell (1980, p. 72) reported spawning in late September in BBWAA lakes. Lake spawning
in northwest Ontario occurs at temperatures between 2 and 6 °C, shortly before lake surfaces
begin to freeze (Anras et al. 1999). Kepler (1973, p. 71) reported spawning in an interior Alaskan
stream from mid-September to early October, at temperatures ranging from 0 to 3 °C.
In interior Alaska, males mature at ages 4 to 6; females at ages 5 to 7 (Alt 1979; Brown 2006, p.
28). Fish are reported to mature at lengths of about 310 to 380 mm (FL; Alt 1979; Brown 2004,
p. 19; Brown 2006, p. 23; Chang-Kue and Jessop 1992, p. 17; Kepler 1973, p. 71), and age and
length at maturity may vary among locations (Alt 1979; Brown 2004, p. 19; Chang-Kue and
Jessop 1992, p. 17). Three females from the lower Nushagak River (ADF&G 2011, sites
FSN0607C08, FSN0607C10), had fork lengths of 435 to 460 mm, were mature, while one 370
mm female was not (Wiedmer unpublished). In interior Alaska, females apparently spawn every
year (Brown 2006, p. 29). Farther north, at least some females do not spawn every year, although
males may (Brown 2004, p. 16, 17).
Humpback whitefish broadcast spawn instead of digging redds; after fertilization their 2- to 3-
mm diameter eggs sink and lodge in the interstitial spaces of the substrate (Anras et al. 1999;
Morrow 1980b, p. 36, 38; Scott and Grossman 1998, p. 271). Fecundity of interior Alaska
humpback whitefish ranges from 8,400 to 65,400 ova for females ranging in length from 320 to
520 mm (Clark and Bernard 1992). The estimated fecundity of three mature females collected in
August in the mouth of Koggiling Creek in the BBWAA (ADF&G 2011, sites FSN0607C08,
FSN0607C10) fell within this range (Wiedmer unpublished).
In Siberian rivers, the time from spawning to hatching is about 185 to 190 days and survival
from egg to fry appears to vary greatly from year to year (Bogdanov et al. 1992). Larval fish,
weighing 4.9 to 6.3 mg, with lengths of 9 to 13 mm, drift downstream immediately after hatching
(Bogdanov et al. 1992; Shestakov 1991). Studies in both Norway and Siberia found that these fry
still have yolk sacs and do not begin feeding for the first several days of their downstream drift
(Naesje et al. 1986; Shestakov 1991). In Siberia's Anadyr River, larvae drift downstream for two
to three weeks, from late May to early June (Shestakov 1991; Shestakov 1992). The scale and
speed of downstream migrations correlate with increases in river discharge (Bogdanov et al.
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1992; Naesje et al. 1986; Shestakov 1991). Russell (1980, p. 72) observed fry in the shallows of
BBWAA lakes by mid-June.
In interior Alaska and northern Canada, immature fish, from age 0 to about age 4, appear to rear
far downstream of spawning areas in off-channel sites such as deltas, lakes, and sloughs, or in
mainstem eddies (Brown 2006, p. 31; Reist and Bond 1988). Age-0 juveniles in the Anadyr
River primarily inhabit lakes that connect to the mainstem during spring high flows (Shestakov
1992). By mid-July, age-0 fish reach 43 mm, with growth faster in floodplain lakes than in
streams (Shestakov 1992).
In the BBWAA, humpback whitefish reach at least age 27 and lengths to 584 mm. (Woody and
Young 2007, p. 8). Elsewhere, maximum age can be up to 57 years (Power 1978). In interior
Alaska, maturing and mature fish show fidelity to both summer feeding (Brown 2006, p. 21;
Brown et al. 2002, p. 16), and fall spawning areas, which can be more than 300 km apart (Brown
2006, p. 22, 31).
Predator-prey relationships
Large humpback whitefish from BBWAA lakes feed predominantly on benthic invertebrates,
particularly mollusks, chironomids (non-biting midges), planktonic crustaceans, and caddis fly
larvae (Metsker 1967, p. 29; Russell 1980, p. 76), but apparently feed on salmon eggs and small
fry when available (Van Whye and Peck 1968, p. 37; Woody and Young 2007, p. 13). Adults
preparing to spawn stop eating earlier than mature non-spawners, and large humpback whitefish
feed little during the spawning migration and while overwintering (Brown 2004, p. 21; Brown et
al. 2002, p. 16). In lakes, young-of-the-year fry initially feed primarily on planktonic crustaceans
(Claramunt et al. 2010; Hoyle et al. 2011). When they reach lengths greater than 40 mm, their
diet transitions to benthic macroinvertebrates, particularly chironomids (Claramunt et al. 2010).
Round whitefish and Arctic grayling feed on humpback whitefish eggs (Brown 2006, p. 23;
Kepler 1973, p. 71), and other species likely do as well. Humpback whitefish are vulnerable to
predation by piscivorous fish, such as lake trout (Van Whye and Peck 1968, p. 37) and in the
BBWAA, northern pike may be important predators (Russell 1980, p. 95).
Abundance and harvest
The total abundance of humpback whitefish in the BBWAA is not known. The estimated mid-
20008 annual subsistence harvests in nine of the villages within the BBWAA totaled over 4,000
fish (Fall et al. 2006, p. 45, 80, 113, 150, 194; Krieg et al. 2009, p. 40, 78, 118, 162, 202). From
the mid-1970s to the mid-2000s, whitefish, the majority of which are humpback whitefish, were
estimated to represent between 8.3 to 26.8% of the total weight of the Kvichak River drainage
non-salmon freshwater fish subsistence harvest (Krieg et al. 2005, p. 214).
The 2009 estimated sport catch of all whitefish species in the BBWAA plus the Togiak River
drainage was 1,118 fish (11% of the total statewide catch of all whitefish species except sheefish
Stenodus leucichthys), and the estimated harvest was 520 (18% of the total statewide harvest of
all whitefish species, except sheefish; Jennings et al. 2011, p. 76). In the mid-1960s, Iliamna
Lake and Lake Clark supported a commercial humpback whitefish fishery (Metsker 1967, p. 8,
10). In 1966 and 1967, humpback whitefish comprised 62% of the total number offish harvested
in a freshwater commercial fishery on Tikchik Lake (Yanagawa 1967, p. 12).
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Stressors
Mature humpback whitefish aggregate in discrete spawning habitats, leaving them at risk to both
acute events during fall spawning and chronic changes to spawning habitat (Brown 2006, p. 32).
Extreme high water events shortly before fall spawning may cause adult whitefish to leave
spawning areas and delay spawning to another year (Underwood et al. 1998, p. 13). The
spawning success of lake-dwelling whitefish is vulnerable to lake level manipulation during the
winter incubating period (Anras et al. 1999) and to elevated substrate sedimentation (Fudge and
Bodaly 1984). Age-0 fish are vulnerable to low flows in spring, which can prevent access to
preferred floodplain lake habitats (Shestakov 1992).
Mature humpback whitefish appear not to feed during spawning migrations or during the winter
(Brown 2004, p. 21; Brown et al. 2002, p. 16). Almost all annual feeding occurs in summer,
often in off-channel lakes. Mature whitefish must have access to and from these lakes, both in
spring to immigrate and in late summer to emigrate (Brown 2006, p. 26).
Fertilized eggs need cold water (optimally around 0.5 °C; Morrow 1980b) during development;
eggs incubating in 10 °C waters suffer 99% mortality rates (Scott and Grossman 1998, p. 272). In
an experiment mimicking Great Lakes summer conditions, Edsall (1999) found juvenile survival
peaked at water temperatures of 10 to 15 °C and declined at lower and warmer temperatures and
that juvenile growth peaked at 18.5 °C. For Great Lakes young-of-the-year acclimated to warmer
waters, the upper lethal temperature was 26.6 °C (Edsall and Rottiers 1976). Metabolically,
whitefish do not swim as efficiently as other salmonids (Bernatchez and Dodson 1985).
Swimming performance peaks at around 12 °C and declines at lower temperatures. Bernatchez
and Dodson (1985) speculate that the timing of seasonal migrations may be a function of the
combined influence of seasonal stream velocities and temperatures. Optimal and lethal
temperatures may be lower for Alaskan populations.
Rainbow trout Oncorhynchusmykiss
Rainbow trout and steelhead are two forms of one species and belong to the same genus
(Oncorhynchus) as the Pacific salmon. Rainbow trout is the common name for individuals with
nonanadromous life histories and steelhead is the common name for individuals with
anadromous life histories. Unlike the Pacific salmon, southwest Alaska rainbow trout/steelhead
are mostly nonanadromous. In Bristol Bay, the Alaska Department of Fish and Game (ADF&G)
documents steelhead only in a few spawning streams near Port Moller, in the southwestern
portion of the basin, outside the BBWAA (Johnson and Blanche 2011, Chignik and Port Moller
1:250,000 quadrangles). As no steelhead are known to occur in the fresh waters of the BBWAA
(e.g., Russell 1977, p. 44), they are not discussed further here.
Freshwater distribution and habitats
The native freshwater range of rainbow trout is largely restricted to Pacific Ocean drainages: in
North America from the Kuskokwim River system in Alaska south to mountain drainages of
central Mexico (MacCrimmon 1971, p. 664), and in Asia in the Kamchatka region (Froese and
Pauly 2011). Native rainbow trout in Alaskan fresh waters are restricted to southwest,
southcentral, and southeast Alaska, from the Holitna River region south to Dixon Entrance
(Morrow 1980b, p. 78). Rainbow trout have been extensively and successfully transplanted
outside their native range, including sites in interior Alaska (MacCrimmon 1971; Morrow 1980b,
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p. 51). While rainbow trout of the Nushagak and Kvichak river drainages are near the northern
limit of their global range, they are widely distributed in the BBWAA, except in Lake Clark and
its tributaries (Minard and Dunaway 1991, p. 2; Minard et al. 1998, p. 32), and the Tikchik Lakes
system, except for Tikchik Lake itself (Burgner et al. 1965, p. 11; Yanagawa 1967, p. 16-17).
They are most often found in medium to large streams and rivers and in lakes (ADF&G 2011;
Meka et al. 2003).
Rainbow trout typically spawn in flowing water, but can spawn along lake shores, near
groundwater upwellings (Northcote and Bull 2007). Rainbow trout in the Naknek River,
downstream of several large lakes, spawn in fast water of the mainstem, with much of the
spawning occurring in the transition between the upstream confined reach and the downstream
unconfined reach (Gwartney 1982, p. 9; Gwartney 1985, p. 47). Females deposit eggs, which are
immediately fertilized by males, into excavated redds (Morrow 1980b, p. 51). In Lower Talarik
Creek, Russell (1977, p. 9) reported that redds were dug in the gravel of side channels, near the
upstream ends of islands, and in pool tails above riffles. Typical water depths at Lower Talarik
Creek redd locations were less than 0.6 m (2 ft.) and current velocities were 0.3 to 0.6 m-s"1. The
most suitable sites for rainbow trout spawning in southcentral Alaska's Copper River system had
water temperatures ranging from 2 to 9 °C, average depths ranging from 0.3 to 0.4 m, average
current velocities of 0.5 to 0.7 m-s"1, and substrate diameters ranging from 20 to 60 mm (Brink
1995, p.71-75). In northern Idaho, rainbow trout spawned after the peak of spring snowmelt, and
redds had a mean area of 1.19 m (standard deviation (SD) = 0.62; range = 0.27 to 2.40 m ), a
mean water depth at the pit head of 0.18 m (SD = 0.08; range = 0.05 to 0.38 m), and a mean
water velocity at the pit head of 0.39 m-s"1 (SD = 0.15; range = 0.08 to 0.67 m-s"1) (Holecek and
Walters 2007). Steelhead in Alaska's Copper River, the size of large BBWAA rainbow trout,
dug redds averaging 3.4 m in area (Brink 1995, p. 125).
As the only spring-spawning member of its genus in the BBWAA, with eggs hatching later in the
summer than other Bristol Bay freshwater fish, young-of-the-year rainbow trout have a very
short time to complete incubation and initial growth before the onset of winter. Therefore,
spawning and early rearing habitats may be limited to locations with warmer summer
temperatures, as fry size in late fall is positively related to winter survival (Smith and Griffith
1994). Spawning areas in southcentral Alaska's Susitna and Copper river tributaries are often at
lake outlets, presumably because of warmer water there (Brink 1995, p. 16-18, 99; Sundet and
Pechek 1985, p. 37). Spawning began in spring when Lower Talarik Creek water temperatures
reached 2 to 3 °C, peaked at 4 to 7 °C, and stopped at temperatures greater than 16 °C (Russell
1977, p. 12).
In streams, rainbow trout summer rearing density increases with pool depth and overhead cover
(Bryant and Woodsmith 2009; Nakano and Kaeiryama 1995). Winter rearing density increases
with increasing availability of multiple cover types (Bjornn and Reiser 1991, p. 135). In summer
in southeast Alaska, rearing juveniles leave small tributaries and are relatively more abundant in
larger streams (>3rd order; sensu Strahler 1952, p. 1120). In spring and fall, juveniles are equally
distributed in both headwater tributaries and larger streams (Bramblett et al. 2002). However,
beginning in September, juvenile rainbow in Idaho move downstream from summer rearing to
winter overwintering areas (Chapman and Bjornn 1968, p. 165). Given the very low winter flows
and water temperatures in southwest Alaska low-order streams (e.g., USGS 2012), BBWAA
juvenile rainbow trout may follow the movement pattern of Idaho fish.
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In southeast Alaska, juvenile rainbow trout rear in streams with gradients up to at least 16%
(Bryant et al. 2004), but there are no reports of trout in such steep streams within the BBWAA
(ADF&G 2011). In streams of southwestern Alaska, in spring and early summer before the
arrival of adult salmon, large rainbow trout are lower in drainages, in slower velocity currents,
often in sloughs (Alt 1986). Later in the summer the distribution of age 1 and older Alaskan
rainbow trout is closely tied to the distribution of spawning salmon (Alt 1986; Brink 1995, p.
102, 104; Meka et al. 2003; Sundet and Pechek 1985, p. 39-40). In fall, after salmon spawning is
complete (except for coho), large southwestern Alaska rainbow trout occupy stream reaches with
moderate currents and gravel substrates, often near grassy banks (Alt 1986). Stream fish may
congregate in discrete overwintering habitats with moderate currents, often in areas of
groundwater upwelling (Sundet and Pechek 1985, p. 40). Groundwater influence may be an
important habitat characteristic because in regions where they are non-native, rainbow trout
invasion can be limited to only groundwater-fed streams with stable flows (Inoue et al. 2009).
Radio telemetry, tagging, and genetic studies indicate the presence of multiple rainbow trout
populations within Bristol Bay watersheds (Burger and Gwartney 1986, p. 22, 26; Gwartney
1985, p. 70-71; Krueger et al. 1999; Meka et al. 2003; Minard et al. 1992, p. 34).
Life cycle
Rainbow trout spawning in the Bristol Bay region is associated with spring ice-out and occurs
from late March through mid-June (Burger and Gwartney 1986, p. 22; Dye 2008, p. 21;
Gwartney 1985, p. 45-46, 51; Minard et al. 1992, p. 2; Russell 1977, p. 41). Pre-spawner
movements to spawning tributaries begins prior to ice-out, in early March (Dye 2008, p. 13).
Within a given drainage, the timing of spawning can vary by several weeks depending on spatial
and interannual stream temperature patterns (Burger and Gwartney 1986, p. 22; Hartman et al.
1962, p. 195; Russell 1977, p. 12). While post-spawners are often in poor physical condition
(Russell 1977, p. 15), BBWAA rainbow trout can spawn in consecutive years and some spawn at
least three years in a row (Minard et al. 1992, p. 17, 22; Russell 1977, p. 15).
In small lakes in southcentral Alaska, males matured at a smaller size than females and
approximately one-third of males smaller than 178 mm (SL, standard length; 7 in) were mature.
In this population most females did not mature until about 300 mm (SL; 12 in), while all males
matured at about 250 mm (SL; 10 in) (Allin 1954, p. 36). In Moose Creek, in the Wood River
lake system, half of the fish over 376 mm (FL) were sexually mature (Dye 2008, p. 22). In
Lower Talarik Creek, most spawners were ages 7 to 9 (Russell 1977, p. 17); in the upper
Kvichak River, from 1989 to 1991, spawners were primarily ages 5 to 7 (Minard et al. 1992, p.
15). Fecundity of Lower Talarkik Creek females (lengths ranging from 533 to 692 mm FL)
averaged 3,431 (n = 16, SD = 1,053) and ova diameter averaged 5.5 mm (n = 25, SD = 0.6,
Russell 1977, p. 18). In the BBWAA rainbow trout can reach at least age 14 (Minard and
Dunaway 1991, p. Ill, 189; determined by scale pattern analysis, a conservative measure; e.g.,
Sharp and Bernard 1988), with lengths to at least 814 mm (FL; Russell 1977, p. 30).
Post-spawning adults exhibit multiple movement patterns (Gwartney 1985, p. 68, 70; Meka et al.
2003). In Bristol Bay watersheds, many adults migrate shortly after spawning in inlet or outlet
streams of large lakes to feeding areas in large lakes (Burger and Gwartney 1986, p. 20; Meka et
al. 2003; Minard et al. 1992, p. 2; Russell 1977, p. 44). After a summer of feeding in lakes, from
September through November these mature rainbow trout move back to, or near, lake inlets and
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outlets to overwinter (Burger and Gwartney 1986, p. 20; Meka et al. 2003; Minard et al. 1992, p.
2; Russell 1977, p. 32). In the Wood River lakes system, mature rainbow trout from many
spawning streams aggregate to feed in the inter-lake rivers and remain there, or nearby in the
adjacent lakes, through the following winter (Dye 2008, p. 13). After spawning in tributaries to
southcentral Alaska's Susitna River, some mature rainbow trout remained near spawning areas,
some moved downstream, some moved into other tributaries, and some moved upstream (Sundet
and Pechek 1985, p. 39). Even in watersheds with large lakes, some fish may remain in outlet
rivers year-round (Meka et al. 2003). Fish grow little in winter (Russell 1977, p. 32).
While some mature fish may not undergo large seasonal migrations, others move considerable
distances (Dye 2008, p. 15; Meka et al. 2003; Minard et al. 1992, p. 33; Russell 1977, p. 23), to
at least 200 km (122 mi) or more (Burger and Gwartney 1986, p. 16). Meka et al. (2003)
speculated that seasonal migrations may be longer in watersheds with large lakes than in
watersheds without large lakes. In southwest Alaska's Goodnews River, most adult fish moved
less than 10 km throughout the year, and the movement that does occur is primarily upstream to
spring spawning locations, and downstream to overwintering locations (Faustini 1996. p. 19-20).
Incubating rainbow trout eggs develop much more rapidly than do those of salmon, and juveniles
emerge from spawning gravels between mid-July and mid-August at about 28 mm long
(ADF&G 2011, e.g., site FSN0616E01; Johnson et al. 1994; Russell 1977, p. 30). Juveniles grow
quickly during late summer and early fall, nearly doubling their length by late September
(Russell 1977, p. 30). Immature fish may remain in their natal stream for several years before
moving to other habitats (Russell 1977, p. 18, 22).
In the Alagnak River, within the BBWAA, Meka et al. (2003) distinguished three unique adult
life history patterns: lake-resident, lake-river, and river-resident. Each of these populations
migrates seasonally, and Meka et al. (2003) suggested that Alagnak rainbow trout evolved these
movements to take advantage of seasonal food sources (salmon eggs and carcasses) and warmer
winter water temperatures. Russell (1977, p. 37) noted that Lower Talarik Creek trout were in
better condition following large Kvichak drainage sockeye salmon escapements than after small
escapements.
Predator-prey relationships
The diet of rearing rainbow trout includes a broad range of aquatic and terrestrial invertebrates
(Nakano and Kaeiryama 1995). When available, sockeye salmon eggs dominated rainbow trout
diet in Lower Talarik Creek. While their diet was highly varied, other important Lower Talarik
Creek rainbow trout food items included aquatic dipterans (chironomids) and caddis fly larvae
(Russell 1977, p. 36). Many larger Lower Talarik Creek rainbow trout appear to feed primarily in
Iliamna Lake and not in the stream (Russell 1977, p. 35). In rivers of the BBWAA, Russell
(1980, p. 103) reported that aquatic insects, salmon eggs, shrews and voles, unidentified fish and
Chinook salmon fry, and salmon carcasses made up the bulk of the summer and fall diet of
rainbow trout.
In the Wood River lakes system, Scheuerell et al. (2007) reported that before the seasonal arrival
of adult salmon, rainbow trout primarily feed on dipterans (39%), stoneflies (18%), mayflies
(12%), and caddis flies (11%). When spawning sockeye salmon arrive, rainbow trout diet shifts
to primarily salmon eggs (64%), larval blowflies (which feed on salmon carcasses; (11%)), and
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salmon carcasses (9%). This diet shift in conjunction with seasonal salmon spawning activity
increases rainbow trout energy intake more than five-fold (Scheuerell et al. 2007).
In the laboratory, slimy sculpin, a ubiquitous species throughout the lakes and streams of the
BBWAA, consume rainbow trout eggs (Fitzsimons et al. 2006). While BBWAA rainbow trout
are certainly consumed by predators, they are not specifically identified in the diet of regional
predatory fish (Metsker 1967, p. 26, 29; Russell 1980, p. 55-56, 62-63, 67, 73, 76, 81-83, 95-97,
103, 108), perhaps due at least in part to their comparatively low abundance relative to other
available prey species.
Abundance and harvest
In the BBWAA total rainbow trout abundance is unknown, but there have been population
estimates of larger fish in selected streams. From 2,000 to 4,500 fish available to hook and line
angling gather in the upper Kvichak River in spring (Minard et al. 1992, p. 30); an average of
950 fish spawn in Lower Talarik Creek (Russell 1977, p. 9); and 950 fish larger than 199 mm
occur in the Tazimina River, north of Iliamna Lake (Schwanke and Evans 2005, p. 9). In the
Wood River lakes system, counts have been as high as 13,700 rainbow trout larger than 250 mm
in the Agulowak River and 2,400 larger than 340 mm in the Agulukpak River (Dunaway 1993,
p. 10, 24).
In the BBWAA and the adjacent Togiak River drainage, sport anglers caught more rainbow trout
in 2009 (an estimated 159,685, or 22% of the statewide total) than all other non-salmon fish
species combined (Jennings et al. 2011, p. 69). In 2009 sport anglers harvested 225 rainbow trout
within the BBWAA and adjacent Togiak River drainage (Jennings et al. 2011, p. 69). Annual
sport harvests have declined, due at least in part to the increasing popularity of catch-and-release
fishing (Dye and Schwanke 2009, p. 6). The State of Alaska's Southwest Alaska Rainbow Trout
Management Plan includes policies to manage BBWAA rainbow trout populations to maintain
historic size and age composition without relying on hatcheries, to provide a range of harvest
opportunities, and to economically develop the sport fishing industry while acknowledging the
intrinsic value of the resource to Alaskans (Dye and Schwanke 2009, p. 32).
From the mid-1970s to the mid-2000s, rainbow trout were estimated to represent between 19 and
30.9% of the total weight of the Kvichak River drainage non-salmon freshwater fish subsistence
harvest (Krieg et al. 2005, p. 214). In the mid-2000s, villagers from nine of the BBWAA
communities annually harvested, as part of their subsistence activities, about an estimated 3,740
rainbow trout (Fall et al. 2006, p. 45, 80, 113, 150, 194; Krieg et al. 2009, p. 40, 78, 118, 162,
202).
Stressors
Low pH (less than or equal to pH 5.5) impairs adult egg and sperm development and reduces
early embryonic survival (Weiner et al. 1986). Pre-emergent embryo survival depends strongly
on elevated DO concentrations and movement of groundwater through redds. Embryo survival is
minimal where mean DO is less than 5.2 mg-1"1; at higher DO levels, embryo survival increases
in relation to the velocity of intergravel flows greater than 5 cm-h"1 (Sowden and Power 1985).
Bjornn and Reiser (1991, p. 84, 85) concluded that upstream migrating large trout need stream
depths no less than 0.18 m, velocities no more than 2.44-sn "*, and DO levels at least 80% of
saturation and never less than 5.0 mgT1. For spawning rainbow trout in the more central part of
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their North American range, Bell (1986, p. 96) recommended water temperatures between 2.2
and 20 °C (36 to 68 °F), and optimally 10 °C (49.5 °F). Russell (1977, p. 12) observed that
Lower Talarik Creek rainbows stopped spawning at stream temperatures above 16 °C. In the
laboratory, at temperatures below 2.8 °C age-0 fry become inactive and seek refuge within the
stream substrate. At temperatures below 5.5 °C, fry stop feeding (Chapman and Bjornn 1968, p.
168).
The survival of incubating embryos rapidly declines as the proportion of fines (sediments less
than 6.35 mm in diameter) increases in spawning gravels, probably because the fines reduce
intragravel flow (Bjornn and Reiser 1991, p. 99, 100). The success rate of fry emergence from
spawning gravels and juvenile rearing density also decline with increasing proportion of fines in
the substrate (Bjornn and Reiser 1991, p. 103, 132). Rainbow trout populations are particularly
vulnerable when adult fish aggregate in spring spawning grounds and overwintering locations.
Ten steelhead population segments in California, Oregon, and Washington are currently listed as
threatened or endangered primarily due to the lack of access to their historic range that has
resulted from constructed barriers to migration and to stream dewatering. Nonanadromous
rainbow trout populations are not listed (NMFS 2006).
Char
The BBWAA is home to three species of char: Arctic char, Dolly Varden, and lake trout. These
char all spawn in fall. Bristol Bay basin Dolly Varden are often anadromous; Arctic char and
lake trout are typically nonanadromous. The habitats of Dolly Varden and Arctic char
occasionally overlap within the BBWAA, and when they do these species may hybridize (Taylor
et al. 2008).
The taxonomic distinctions between Arctic char and Dolly Varden historically have been
inconsistent. Some earlier authors (e.g., Craig 1978; Craig and Poulin 1975; Yoshihara 1973)
called riverine and anadromous Alaskan char "Arctic char" Salvelinus alpinus. More recent
assessments suggest these fish are Dolly Varden (Behnke 1980, p. 454; Cavender 1980, p. 319-
320; Taylor et al. 2008). In general, researchers currently believe that the North American char
west of Canada's Mackenzie River living primarily in flowing water are Dolly Varden, and
Arctic char (and lake trout) are largely limited to lakes and adjacent reaches of their inlet and
outlet streams (Reist et al. 1996).
The State of Alaska's 2011 edition of the Catalog of Waters Important for the Spawning,
Rearing or Migration of Anadromous Fishes, or "Anadromous Waters Catalog" (AWC; e.g.,
Johnson and Blanche 2011) identifies Dolly Varden as the anadromous char across most of the
state. However, in Bristol Bay the AWC identifies some streams as anadromous Dolly Varden
habitat and some as anadromous Arctic char habitat. The AWC lists both anadromous Dolly
Varden and anadromous Arctic char in the Kvichak River drainage, but only anadromous Arctic
char in the Nushagak River drainage. These distinctions result from the history of regional
variations in species naming and do not accurately reflect the ranges of different species and life
histories. Current terminology labels the river-dwelling BBWAA char Dolly Varden. That is, the
rivers and streams in the AWC currently designated as Arctic char habitat should, in almost all
cases, be interpreted as Dolly Varden habitat. As a result of recent field work, ADF&G
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concluded that the Nushagak River, and the Koktuli River in particular, likely supported
anadromous Dolly Varden (Schwanke 2007, p. 14).
Arctic char Salvelinus alpinus
Freshwater distribution and habitats
The Arctic char is a circumpolar species, distributed at high latitudes across the northern
hemisphere (Brunner et al. 2001). In fresh water, Arctic char range closer to the North Pole than
any other fish species (Johnson 1980, p. 16). In the fresh waters of North America, Arctic char
are not typically far from the ocean. They range from Maine and New Hampshire north to the
Canadian mainland Arctic Coast and through the Canadian Arctic archipelago (Scott and
Grossman 1998, p. 203). The Alaskan Arctic char distribution is disjunct. They occur in the
Brooks Range, on the North Slope and the Seward Peninsula, in Bristol Bay, and a few other
isolated locations in southcentral and interior Alaska (Mecklenburg et al. 2002, p. 199). Multiple
distinct Arctic char races, differing in growth rate and life history, can occupy a single lake
(Baroudy and Elliott 1994; Sandlund et al. 1992).
Alaskan Arctic char appear primarily restricted to lakes and adjacent reaches of their inlet and
outlet streams in well-drained areas (Morrow 1980b, p. 58; Scanlon 2000, p. 56, 58; Taylor et al.
2008) and do not appear to undertake extensive seasonal migrations outside their home lakes
(McBride 1980, p. 17). However, some Alaskan Arctic char are known to move 15 to 20 km
upstream and downstream between connected lakes (Troyer and Johnson 1994, p. 49) and
Scanlon (2000, p. 43-48) suggested some move seasonally to estuarine or marine areas. Within
the BBWAA, they are reported in the Tikchik and Wood River lakes, Hiamna Lake, and other
upland lakes (Bond and Becker 1963; Burgner et al. 1965; Russell 1980, p. 49; Taylor et al.
2008). Metsker (1967, p. 23) believed that Intricate Bay in Diamna Lake is a particularly
important spawning area. Adults and juveniles are common in the east end of Iliamna Lake, but
not in tributaries (Bond and Becker 1963).
The depth of Arctic char lake spawning habitat can vary from 1 to 100 m (reviewed in Johnson
1980, p. 44), but is often in gravel shoals less than 5 m deep (Klemetsen et al. 2003, p. 31).
McBride (1980, p. 6) found Wood River lakes spawners concentrated in the mouths of small
tributary streams. DeLacy and Morton (1943) concluded that Kodiak Island's Karluk Lake
Arctic char spawn in the lake and not in the tributary streams.
During the spring and early summer, McBride (1980, p. 20) estimated that approximately 40%
(approximately 65,000) of the Wood River lakes Arctic char population greater than 300 mm
long congregated in the inlets and outlets of the inter-lake rivers to feed on the sockeye salmon
smolt outmigration. In Bristol Bay's Ugashik lakes, Plumb (2006, p. 14-15) found Arctic char at
depths greater than 75 m; but 90% of her catch was in waters less than 10 m deep. Fish sizes
were not segregated by depth (Plumb 2006, p. 19-20). Similar to Dolly Varden (discussed
below), Arctic char often occupy different habitats depending on the presence or absence of
competitors (reviewed in Klemetsen et al. 2003, p. 29-30)
Life cycle
Arctic char in Bristol Bay are thought to be primarily nonanadromous (e.g., Reynolds 2000, p.
16), but Scanlon (2000, p. 43-48) suggested that some Becharof Lake Arctic char were
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anadromous. In BBWAA lakes, maturity is reached at around ages 3 to 6, at a length of
approximately 330 mm (FL; 13 in.) (Metsker 1967, p. 23; Russell 1980, p. 48, 54). Metsker
(1967, p. 23, 26) concluded that individual Diamna Lake Arctic char spawned in alternating
years, but McBride (1980, p. 16) provided evidence that at least some Lake Aleknagik Arctic
char return annually to spawning locations. Wood River lakes Arctic char demonstrated high
level of interannual fidelity to both spawning and feeding sites (McBride 1980, p. 6, 8, 19). Lake
Aleknagik Arctic char periodically provide eggs for Alaska's sport fish hatcheries (Dunaway and
Jaenicke 2000, p. 138).
In the BBWAA Russell (1980, p. 48) found individuals ready to spawn in mid-September and
McBride (1980, p. 6) collected Wood River lakes spawning fish between mid-September and
mid-October. Ripening females in Brooks Range lakes have ova diameters ranging from 1.62 to
4.75 mm and fecundity ranges from 3,200 to 4,000 ova (Troyer and Johnson 1994, p. 41). If the
substrate is not too coarse (approximately 10 cm or more, Sigurjonsdottir and Gunnarsson 1989)
females excavate redds into which they deposit their ova, which males immediately fertilize
(Johnson 1980, p. 45). The incubating eggs and alevins remain in spawning gravels until the
following spring (summarized in Johnson 1980, p. 47-48). Bristol Bay Arctic char live at least 15
years (Plumb 2006, p. 19), are particularly slow growing (Russell 1980, p. 48), reach fork
lengths to at least 684 mm, and weights to at least 3.8 kg (Scanlon 2000, Appendix Table A). As
with Dolly Varden, multiple life history patterns and morphologies (Klemetsen et al. 2003, p. 36)
occur with the basin (Russell 1980, p. 48; Scanlon 2000, p. 63-64). Tagging studies indicated
that the Wood River lakes supported at least 20 discrete stocks (McBride 1980, p. 20).
Predator-prey relationships
The diet of young-of-the-year is poorly understood, but is thought in general to be dominated by
small benthic and planktonic invertebrates (reviewed in Klemetsen et al. 2003, p. 32). In larger
Brooks Range fish, planktonic crustaceans, insects, and snails were the most frequently
occurring food items and fish were not an important part of the diet (Troyer and Johnson 1994, p.
44). In Iliamna Lake, summer diet was dominated by snails (Bond and Becker 1963) and winter
diet was dominated by threespine stickleback (Metsker 1967, p. 26, 28). In other BBWAA lakes,
mollusks and caddis fly larvae were the dominant benthic organisms consumed (Russell 1980, p.
55-56). In summer, freshwater crustaceans dominated the diet of Ugashik Lakes Arctic char
(Plumb 2006, p. 27) and crustaceans, sticklebacks, insects, pygmy whitefish, sculpins, and
juvenile sockeye salmon dominated the diet of Becharof Lake Arctic char (Scanlon 2000, p. 51,
53-54).
In the BBWAA, larger Arctic char eat outmigrating sockeye salmon smolt, often in spring and
early summer at lake outlets (McBride 1980, p. 1; Metsker 1967, p. 29). Karluk Lake Arctic char
eat mostly insects until the arrival of spawning sockeye, when their diet shifts to drifting salmon
eggs, benthic invertebrates dislodged by salmon redd excavation, and adult salmon carcasses
(DeLacy and Morton 1943).
Arctic char are eaten by other predatory fish, including lake trout (Troyer and Johnson 1994, p.
42) and larger Arctic char (Klemetsen et al. 2003, p. 33). Mink eat mature Wood River lakes
Arctic char when they have the opportunity (Dunaway and Jaenicke 2000, p. 138).
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Abundance and harvest
In the BBWAA total Arctic char abundance is unknown. Meacham (reported in McBride 1980,
p. 20) estimated that in the 1970s the Wood River lakes supported between 135,000 and 210,000
(presumably larger) Arctic char. Russell (1980, p. 48, 49) considered them common in some
lakes in the Lake Clark area, but absent or rare in lakes of the upper Mulchatna River watershed
and Lake Clark itself. In the mid-1960s, Hiamna Lake supported a commercial fishery and char
made up 84% (2,979 kg, 6,553 Ib) of the total dressed weight harvest (Metsker 1967, p. 9). These
fish are thought to be mostly Arctic char (Bond and Becker 1963; Taylor et al. 2008).
Between 1971 and 1980, the annual estimated abundance of Arctic char larger than 249 mm
ranged from 8,000 to 12,000 fish at the mouth of the Agulowak River and 4,300 to 7,800 fish at
the mouth of the Agulukpak River (Minard et al. 1998, p. 131). By 1993 the estimated
abundance of the Agulowak River population declined to only 5,400 fish, prompting a
substantial reduction in bag limits and harvest means (Minard and Hasbrouck 1994, p. 13, 22).
While excessive sport harvests were thought to be responsible for the decline (Minard et al.
1998, p. 16), anecdotal reports suggest that the more conservative sport harvest regulations were
leading to the recovery of the stock (Dunaway and Sonnichsen 2001, p. 131). Minard et al.
(1998, p. 16) also reported a similar apparent significant decline in Iliamna River stocks, both in
overall abundance and in larger, older age classes. These observations prompted adoption of a
catch-and-release fishing regulation.
The State of Alaska's sport and subsistence fisheries statistics do not distinguish between Arctic
char and Dolly Varden. Sport anglers caught an estimated 48,438 Arctic char/Dolly Varden in
the BBWAA and the adjacent Togiak River system in 2009 (8% of the statewide total) and
harvested (kept) an estimated 2,159 (5% of the statewide total; Jennings et al. 2011, p. 73).
Arctic char/Dolly Varden consistently support the greatest sport harvest of any non-salmon
freshwater fish in Bristol Bay (Dye and Schwanke 2009, p. 8). Sport harvests have declined, due
at least in part to both lower bag limits and the increasing popularity of catch-and-release fishing
(Dye and Schwanke 2009, p. 6).
In the mid-2000s, villagers from ten of the BBWAA communities annually harvested, as part of
their subsistence activities, about an estimated 3,450 Arctic char and Dolly Varden combined
(Fall et al. 2006, p. 45, 80, 113, 150, 194; Krieg et al. 2009, p. 40, 78, 118, 162, 202). Arctic char
and Dolly Varden combined were the most important non-salmon fish harvested in the villages
of Iliamna, Newhalen, and Pedro Bay (Fall et al. 2006, p. 49, 84, 117). From the mid-1970s to
the mid-2000s, Arctic char/Dolly Varden were estimated to represent between 16.2 and 26.9% of
the total weight of the Kvichak River drainage non-salmon freshwater fish subsistence harvest
(Krieg etal. 2005, p. 214).
Stressors
Arctic char are not tolerant of warm water. In tests of European Arctic char, egg mortality was
100% at temperatures at or above 12 to 13 °C (Jungwirth and Winkler 1984). Even when
acclimated to water temperatures between 15 and 20 °C, pre-emergent fry could not survive
exposures to temperatures above 26.6 °C for more than 10 minutes and could not survive
temperatures over 21° C for more than a week (Elliott and Klemetsen 2002). Apparent over-
harvests have been implicated for historic population declines within the BBWAA (Minard et al.
1998, p. 16).
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Dolly Varden Salvelinus malma
Dolly Varden is a highly plastic species: multiple genetically, morphologically, and ecologically
distinct morphs (e.g., benthic specialist, riverine specialist, lacustrine generalist, specialized
piscivore) can exist in the same water body (Ostberg et al. 2009). Researchers currently
recognize two geographically distinct forms of Dolly Varden: northern and southern, based on
differences in life history (Armstrong and Morrow 1980, p. 107-130), phenotype (Behnke 1980,
465-466; Cavender 1980, p. 299-318), and genotype (Taylor et al. 2008). Dolly Varden in the
BBWAA are of the northern form (Behnke 1980, p. 465).
Freshwater distribution and habitats
The global native freshwater range of Dolly Varden is restricted to waters draining to the
Beaufort, Chukchi and Bering seas and the North Pacific. The North American range extends
from the Arctic coast of Alaska and Canada west of the Mackenzie River south to northern
Washington. The Asian range stretches from the Chukotka Peninsula south to Japan and Korea
(Mecklenburg et al. 2002, p. 200). In Alaska, Dolly Varden are found in waters draining to all
coasts (Mecklenburg et al. 2002, p. 200) and the Alaska Peninsula divides the northern and
southern forms (Behnke 1980, p. 453). Dolly Varden are known to occur widely in Bristol Bay,
but their true distribution across the waters of the BBWAA is underreported. Within the
BBWAA, popular sport fishing areas include the Alagnak, Newhalen, Nushagak, Mulchatna, and
the Wood River-Tikchik Lakes systems (Minard et al. 1998, p. 188).
As in southeast Alaska (Bryant et al. 2004), BBWAA Dolly Varden occur farther upstream in
high-gradient headwater streams than other fish species (ADF&G 2011, e.g., site FSN0604E01).
In both southeast Alaska (Bramblett et al. 2002; Wissmar et al. 2010) and the BBWAA (ADF&G
2011, e.g., site FSN0616E01; e.g., Tazimina Lakes, Russell 1980, p. 31-32, 73), resident Dolly
Varden occur above migratory barriers that currently prevent access to anadromous salmon
populations.
Spawning occurs well upstream from areas used for overwintering (DeCicco 1992). Northern-
form anadromous Dolly Varden overwinter primarily in lakes and in lower mainstem rivers
where sufficient groundwater provides suitable volumes of free-flowing water (DeCicco 1997;
Lisac 2009, p. 13, 15-16). In stream systems, spawning occurs in fast-flowing channels,
primarily in upper reaches (Bramblett et al. 2002; Fausch et al. 1994; Hagen and Taylor 2001;
Kishi and Maekawa 2009; Koizumi et al. 2006) and small, spring-fed tributaries (Hagen and
Taylor 2001). Stream-resident Dolly Varden are reported to spawn in channels that are 1 to 3 m
wide and 10 to 35 cm deep (Hino et al. 1990; Maekawa et al. 1993), with a mean depth of 9 cm,
mean velocity of 21 cm-s"1, and median substrate diameter of 1.6 cm (Hagen and Taylor 2001).
Stream-resident females select spawning sites where gravel is prevalent (Kitano and Shimazaki
1995). Spawning site substrate and current velocity do not correlate significantly with female
size, but redd depth does (Kitano and Shimazaki 1995). Anadromous individuals spawn in
deeper water than resident fish, ranging from 20 to 60 cm (Blackett 1968). They construct redds
approximately 30 cm long, 15 to 25 cm wide, and 15 cm deep (Blackett 1968); composite redds,
potentially containing several individual nests can be up to 3.5 m long and 1.2 m wide
(Yoshihara 1973, p. 47).
In Kamchatka Eberle and Stanford (2010) found rearing Dolly Varden in floodplain
springbrooks and 7* -order mainstem channels. Within the BBWAA, juveniles appear to be
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limited primarily to low-order headwaters (ADF&G 2011), and infrequently to side channels and
the main channel of larger rivers downstream to the confluence of 5th-order streams (ADF&G
2011, e.g., site FSN0609A02). In southeast Alaska Dolly Varden rear in channels with gradients
steeper than 20% (Wissmar et al. 2010), but in the BBWAA, Dolly Varden have been reported
only in gradients of 12% or less (ADF&G 2011, e.g., site FSM0503A07). Rearing Dolly Varden
normally stay close to the stream bottom over gravels and cobbles (Dolloff and Reeves 1990;
Hagen and Taylor 2001; Nakano and Kaeiryama 1995). Fry density is inversely related to stream
depth (Bryant et al. 2004) and use of shallows increases if cover is available (Bugert et al. 1991).
Different juvenile age classes can segregate in different micro- (Bugert et al. 1991; Dolloff and
Reeves 1990) and macro- (ADF&G 2011; Denton et al. 2009) habitats. Affinity for cover,
including cobbles and boulders, increases with age and tolerance for other Dolly Varden declines
(Dolloff and Reeves 1990). Gregory (1988, p. 49-53) found stream-resident juvenile Dolly
Varden in beaver ponds, where they grow faster than fish in adjacent streams, because of
relatively warmer water temperatures and increased productivity.
Dolly Varden occur in upland Bristol Bay lakes, often in large numbers, feeding both at the
surface and on the lake bottom, but they are uncommon or absent in lakes supporting Arctic char
populations (Russell 1980, p. 49, 69-72; Scanlon 2000, p. 56). Dolly Varden will use all lake
habitats in the absence of competitors (other salmonids), but concentrate in offshore and near-
bottom habitats where competitors occupy nearshore and near-surface habitats (Andrew et al.
1992; Jonsson et al. 2008; Schutz and Northcote 1972). In the absence of competitors, lake-
dwelling Dolly Varden move from deeper offshore waters, where they spend the day, perhaps in
loose aggregations, to spend the night in onshore waters, near the surface (Andrusak and
Northcote 1971). Dolly Varden vision is more sensitive to low light than competing salmonids
(Henderson and Northcote 1985; Henderson and Northcote 1988; Schutz and Northcote 1972),
allowing them to feed in deeper water and at night.
Life cycle
Northern-form Dolly Varden express several life history patterns, including anadromous,
nonanadromous stream-resident, nonanadromous spring-resident, nonanadromous lake-resident,
nonanadromous lake-river-resident, and nonanadromous residuals (nonanadromous male
offspring of anadromous parents; (Armstrong and Morrow 1980, p. 107-130; Behnke 1980, p.
466). The Bristol Bay basin supports Dolly Varden with both anadromous (Reynolds 2000, p.
16-17; Scanlon 2000, p. 48-51) and nonanadromous (Denton et al. 2009; Scanlon 2000, p. 48-51)
life histories.
Anadromous Dolly Varden exhibit very complex migratory patterns (Armstrong and Morrow
1980, p. 108-109), frequently leaving one drainage, traveling through marine waters, and
reentering distant drainages, including those on separate continents (DeCicco 1992; DeCicco
1997; Lisac 2009, p. 14; Morrow 1980a). Even apparently nonanadromous fish can seasonally
move more than 200 km within complex Bristol Bay watersheds (Scanlon 2000, p. 60).
Anadromous Dolly Varden of the Togiak River system, just west of the BBWAA, spawn from
approximately mid-September to mid-October, overwinter downstream from spawning locations,
and migrate annually to sea, where they spend approximately six weeks feeding (Lisac and Nelle
2000, p. 31-34). The timing of adult seaward migration generally corresponds with spring ice-out
and high water, with adults migrating to sea in May and June. Their return to fresh water appears
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to relate to decreased stream discharge (Lisac and Nelle 2000, p. 33-34, 35). Anadromous Dolly
Varden migrate upstream from the ocean to spawning areas in July and August (Lisac 2011).
Russell (1980, p. 72) observed Dolly Varden spawning in the upper Mulchatna River system in
mid-September.
Anadromous Dolly Varden home to spawn (Crane et al. 2003; Lisac and Nelle 2000, p. 31), but
stocks can mix at sea and in overwintering areas (DeCicco 1992). In northwest Alaska
anadromous Dolly Varden usually undertake three to five ocean migrations before reaching
sexual maturity (DeCicco 1992). In the Togiak River, some anadromous fish mature at age 2 and
most mature at age 4 (Lisac and Nelle 2000, p. 31; Reynolds 2000). Bristol Bay Dolly Varden
can live at least 14 years (Plumb 2006, p. 19; Scanlon 2000, Appendix Table B) and reach
lengths of 740 mm or more (Faustini 1996, p. 16). The minimum length of anadromous spawners
in southwest Alaska's Goodnews River is about 330 to 360 mm (Lisac 2010, p. 4).
Stream-residents mature from age 2 to 5 (Blackett 1973; Craig and Poulin 1975; Maekawa and
Hino 1986; Russell 1980, p. 72) and live at least to age 7 (Blackett 1973). They are smaller than
their anadromous counterparts, ranging at maturity from 113 mm (Hagen and Taylor 2001) to
520 mm (Gregory 1988, p. 29) in length, with most less than 200 mm (Gregory 1988, p. 21-25).
Like anadromous individuals, after spawning stream-resident adults move quickly to
downstream overwinter areas (Maekawa and Hino 1986).
Although anadromous Dolly Varden in northern Alaska tend to spawn only every second year
(DeCicco 1997), Lisac and Nelle (2000, p. 31) speculated that most anadromous Dolly Varden in
the Togiak River near the BBWAA can spawn in consecutive years. Female fecundity is a
function of size (Jonsson et al. 1984), and anadromous females can produce up to 7,000 ova
(Armstrong and Morrow 1980, p. 102), a productivity more than 50 times that of resident
females (Blackett 1973). Ripe eggs of anadromous females are 3.5 to 6 mm in diameter; ripe
eggs of resident females can be as small as 2.8 mm (Armstrong and Morrow 1980, p. 101, 102).
In most cases, a spawning group consists of one female and several males, one of which is a
dominant male that actively courts the female (Hino et al. 1990; Maekawa et al. 1993). Females
excavate redds in stream gravels, and then deposit their eggs while a male fertilizes them. Chars
show little evidence of nest-guarding behavior (Kitano and Shimazaki 1995). Males appear to
suffer a much higher post-spawning mortality than do females (Armstrong 1974).
In streams on both sides of the Bering Strait, egg hatching peaks from the end of April to mid-
May (Radtke et al. 1996). Embryos are 15 to 20 mm long at hatching and remain in the spawning
substrate while they absorb their yolk sac. Alevins emerge from the nest around the time of ice
break-up (April to June), at a length of about 25 mm (Armstrong and Morrow 1980, p. 108).
Radtke et al. (1996) found that first feeding begins from June to early July, 42 to 52 days after
hatching. Newly emerged alevins tend to stay on the bottom of pools and are relatively inactive
except when feeding (Armstrong and Morrow 1980, p. 108). Growth greatly increases through
the summer as water becomes warmer; by September, age-0 fish average about 60 mm long
(Armstrong and Morrow 1980, p. 108). Young anadromous Togiak River Dolly Varden make
their first seaward migration between their first summer and age 3 (Reynolds 2000, p. 15). Size,
rather than age, appears to govern the timing of initial smolt out-migration (Armstrong 1970).
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Predator-prey relationships
Dolly Varden primarily target benthic invertebrates in streams (Eberle and Stanford 2010;
Russell 1980, p. 73; Stevens and Deschermeier 1986) and lakes (Scanlon 2000, p. 53-55; Schutz
and Northcote 1972). During the day, foraging from stream drift (food drifting in the current) is
more important than benthic foraging, but the relative importance of benthic foraging increases
at night; surface feeding is not important (Hagen and Taylor 2001). Dolly Varden also switch to
benthic feeding when drift availability is limited (Fausch et al. 1997; Nakano et al. 1999; Nakano
and Kaeiryama 1995).
Dolly Varden eat juvenile salmon (Armstrong 1970; Bond and Becker 1963), but they have been
largely exonerated (Armstrong and Morrow 1980, p. 133; DeLacy and Morton 1943; Morton
1982) from earlier accusations that they were salmon run destroyers. From 1921 to 1939,
Alaskan Dolly Varden were the target of a bounty program designed to increase salmon
abundance. Now it is believed that Dolly Varden were not responsible for the declines in salmon
abundance (Harding and Coyle 2011, p. 19). When spawning salmon are present, salmon eggs-
probably those flushed by high flows and superposed redd construction-can be important food
(Armstrong 1970, p. 53-54; Scanlon 2000). Denton et al. (2009) reported that resident age-1 and
older Dolly Varden in certain ponds near Iliamna Lake feed on sockeye salmon fry for a brief
time in late June to mid-July, then migrate to sockeye spawning areas and feed almost
exclusively on eggs from late July to mid-September. From late August through September they
also eat blowfly larvae that had fed on adult sockeye salmon carcasses. Salmon eggs are too big
for age-0 fry to consume, but blowfly maggots, when available, dominate their diet. Resident
Dolly Varden actively follow adult sockeye salmon to spawning areas and grow significantly
faster after the arrival of spawning salmon (Denton et al. 2009; Wipfli et al. 2003). In May in
Iliamna Lake tributaries such as the Copper River, Dolly Varden feed heavily on the spawning
run of mature pond smelt (Richard Russell, Alaska Department of Fish and Game (retired), King
Salmon, AK, personal communication).
The summer diet of stream-resident Dolly Varden in northcentral British Columbia is primarily
adult dipterans (true flies; 33.6%) and hymenopterans (wasps, bees, and ants; 7.5%), with other
aquatic insects comprising the remainder (Hagen and Taylor 2001). In southeast Alaska Dolly
Varden also feed on terrestrial insects, but do so less than other salmonids occupying the same
habitat (Wipfli 1997). Juvenile stream-rearing Dolly Varden consume a wide variety of
predominantly aquatic invertebrates (Eberle and Stanford 2010), preferentially selecting
immature blackflies, non-biting midges (chironomids), and mayflies (Milner 1994; Nakano and
Kaeiryama 1995), but also feed on terrestrial invertebrates (Baxter et al. 2007; Nakano et al.
1999), particularly in the absence of competing salmonids (Baxter et al. 2004; Baxter et al.
2007). Some juvenile Dolly Varden eat age-0 Arctic grayling (Stevens and Deschermeier 1986).
In the absence of competitors, lake-dwelling Dolly Varden feed heavily in summer on terrestrial
insects and during fall on zooplankton. In the presence of competition, they feed heavily on
chironomids (both pupae and larvae) and trichopterans (caddis flies; Andrusak and Northcote
1971;Hindaretal. 1988).
River otters Lutra canadensis can extensively prey on rearing Dolly Varden (Dolloff 1993).
Armstrong and Morrow (1980, p. 110) noted that bears and wolves take some mature fish from
spawning areas (also observed by Wiedmer; ADF&G 2011, site FSS0424A07) and speculated
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that fish-eating birds also take a few. Fish-eating birds such as harlequin ducks Histrionicus
histrionicus, common Mergus merganser and red-breasted M. serrator mergansers, and bald
eagles Haliaeetus leucocephalus are common in southwest Alaska throughout the year and
ospreys (Pandion haliaetus, a fish-eating raptor) are more abundant along the waters of Bristol
Bay than elsewhere in Alaska (Armstrong 1980, p. 69, 80, 81, 89, 92). Russell (1980, p. 81)
reported that Lake Clark National Park and Preserve lake trout feed on Dolly Varden. Perhaps
the greatest predators on smaller Dolly Varden are larger Dolly Varden (Armstrong and Morrow
1980, p. 110; Russell 1980, p. 73). Wiedmer (ADF&G 2011, site FSS0406A01) collected a 195
mm (FL) northern- form Dolly Varden that had partially swallowed a 98 mm Dolly Varden.
Abundance and harvest
In the BBWAA total Dolly Varden abundance is unknown. Annual runs of anadromous Dolly
Varden to southwest Alaska's Kanektok River average 13,115 (range: 8,140 to 43,292, Lisac
2011). The State of Alaska's sport and subsistence fisheries statistics do not distinguish between
Arctic char and Dolly Varden. Sport anglers caught an estimated 48,438 Arctic char/Dolly
Varden in the BBWAA and the adjacent Togiak River system in 2009 (8% of the statewide total)
and harvested (kept) an estimated 2,159 (5% of the statewide total; Jennings et al. 2011, p. 73).
The combination of Arctic char/Dolly Varden consistently support the greatest harvest of any
non-salmon freshwater fish in Bristol Bay (Dye and Schwanke 2009, p. 8). Sport harvests have
declined, due at least in part to both lower bag limits and the increasing popularity of catch-and-
release fishing (Dye and Schwanke 2009, p. 6).
In the mid-2000s, villagers from ten of the BBWAA communities annually harvested, as part of
their subsistence activities, about an estimated 3,450 Dolly Varden and Arctic char combined
(Fall et al. 2006, p. 45, 80, 113, 150, 194; Krieg et al. 2009, p. 40, 78, 118, 162, 202). Dolly
Varden and Arctic char combined were the most important non-salmon fish harvested in the
villages of Iliamna, Newhalen, and Pedro Bay (Fall et al. 2006, p. 49, 84, 117). From the mid-
1970s to the mid-2000s, Dolly Varden/Arctic char were estimated to represent between 16.2 and
26.9% of the total weight of the Kvichak River drainage non-salmon freshwater fish subsistence
harvest (Krieg et al. 2005, p. 214).
Stressors
Dolly Varden are not tolerant of warm water (Fausch et al. 1994; Kishi et al. 2004; Nakano et al.
1996). Feeding activity declines to low levels at water temperatures above 16 °C and their upper
lethal limit is 24 °C (Takami et al. 1997). As a result, activities that increase water temperatures
beyond tolerance levels will reduce available habitat (Kishi et al. 2004; Nakano et al. 1996),
including the refuge from potential competitors that cold stream temperatures provide (Fausch et
al. 2010).
Total dissolved solids (TDS) do not have a significant impact on Dolly Varden fertilization, up to
the highest concentrations evaluated (1,817 mg-F1); however, elevated TDS did significantly
affect embryo water absorption at concentrations as low as 964 mg-1 l (Brix et al. 2010). Brix et
al. (2010) concluded that the water-hardening phase immediately following fertilization was the
most sensitive life stage to elevated TDS.
McDonald et al. (2010) reported that Dolly Varden are relatively insensitive to selenium
exposure (perhaps due to low rearing temperatures) and estimated that concentrations of 44 and
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49 mg-kg"1, dry weight affected 10 and 20% of the study population, respectively. Dolly Varden
in fresh water metabolize naphthalene much more rapidly than seawater, which may explain the
greater toxicity of naphthalene to fish when in seawater (Thomas and Rice 1980). Whether in
fresh water or sea water, toluene is more readily metabolized by Dolly Varden than is
naphthalene (Thomas and Rice 1986b), and toluene is more rapidly metabolized in warmer water
(Thomas and Rice 1986a).
In southeast Alaska Dolly Varden are typically the first salmonid colonizers of new streams
formed by glacial retreat, suggesting they have lower requirements for microhabitat features
(e.g., pools) that are a function of stream age (Milner 1994). Because they often use small
isolated stream habitats and spawning populations can be small, both anadromous and
nonanadromous Dolly Varden are particularly vulnerable to barriers to migration (Dunham et al.
2008; Fausch et al. 2010; Kishi and Maekawa 2009; Koizumi 2011; Koizumi and Maekawa
2004) and to alterations of the small headwater streams in which they spawn and rear
(Armstrong and Morrow 1980, p. 133). The closely related bull trout S. confluentus is listed as
threatened in the contiguous United States (USFWS 1999), due in large part to habitat
fragmentation and warming stream temperatures.
Lake trout Salvelinus namaycush
Freshwater distribution and habitats
The global native distribution of lake trout is limited almost entirely to Canada and Alaska, from
the just south of the southern border of Canada north to the Canadian Arctic archipelago and
from Canada's eastern maritime provinces west to near the Bering Sea coast (Martin and Olver
1980, p. 209-210). This native range is almost entirely restricted to the limits of North American
late-Pleistocene glaciations (Lindsey 1964). In Alaska lake trout occur in suitable habitats across
most of the state except for southern southeast Alaska, much of western Alaska, and maritime
islands (Mecklenburg et al. 2002, p. 198), but within that broad range, there is great discontinuity
between occupied habitats (Lindsey 1964). Bristol Bay marks the westernmost limit of the lake
trout's native range (Mecklenburg et al. 2002, p. 198). Bristol Bay lake trout appear to be
restricted to upland lakes and their inlet and outlet streams (ADF&G 2011, FSN0616C03;
Burgner et al. 1965; Metsker 1967, p. 9, 11; Russell 1980, p. 47, 78, 79; Yanagawa 1967, p. 10).
They are common in the Tikchik Lake system but absent from the main Wood River lakes
(Burgner et al. 1965). Russell (1980, p. 77) considered them widely distributed in the Lake Clark
area and their diet indicated they fed at lake surfaces and bottoms, and through the water
columns. Anglers target lake trout in many BBWAA upland lakes, particularly Lake Clark,
Iliamna Lake, and the Tikchik Lakes (Minard et al. 1998, p. 152-155).
Almost all spawning occurs along lake shorelines or shoals, above coarse, often angular substrate
(Martin and Olver 1980, p. 218; Scott and Crossman 1998, p. 222; Viavant 1997, p. 6-7). Lake
trout typically spawn along exposed shorelines off points or islands or in mid-lake shoals (Martin
and Olver 1980, p. 218). Russell (1980, p. 77) reported apparent spawning habitat on shoals
around islands in Lake Clark. Spawning can occur in very spatially discrete locations (Viavant
1997, p. 6-7). Spawning areas appear to be kept clean of fine sediments by wind-driven or deep-
water currents and not by springs or seeps. The maximum depth of spawning may be positively
related to lake size, particularly fetch length, but is often less than 6 m (Martin and Olver 1980,
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p. 218; Royce 1951). In lakes that thermally stratify, lake trout may migrate seasonally from
warming surface waters to cool deep waters (Martin and Olver 1980, p. 228-230).
Life cycle
Compared to many other salmonids, lake trout exhibit little tendency toward anadromy
(Rounsefell 1958), but some individuals in far northern areas do migrate seasonally to marine
waters (Swanson et al. 2010). Like other char, lake trout is a highly variable species and multiple
forms, differing in diet, growth, and life span can occupy a single lake (Martin 1966). Adults can
live to at least 51 years (Keyse et al. 2007); in the BBWAA, lake trout are known to live at least
29 years, begin to reach maturity at about 6 years (Russell 1980, p. 77), reach lengths of at least
910 mm (FL; Wiedmer unpublished), and weights of at least 14.5 kg (Richard Russell, Alaska
Department of Fish and Game (retired), King Salmon, AK, personal communication). In some
southcentral Alaska lakes, lake trout mature at ages 7 to 10 at lengths of 450 to 550 mm (FL;
Van Whye and Peck 1968, p. 35). In the BBWAA and lakes in southcentral and interior Alaska,
lake trout spawn in mid- to late September and perhaps later (Russell 1980, p. 77; Van Whye and
Peck 1968, p. 35; Viavant 1997, p. 6). Mature lake trout, particularly those in more northern
habitats, may not spawn annually, but will skip one or two years between spawning events
(Martin and Olver 1980, p. 215). Most lake trout appear to home each year to specific spawning
sites, but not all do (Martin and Olver 1980, p. 218).
The number of ova produced by mature females is a function of size and perhaps stock; reported
average fecundities range from 996 to 15,842, and the diameter of ripe ova range from 3.7 to 6.8
mm (Martin and Olver 1980, p. 211, 213, 214). Lake trout may clean fine debris from the general
area of spawning locations, but they do not construct redds, nor cover or guard their fertilized
eggs (Royce 1951). Eggs and alevins incubate in spawning substrates until the following spring
(Martin and Olver 1980, p. 224). The movements of young-of-the-year fry are poorly
understood, but they are suspected to move to deeper water, often using the cover of coarse
substrates (Martin 1966, p. 224, 226; Royce 1951). Larger fish can be nomadic within their home
lake (Martin and Olver 1980, p. 226-227), and may move short distances between lakes (Scanlon
2010, p. 22). One probably mature, and apparently healthy 565 mm (FL) lake trout was captured
in mid-August in the Tikchik River approximately 14 km from the nearest large lake (ADF&G
2011, site FSN0616C03). As a result of spawning stress, some adults move from lakes
downstream into outlet rivers, and many likely do not survive to return to their natal waters
(Richard Russell, Alaska Department of Fish and Game (retired), King Salmon, AK, personal
communication).
Predator-prey relationships
In Lake Clark, growth remains fairly constant up to lengths of about 560 mm (FL), after which
the relationship between weight and length significantly increases. Metsker (1967) attributed this
to a transition, occurring at a length of about 480 mm (FL), from a diet of invertebrates to a diet
offish, primarily least cisco. A similar diet transition from insects and mollusks to fish, coupled
with a potential influence on growth rate, was observed in lake trout from lakes in southcentral
Alaska (Van Whye and Peck 1968, p. 30, 37).
Aquatic and terrestrial insects and small crustaceans are important foods for young-of-the-year
fry (Martin and Olver 1980, p. 234). In Alaskan lakes, Arctic grayling, sculpins, humpback,
round, and pygmy whitefish, least cisco, sockeye salmon fry, salmon eggs, ninespine stickleback,
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longnose suckers, Dolly Varden, Arctic char, rodents, shrews, and smaller lake trout are all prey
items for large lake trout (Plumb 2006, p. 29; Russell 1980, p. 81-83; Troyer and Johnson 1994,
p. 42; Van Whye and Peck 1968, p. 37). In the absence of fish prey, large lake trout in arctic
Alaskan lakes are generalist feeders and feed primarily on benthic invertebrates (Keyse et al.
2007). In the presence of large lake trout, small lake trout limit their use of available habitats to
avoid predation (Hanson et al. 1992; Keyse et al. 2007; McDonald and Hershey 1992).
In the laboratory, slimy sculpin consume lake trout eggs (Fitzsimons et al. 2006). In the wild
small lake trout (Royce 1951), are known to feed on lake trout eggs, as are round whitefish
(Loftus 1958), which are found throughout the BBWAA (ADF&G 2011). Royce (1951)
suspected humpback whitefish, which are found in many of the same BBWAA lakes as lake
trout, also feed on lake trout eggs. Burbot and large lake trout in the BBWAA feed on small lake
trout (Russell 1980, p. 67, 82-83). Power and Gregoire (1978) concluded that, of the all the
members of the fish community in Lower Seal Lake, Quebec, lake trout were the species most
affected by freshwater seal Phoca vitulina predation. In 1998, Small (2001) reported that Iliamna
Lake in the BBWAA supported a minimum harbor seal population of 321.
Abundance and harvest
In Bristol Bay total lake trout abundance is unknown, but in 2009 the BBWAA and the adjacent
Togiak River system supported an estimated sport catch of 3,651 (12% of the statewide total)
and harvest of 588 (11% of the statewide total; Jennings et al. 2011, p. 72). Dye and Schwanke
(2009, p. 6) speculated that the trend of decreasing sport harvests are due in part to increasing
catch-and-release practices.
In the mid-1960s, Iliamna Lake and Lake Clark supported a commercial winter lake trout fishery
(Metsker 1967, p. 8, 10). In 1966 and 1967 Tikchik Lake also supported an experimental
commercial freshwater fishery (Yanagawa 1967). Lake trout were the second-most commonly
harvested species in that fishery, representing 30% of the overall harvest. The Tikchik Lake
fishery harvested 1,502 fish, which averaged 2.2 kg in weight, and ranged in length from 500 to
575 mm and in age to more than 15 years (Yanagawa 1967).
In the mid-2000s, villagers from ten of the BBWAA communities annually harvested, as part of
their subsistence activities, about an estimated 1,030 lake trout (Fall et al. 2006, p. 45, 80, 113,
150, 194; Krieg et al. 2009, p. 40, 78, 118, 162, 202). From the mid-1970s to the mid-2000s, lake
trout were estimated to represent between 4.6 and 11.8% of the total weight of the Kvichak River
drainage non-salmon freshwater fish subsistence harvest (Krieg et al. 2005, p. 214).
Stressors
As with lake-spawning humpback whitefish, excessive variation in lake level is suspected to
reduce egg and alevin survival (Martin and Olver 1980, p. 223). Sedimentation of lake spawning
areas has resulted in declines or elimination of successful reproduction (reviewed in Martin and
Olver 1980, p. 223-224). In nature, lake trout are reported in water temperatures ranging from
-0.8 to 18 °C, appear to prefer summer temperatures around 6 to 13 °C (Martin and Olver 1980.
p. 230-231), and to have an upper lethal temperature of approximately 23.5 °C (Gibson and Fry
1954). Martin and Olver (1980, p. 231) concluded that a DO level of approximately 4 mgl -1 is
the minimum tolerated by lake trout. Late maturity, long life, and slow growth make lake trout
particularly vulnerable to over-harvest (Martin and Olver 1980, p. 259). Like the similarly long-
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lived piscivore, northern pike, lake trout bioaccumulate and biomagnify atmospherically
deposited mercury (Swanson et al. 2011). Lake acidification has extirpated lake trout from some
Canadian lakes (Matuszek et al. 1992).
Arctic grayling Thymallus arcticus
Freshwater distribution and habitats
Arctic grayling are found in fresh waters of higher latitudes of the Northern Hemisphere, from
Hudson Bay west across the Bering Strait to the Ob and Kara river drainages east of Asia's Ural
Mountains. In North America, the current native distribution of Arctic grayling is almost entirely
restricted to northwestern Canada and Alaska (Scott and Grossman 1998, p. 301, 302). Arctic
grayling native to northern Michigan were extirpated by around 1936 (Scott and Grossman 1998,
p. 301), and by the 1990s their former broad distribution in streams of the Upper Missouri River
were limited to the Big Hole River in southwestern Montana (Lohr et al. 1996). In Alaska, the
Arctic grayling native range stretches across the entire mainland, but they are absent from most
islands, except those formerly part of the Bering land bridge (Morrow 1980b, p. 145-146).
Throughout their range, Arctic grayling are primarily restricted to fresh waters. Along the Arctic
Ocean coast, they will descend downstream to feed in nearshore marine waters, but they appear
to remain in the low salinity plume at the mouths of rivers or in lagoons (Furniss 1975; Tack
1980, p. 26).
Arctic grayling are widely distributed in Bristol Bay lakes (Burgner et al. 1965; Russell 1980, p.
49, 57; Yanagawa 1967, p. 12) and streams (Coggins 1992). They can occur in slow-flowing
lowland streams where salmon, rainbow trout, and Dolly Varden are absent (ADF&G 2011), but
they do not occur in many of the small shallow ponds on the coastal plain (Hildreth 2008, p. 9).
Their range does not extend quite as far up the BBWAA's higher gradient headwater streams as
do Dolly Varden and rearing coho salmon, but they are found, at some time of the year, in most
tributaries and downstream to the lower Nushagak River (ADF&G 2011; Krieg et al. 2009, p.
383). Sport anglers catch Arctic grayling across most of the BBWAA, with a particular focus on
the Kvichak, Alagnak, Newhalen, Tazimina, Nushagak, Mulchatna, and Koktuli rivers, Lake
Clark, and the Wood River and Tikchik lake systems (Minard et al. 1998, p. 189).
BBWAA stream spawning locations may represent sites that provide both warm spring and
summer temperatures and suitable hydrology (Tack 1980, p. 3-4, 14-16, 27; Warner 1957). Some
spawning may occur in lakes, at stream outlets (Warner 1957). Arctic grayling and rainbow trout
are the only spring-spawning salmonids in the BBWAA, and both likely seek spawning sites that
enhance incubation rates and early fry growth. Tack (1980, p. 14) reported that most interior
Alaska spawning occurred in riffles with sand and gravel substrates and minimal silt, in currents
ranging from 0.25 to 1 ms"1. Reed (1964, p. 14) concluded that Alaskan Arctic grayling did not
target specific spawning substrates.
Best egg survival in the closely-related European grayling T. thymallus was 6 to 13.5 °C
(Jungwirth and Winkler 1984). For much of the summer, age-0 fish tend to remain near the sites
where they emerged from the spawning substrate (Craig and Poulin 1975; MacPhee and Watt
1973, p. 14, 15; Tack 1980, p. 27; Tripp and McCart 1974, p. 56). Given the August distribution
of age-0 fry in the Nushagak-Mulchatna drainage (ADF&G 2011), it appears that most Arctic
grayling spawning in this system occurs in tributaries.
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When food is not limiting, optimal growth for age-0 juveniles in interior Alaska is at about 17 °C
(Dion and Hughes 2004; Mallet et al. 1999). Older age classes may segregate to different
habitats (Craig and Poulin 1975; Tack 1980, p. 29; Vincent-Lang and Alexandersdottir 1990, p.
50), but the details of that segregation may depend on the drainage-specific patterns of water
temperature and food availability (Hughes 1998).
After spawning, adults may migrate further upstream (Hughes and Reynolds 1994; Vascotto
1970, p. 77; Wojcik 1954), or descend back to the mainstem (Craig and Poulin 1975; MacPhee
and Watt 1973, p. 14; Tripp and McCart 1974, p. 49-51; Warner 1957), often using the same
summer feeding areas annually (Ridder 1998, p. 17; Tack 1980, p. 21). Juveniles age 1 and older
often follow adults, perhaps to imprint the complex migratory routes (Tack 1980, p. 20). In
interior and southcentral Alaska, adult Arctic grayling overwinter in deep lakes and large rivers
(Reed 1964, p. 13; Ridder 1998, p. 10-15; Sundet and Pechek 1985, p. 44; Tack 1980, p. 8, 28).
Available evidence suggests the same pattern applies in the Nushagak River drainage. In August,
Arctic grayling are absent or uncommon in the lower mainstem of the Nushagak River (ADF&G
2011). However, in this same area, local residents harvest large numbers of Arctic grayling
through the ice during winter (Krieg et al. 2009, p. 220, 383).
Life cycle
Arctic grayling are nonanadromous, but often do undertake extensive seasonal migrations. Prior
to spring breakup, large fish concentrate in mainstem rivers, at the mouths of tributaries. During
and immediately after breakup, fish begin entering tributaries, even below ice cover and through
channels on the ice surface (Reed 1964, p. 12-13; Warner 1957). In at least parts of Alaska, the
upstream migration correlates with the peak of the spring freshet (Tack 1980, p. 13) and adults
appear to show some fidelity to spawning areas (Craig and Poulin 1975; Tack 1980, p. 27).
BBWAA Arctic grayling spawn in May through early June, shortly after breakup (Dye 2008, p.
26; Russell 1980, p. 57).
Mature female fecundity probably averages between about 4,000 and 7,000 ova, with some large
fish producing much more (Scott and Grossman 1998, p. 303). Water-hardened eggs have an
average diameter of around 3 mm and are non-adhesive (Reed 1964, p 14). Spawning adults do
not actively construct redds (Craig and Poulin 1975), but their actions may create slight
depressions in the stream substrate (Reed 1964, p. 13-14). Fertilized eggs fall into interstitial
spaces, hatch in 2 to 3 weeks at lengths of about 8 mm (Scott and Grossman 1998, p. 303), and
fry start feeding a few days later (Morrow 1980b, p. 146). Some BBWAA age-0 fish are free-
swimming in early June, and perhaps even earlier in certain locations (Russell 1980, p. 57). Early
growth rates appear related to temperature and benthic invertebrate densities (Tripp and McCart
1974, p. 21); on Alaska's North Slope, growth rates of age-0 Arctic grayling correlate positively
to stream temperature (Luecke and MacKinnon 2008).
In the BBWAA, age-0 fish reach a mean fork length of about 69 mm (n = 700, SD = 13.6 mm)
by August (calculated from data provided by ADF&G 2011). After age 0, BBWAA Arctic
grayling grow about 47 mrry -1 until age 5 when growth begins to slow (Russell 1980, p. 60).
Fish begin maturing at lengths of about 300 mm (FL), and once mature, grayling appear to
spawn every year (Craig and Poulin 1975; Engel 1973, p. 8; Tripp and McCart 1974, p. 34).
Bristol Bay Arctic grayling mature around age 5 (Russell 1980, p. 57), can live up to at least 13
years (Plumb 2006, p. 56), reach lengths of at least 650 mm (FL; MacDonald 1995, Table 7) and
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weights at least 0.9 kg (Russell 1980, p. 57). Alaskan Arctic grayling may travel over 320 km
between spawning, summer feeding, and overwintering locations (Reed 1964, p. 13; Ridder
1998, p. 10; Tripp and McCart 1974, p. 53).
Predator-prey relationships
Arctic grayling appear to feed on whatever is available to them, primarily aquatic and terrestrial
insects, sequentially taking advantage of temporary peaks of abundance of different invertebrate
populations (Plumb 2006, p. 62; Reed 1964, p. 20; Scheuerell et al. 2007; Tripp and McCart
1974, p. 60-61). Arctic grayling typically feed at the surface and mid-depth in the water column
(Vascotto 1970), but food items include benthic slimy sculpin and slimy sculpin eggs (Bond and
Becker 1963) and humpback whitefish eggs (Kepler 1973, p. 71). Scheuerell et al. (2007)
discovered that, in the Wood River lakes system, after the arrival of spawning sockeye salmon,
the energy intake of Arctic grayling increases more than five-fold, due primarily to the increased
availability of benthic invertebrates. As spawning salmon construct redds and bury fertilized
eggs, they disturb the substrate, displacing benthic macroinvertebrates, thus making them more
available to Arctic grayling predation. In addition, Arctic grayling feed on salmon eggs and the
larval blowflies that colonize salmon carcasses. These salmon-derived resources contribute a
large majority of the energy necessary for the annual growth of resident Arctic grayling
(Scheuerell et al. 2007). In lakes, Arctic grayling can be the most important prey species of lake
trout (Troyer and Johnson 1994, p. 42). In Alaskan Arctic streams, Stevens and Deschermeier
(1986) found that some juvenile Dolly Varden eat age-0 Arctic grayling fry .
Abundance and harvest
In Bristol Bay total Arctic grayling abundance is unknown, but in 2009 the BBWAA and the
adjacent Togiak River drainage supported an estimated sport fish catch of 44,762 fish (11% of
the statewide total) and a harvest of 1,094 (4% of the statewide total; Jennings et al. 2011, p. 74).
Dye and Schwanke (2009, p. 6) speculated that the trend of decreasing sport harvests are due in
part to increasing catch-and-release practices.
In the mid-2000s, villagers from nine of the BBWAA communities annually harvested, as part of
their subsistence activities, about an estimated 7,790 Arctic grayling (Fall et al. 2006, p. 45, 80,
113, 150, 194; Krieg et al. 2009, p. 40, 78, 118, 162, 202). From the mid-1970s to the mid-
20008, Arctic grayling were estimated to represent between 6.9 and 9.7% of the total weight of
the Kvichak River drainage non-salmon freshwater fish subsistence harvest (Krieg et al. 2005, p.
214).
Stressors
Total dissolved solids up to 2,782 mg-F1 do not have a significant impact on Arctic grayling egg
fertilization; however, concentrations as low as 1,402 mg-1 l do significantly affect water
absorption during the water-hardening phase immediately following fertilization, when embryos
gain resistance to mechanical damage (Brix et al. 2010). As a result, Brix et al. (2010) identified
that period as the most sensitive early developmental stage.
Egg mortality in the closely-related European grayling T. thymallus was 100% at temperatures
over 16 °C or under 4 °C (Jungwirth and Winkler 1984). In interior Alaska, the minimum and
maximum temperatures at which growth occurs are 4.5 °C and 21 °C (Dion and Hughes 2004;
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Mallet et al. 1999). In interior Alaska, age-0 fish are more tolerant of high water temperatures
than alevins and older juveniles, with a median tolerance limit in excess of 24.5 °C, compared to
20 to 24.5 °C for the other life stages (LaPerrier and Carlson 1973, p. 29).
In North Slope streams, the growth of age-0 fry is positively correlated with temperature, while
adult growth has no temperature correlation (Deegan et al. 1999; Luecke and MacKinnon 2008).
Adult and age-0 juveniles may also respond differently to stream discharge. Adult growth in
North Slope streams is positively correlated with discharge, while age-0 growth is negatively
correlated with it (Deegan et al. 1999; Luecke and MacKinnon 2008). Wojick (1954, p. 67)
speculated that elevated stream discharges during the incubation and early fry rearing stage
would harm Arctic grayling stocks.
Although reasons for the dramatic contraction in the native range of stream-resident Upper
Missouri River Arctic grayling is not well understood, constructed barriers to fish migration and
stream dewatering appear to be major contributing factors (Barndt and Kaya 2000).
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Table 1. Fish species reported in the EPA Bristol Bay Watershed Assessment Area (ADF&G 2011; Mecklenburg et al.
2002; Morrow 1980).
Scientific/Common
Family Name
Common Name
Scientific Name
Principal Life History'
Petromyzontidae/lampreys
Clupeidae/herrings
Catostomidae/suckers
Esocidae/pikes
Umbridae/mudminnows
Osmeridae/smelts
Salmonidae/salmonids
Gadidae/cods
Gastero steidae/sticklebacks
Cottidae/sculpins
Pleuronectidae/righteye flounders
Arctic lamprey
Alaskan brook lamprey
Pacific lamprey
Pacific herring
longnose sucker
northern pike
Alaska blackfish
rainbow smelt
pond smelt
eulachon
Bering cisco
humpback whitefish
least cisco
pygmy whitefish
round whitefish
coho salmon
Chinook salmon
sockeye salmon
chum salmon
pink salmon
rainbow trout
Arctic char
Dolly Varden
lake trout
Arctic grayling
burbot
Pacific cod
saffron cod
threespine stickleback
ninespine stickleback
coastrange sculpin
slimy sculpin
Pacific staghom sculpin
Arctic flounder
starry flounder
Lethenteron camtschaticum2
L. alaskense
Entosphenus tridentatus2
Clupea pallasii
Catostomus catostomus
Esox lucius
Dallia pectoralis
Osmerus mordax
Hypomesus olidus
Thaleichthys pacificus
Coregonus laurettae
C. pidschian
C. sardinella
Prosopium coulterii
P. cylindraceum
Oncorhynchus kisutch
O. tshawytscha
O. nerka
O. keta
O. gorbuscha
O. mykiss
Salvelinus alpinus
S. malma
S. namaycush
Thymallus arcticus
Lota lota
Gadus macrocephalus
Eleginus gracilis
Gasterosteus aculeatus
Pungitius pungitius
Cottus aleuticus
C. cognatus
Leptocottus armatus
Pleuronectes glacialis
Platichthys stellatus
Anadromous
Nonanadromous
Anadromous
Amphidromous
Nonanadromous
Nonanadromous
Nonanadromous
Anadromous
Nonanadromous
Anadromous
Nonanadromous and Anadromous3
Nonanadromous and Anadromous3
Nonanadromous and Anadromous
Nonanadromous
Nonanadromous
Anadromous
Anadromous
Anadromous
Anadromous
Anadromous
Nonanadromous
Nonanadromous
Nonanadromous and Anadromous
Nonanadromous
Nonanadromous
Nonanadromous
Amphidromous
Amphidromous
Nonanadromous and Anadromous
Nonanadromous
Nonanadromous
Nonanadromous
Amphidromous
Amphidromous
Amphidromous
1 Anadromous: fishes that spawn in fresh waters and migrate to marine waters to feed; Nonanadromous: fishes that spend their entire life in
fresh waters, with possible migrations between habitats within a drainage (resident fish); Anadromous and
Nonanadromous/Nonanadromous and Anadromous: fish populations in which some individuals have anadromous life histories and some
have potamodromous life historiesthe most common form listed first; Amphidromous: fishes that may move between fresh and salt water
during some part of life cycle, but not for spawning. The Bristol Bay fishes classified amphidromous are primarily marine species that may
move into estuaries and lower rivers for feeding, particularly as juveniles.
2 Nomenclature follows Brown et al. (2009).
3 Anadromy known in other Alaskan locations, but not verified within the BBWAA.
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Appendix C k%
Wildlife Resources of the Nushagak
and Kvichak River Watersheds
C-l
-------
U. S. Fish and Wildlife Service
Report to the
U.S. Environmental Protection Agency
for the
Bristol Bay Watershed Assessment
Wildlife Resources of the
Nushagak and Kvichak River Watersheds
External Review Draft
January 2012
Edited by: Philip J. Brna and Lori A. Verbrugge
Anchorage Fish and Wildlife Field Office
FU.S.
FISH & WILDLIFE
SERVICE
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ACKNOWLEDGEMENTS
This report is the result of the work of many people and all those who contributed are listed in
Appendix 1. If we forgot to acknowledge anyone we sincerely apologize. Kirsti Sanchez and
Greg Aull, who were 2011 summer hires in the Anchorage Fish and Wildlife Field Office
(AFWFO), completed much of the initial literature review and prepared initial drafts of some of
the species accounts. Retired biologists Colleen Matt, Ken Whitten, Chuck Schwartz, Lowell
Suring, and Tom Rothe agreed to assist with the project under small contracts by reviewing and
improving species accounts for brown bear, caribou, moose, bald eagle, and waterfowl,
respectively. U.S. Fish and Wildlife Service (USFWS) Alaska Peninsula-Becharof National
Wildlife Refuge (NWR) Biologist Susan Savage prepared the shorebirds species account and
USFWS AFWFO biologist Maureen de Zeeuw prepared the landbirds account and the initial
draft of the bald eagle account and they collaborated on the species list. Marcus Geist of The
Nature Conservancy provided GIS support including maps, watershed area calculations and land
cover information and analysis. Numerous Federal and State agency wildlife biologists
generously gave time and effort to review and improve sections of the report. Of special note are
United States Geological Survey (USGS) biologist Layne Adams, National Park Service (NFS)
biologists Buck Mangipane and Grant Hildebrand, retired NFS biologist Page Spencer, and
USFWS biologists Andy Aderman (Togiak NWR) and Dom Watts (Alaska Peninsula-Becharof
NWR) who provided significant reviews, edits, information and insight based on their
considerable scientific expertise and knowledge. Heather Dean of the Environmental Protection
Agency provided thorough and impressive editing. Her attention to detail makes this a far more
consistent and readable report. Finally, this project would not have been possible without the
support and positive encouragement from Ann Rappoport, AFWFO Field Supervisor.
This document should be cited as:
Brna, Philip J. and Verbrugge, Lori A. (Eds.) 2012. Wildlife resources of the Nushagak and
Kvichak River watersheds. Report to the U.S. Environmental Protection Agency for the Bristol
Bay Watershed Assessment. Anchorage Fish and Wildlife Field Office, U.S. Fish and Wildlife
Service, Anchorage Alaska. 160 pp.
For information on this report contact:
USFWS/AFWFO
605 W. 4th Avenue, Room G-61
Anchorage, Alaska 99501
907-271-2888
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EXECUTIVE SUMMARY
Background
At the request of the U.S. Environmental Protection Agency (EPA), the U.S. Fish and Wildlife
Service (USFWS) prepared this report which summarizes known information related to brown
bear, moose, caribou, wolf, waterfowl, bald eagle, shorebirds, and landbirds in the Bristol Bay
region of Alaska, with a focus on the Nushagak and Kvichak watersheds. These species were
selected for review because of their importance to ecosystem function, their direct link to
salmon, or their importance to local and Alaska residents. EPA is conducting a watershed
assessment in the Nushagak and Kvichak watersheds in response to requests from various
organizations under the authority of the Clean Water Act, and requested assistance from the
USFWS, as the agency with responsibilities and expertise for the nation's fish and wildlife
resources. This report is a small portion of the larger EPA effort which includes evaluation of
fish resources of the region, hydrology, and an ecological risk assessment related to potential
effects of large scale mineral development on fish, wildlife, water quality, and humans. In
addition to being part of the EPA Bristol Bay Watershed Assessment, the information in this
report will be useful for Statewide or regional land use planning, completion of environmental
documentation for permitting of development projects, or activities related to Landscape
Conservation Cooperatives in Alaska.
In this report, we describe: habitat use; food habits; behavior; interspecies interactions;
productivity, mortality and survivorship; populations, subpopulations, and genetics; human use
and interactions, and management for wildlife with a focus on the Nushagak and Kvichak
watersheds, to the extent that existing data allow. We describe the relationship of these wildlife
species (brown bear, moose, caribou, wolf, and bald eagle) or species guilds (waterfowl,
shorebirds and landbirds) with salmon. We describe the dependence of wildlife on marine
derived nutrients (MDN) transported to these watersheds by salmon and the role of wildlife in
distributing MDN through the ecosystem to the extent this information is available.
About 40 current or retired biologists and scientists from USFWS, National Park Service, Bureau
of Land Management, U.S. Geological Survey, U.S. Forest Service, and Alaska Department of
Fish and Game were involved with preparation and review of this report. Collectively, these
biologists and scientists have significant experience with research and management of wildlife in
Alaska and many have extensive experience in the Nushagak and Kvichak watersheds.
The Importance of Marine-Derived Nutrients to Bristol Bay
One of the most important ecological functions of salmon is to transfer large quantities of
nutrients from the marine environment into terrestrial and freshwater ecosystems within the
watersheds where adults return to spawn. Marine-derived nutrients (MDN), in combination with
other ecosystem features such as suitable spawning habitat and oceanic carrying capacity, are
essential for the survival and growth of the next generation of salmon, and also greatly benefit
other fish and wildlife species. Herbivores benefit from increased vegetative growth in riparian
areas stimulated by MDN, while carnivores and scavengers directly consume migrating salmon
or their carcasses. In both aquatic and terrestrial ecosystems, MDN are also integrated into the
base of food chains, and increased productivity is transferred to species at higher trophic levels.
It is likely that southwest Alaska and the Nushagak and Kvichak watersheds support large
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populations of wildlife both because habitat in the region is almost totally intact and because of
the presence of millions of salmon annually spawning, rearing, and migrating to sea and back.
The annual introduction of millions of pounds of MDN from salmon and the lack of significant
anthropogenic watershed disturbances make Bristol Bay relatively unique in the world.
Brown bears within the Nushagak and Kvichak watersheds depend on salmon for food and
survival. Accumulation of fat reserves is important for successful hibernation, and for female
reproductive success. Wolves also consume salmon seasonally when available, and this marine-
derived source of food is a major component of the lifetime total diet of wolves in areas where
the two species co-exist. Brown bears and wolves play an important role in distributing MDN
from streams to the terrestrial environment, both by transporting salmon carcasses prior to
consumption and through excretion of wastes rich in salmon nutrients. Both brown bears and
wolves have been documented travelling long distances to feed on salmon.
Waterfowl benefit from salmon which provide a large influx of nutrients to riverine and
terrestrial systems, both directly as sources of prey and carrion, and indirectly as nutrient drivers
of aquatic systems. Of the 24 duck species that regularly occur in Bristol Bay, at least eleven
species are known to prey on salmon eggs, parr, smolts, and scavenge on carcasses. Of these,
greater and lesser scaup, harlequin duck, bufflehead, common and Barrow's goldeneyes, and
common and red-breasted mergansers exhibit directed foraging on salmon. Among dabbling
ducks, mallards feed most on salmon because they are distributed across a diversity of summer
habitats in spawning areas, and they are the principal wintering dabbling duck on the North
Pacific coast where fall-winter salmon runs occur. Fish predators like mergansers feed
extensively on salmon fry and smolts. Other duck species may prey on smolt incidentally.
Salmon eggs are a seasonally rich food source for harlequin ducks, goldeneyes and scaup that
frequent rivers and streams and probably for other opportunistic ducks. Species ranging from
dabbling ducks (mallard, green-winged teal) and diving ducks to sea ducks that inhabit spawning
waters probably opportunistically scavenge easy protein-rich meals from salmon carcasses.
Spawned-out salmon carcasses provide an ideal food resource to bald eagles. The abundance of
salmon in the region affects bald eagle population size, distribution, breeding and behavior. In
one studied population, salmon availability in spring was tightly correlated with whether adult
bald eagles laid eggs in a given year, and also influenced the timing of egg-laying. As with other
salmon consumers, bald eagles affect the ecosystems within their range by distributing MDNs in
their excretions.
Direct or indirect interactions between shorebirds and salmon are not well-documented. Some
shorebird species are observed to consume dead salmon and salmon eggs. No studies have been
conducted to deduce the contribution of salmon to the energetics of shorebird populations;
however, the abundance of invertebrates in the intertidal zone is very likely due in part to MDN
from salmon that die on the coast and in the rivers feeding Bristol Bay. Shorebirds play a role in
distributing MDNs into the terrestrial system, especially during the migratory period, but this has
not been quantified. Some of the longest migrations known for birds involve shorebird species
(bar-tailed godwit) that use Bristol Bay intertidal areas in autumn. Such flights are possible not
only due to the extreme abundance of intertidal invertebrates (polychaetes, crustaceans,
gastropods, and bivalves) in the region, but also because the adjacent uplands are usually rich in
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fruits of ericaceous plants or tubers that birds like plovers, whimbrels, and godwits, regularly
feed on. Some species like whimbrel, Hudsonian godwit, and black and ruddy turnstones feed on
herring roe, carrion (including salmon carcasses), and salmon eggs. Many shorebirds make use
of freshwater invertebrates; small fish may be consumed by yellowlegs and phalaropes.
Landbirds also benefit from salmon carcasses when available. Aquatic invertebrate larvae feed
on salmon carcasses, overwinter in the soil, and emerge in the spring as adults. These
invertebrate adults become prey for a variety of landbird species in the spring, and serve as an
important seasonal subsidy during a period when terrestrial invertebrate biomass is low. Salmon
also benefit landbirds by increasing plant productivity due to MDN inputs, potentially resulting
in an abundance of berries and seeds for avian consumption. Some birds, such as the American
Dipper, directly consume salmon eggs, fry, and small bits of carcasses when available.
In summary, MDN from salmon cycle throughout the ecosystem of watersheds with healthy
salmon runs, benefitting wildlife, increasing vegetation productivity, and promoting the
production of periphyton, aquatic macroinvertebrates, resident freshwater fish, and juvenile
salmon. This nutrient cycling is in turn dependent on interactions with wildlife, which distribute
MDN into the terrestrial environment through both transport of carcasses and excretion of
wastes. The loss of either salmon or key wildlife species may result in significant changes to the
productivity, diversity and physical structure of the ecosystem, via mechanisms that extend
beyond simple "food chain" interactions.
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CONTENTS
ACKNOWLEDGEMENTS ii
EXECUTIVE SUMMARY iii
Background iii
The Importance of Marine-Derived Nutrients to Bristol Bay iii
LIST of FIGURES x
LIST of TABLES x
INTRODUCTION 11
METHODOLOGY 12
Geographic Scope of USFWS Evaluation 13
Selection of Wildlife Species for Characterization 13
The Characterization Process 15
CHARACTERIZATION OF WILDLIFE 15
Overview of Wildlife Species 15
Alaska- 15
Southwest Alaska Region- 16
The Importance of Marine-Derived Nutrients to Bristol Bay Watershed Ecosystems 16
Overview of Land Cover and Habitat Types 19
BROWN BEARS 21
Introduction 21
Habitat 21
Food Habits 22
Behavior 23
Interspecies Interactions 25
Mortality, Productivity, and Survivorship 25
Population 27
Human Use/Interaction/Management 28
Sport Hunting for Brown Bears 28
Game Management Unit 9- 28
Game Management Unit 17- 29
Subsistence Hunting for Brown Bears- 29
Bear Viewing- 30
Other Human-Bear Interactions 30
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Local Residents and Bear-Human Conflicts- 30
Other Recreational Users and Bear-Human Conflicts- 30
MOOSE 31
Introduction 31
Habitat 31
Winter Habitat- 34
Food Habits 35
Behavior 37
Movements and Home Ranges- 37
Sexual Segregation and Grouping Behaviors- 40
Mating and Maternal Behaviors- 41
Activity Budgets- 42
Interspecies Interactions 43
Mortality, Productivity, and Survivorship 45
Population, Subpopulations, and Genetics 48
Human Use (Subsistence, Recreation)/Interaction/Management) 49
BARREN GROUND CARIBOU 51
Introduction 51
Population History of Caribou in the Upper Bristol Bay Region 51
Habitat 52
Seasonal Preference- 52
Food Habits 52
Spring- 52
Summer- 53
Fall- 53
Winter- 53
Behavior 53
Seasonal Range Use and Migrations- 53
Response to Disturbance- 54
Interspecies Interactions 55
Mortality, Productivity, Survivorship 55
Mortality- 55
Breeding- 55
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Human Use/Interaction/Management 56
WOLF 58
Introduction 58
Habitat 58
Food Habits 58
Diet- 58
Salmon as a Food Source 59
Dispersal of Marine-Derived Nutrients (MDNs) by Wolves- 60
Behavior 61
Wolf Packs- 61
Range- 61
Dispersal (Emigration)- 62
Seasonal Movements- 63
Interspecies Interactions; Response to Change in Salmon Populations/ Distribution 63
Mortality, Productivity, and Survivorship 65
Population Estimates 66
Human Use/Interaction/Management 67
WATERFOWL 69
Introduction 69
Regional Overview- 69
History of Waterfowl Surveys- 70
Waterfowl Resources and Seasonal Occurrence 71
Estuaries and Inner Bristol Bay- 71
Bristol Bay Lowlands- 75
Inland Tundra/Taiga- 78
Nutrients, Trophic Relations and Foods 80
Nutrients and Habitat Productivity- 80
Food Habits- 81
Importance of Marine-Derived Nutrients (Salmon and Herring) to Waterfowl- 84
Populations, Subpopulations, and Genetics 85
Swans- 86
Geese- 86
Dabbling and Diving Ducks- 86
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Sea Ducks- 86
Human Use 87
Nonconsumptive Uses- 87
Recreational Harvest- 88
Subsistence Harvest- 88
BALD EAGLES 91
Introduction 91
Habitat 91
Food Habits 93
Diet- 93
Significance of MDNs- 94
Foraging Methods- 95
Behavior 95
Territoriality- 95
Flocking- 96
Migration and Local Movements- 96
Interspecies Interactions 97
Breeding, Productivity, and Mortality 97
Breeding- 97
Productivity and Survivorship- 98
Mortality- 99
Population, Distribution, and Abundance 101
Human Use 102
SHOREBIRDS 104
Introduction 104
Habitat 106
Food Habits 108
Behavior (Movements) 109
Interspecies Interactions 110
Population, Subpopulations, and Genetics 112
Human Use and Threats 113
LANDBIRDS 114
Introduction 114
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Habitat 114
Interspecies Interactions 115
APPENDIX 1: LIST OF AUTHORS AND REVIEWERS 117
APPENDIX 2: SOUTHWEST ALASKA TERRESTRIAL VERTEBRATE SPECIES 119
APPENDIX 3: LITERATURE CITED 128
LIST of FIGURES
Figure 1. Nushagak and Kvichak River watersheds and inner Bristol Bay 12
Figure 2. National Land Cover Dataset, land cover types for the Nushagak River watershed and
Kvichak River watershed 19
Figure 3. Nushagak and Kvichak River watersheds and associated Alaska Department of Fish
and Game, game management units 29
LIST of TABLES
Table 1. Size of subbasins (sq. mi.) within the Nushagak and Kvichak River watersheds and
percent of watershed or subbasin within each National Land Cover Database land cover type.. 20
Table 2. Mean (range) home range size (km2) for selected moose populations in Alaska 39
Table 3. Moose activity budgets in winter1 and spring/summer2 in Denali National Park and
Preserve (averages) 43
Table 4. Subsistence statistics for moose harvest in Alaska Dept. of Fish and Game, game
management units (GMU) 9B, 17B, and 17C 50
Table 5. Mulchatna caribou herd- estimated population size and harvest 56
Table 6. Average territory sizes (km2) of wolf packs in Alaska* 62
Table 7. Total wolf harvests in GMUs 9, 10 and 17 reported to Alaska Department of Fish and
Game 67
Table 8. Average abundance indices and densities of species/groups recorded in late May on the
Alaska Yukon Waterfowl Breeding Population Survey, Bristol Bay Lowlands (Stratum 8) 76
Table 9. General food habits and consumption offish by duck species of Bristol Bay 82
Table 10. Reported survival of bald eagle nestlings in Alaska 100
Table 11. Summary of surveys for bald eagle nests in the Bristol Bay study area 102
Table 12. Shorebirds found in the Bristol Bay Watershed 105
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INTRODUCTION
The U.S. Environmental Protection Agency (EPA) is conducting a watershed assessment in the
Nushagak and Kvichak River watersheds of Bristol Bay, Alaska in response to requests from nine
Tribal Governments and other interests. The tribes have requested that EPA take action to protect
the Bristol Bay watershed from the adverse impacts of potential large-scale hard rock mining
projects utilizing EPA's statutory authority, including Section 404(c) of the Clean Water Act
(CWA). Section 404(c) allows EPA to prohibit or restrict discharges of dredged or fill material
into waters of the United States, including wetlands, when it determines that such discharges
would have an unacceptable adverse effect on municipal water supplies, shellfish beds and fishery
areas (including spawning and breeding areas), wildlife, or recreational areas. EPA has also
received requests from two tribes and other interests to refrain from taking advance action and to
wait for specific permit applications for mining projects to be reviewed by the U.S. Army Corps of
Engineers (Corps) pursuant to Section 404 of the CWA and the National Environmental Policy
Act (NEPA). EPA is conducting the Bristol Bay Watershed Assessment (BBWA) under its Clean
Water Act Section 104(a) authority, which directs them to: "...conduct and promote the
coordination and acceleration of, research, investigations, experiments, training, demonstrations,
surveys, and studies relating to the causes, effects, extent, prevention, reduction, and elimination
of pollution."
EPA requested that the U.S. Fish and Wildlife Service (USFWS) provide assistance in conducting
the BBWA. The mission of the USFWS is to work with others to conserve, protect and enhance
fish, wildlife, and plants and their habitats for the continuing benefit of the American people. The
USFWS works to protect a healthy environment for people, fish and wildlife, and helps Americans
conserve and enjoy the outdoors and our living treasures. The USFWS has responsibility for fish
and wildlife resources with specific emphasis on migratory birds, endangered plants and animals,
certain marine mammals, and freshwater and anadromous fish. USFWS expects that this report
will be useful to EPA in completing the BBWA, but it should also provide a comprehensive
summary of wildlife information for others interested in southwest Alaska. These uses might
include: completion of environmental documentation for oil and gas leasing or any development
project; activities related to the Western Alaska Landscape Conservation Cooperative; the
Southwest Alaska Salmon Habitat Partnership; or any land use planning effort.
The Bristol Bay watershed is comprised of six major drainages: the Togiak River, Nushagak
River, Kvichak River, Naknek River, Egegik River, and Ugashik River. The Kvichak and
Nushagak River watersheds are the principle Bristol Bay drainages that have lands open to large-
scale development. Much of the rest of the region is within National Parks or National Wildlife
Refuges and is protected from such development. EPA's analysis therefore, focuses on the
Nushagak and Kvichak watersheds (Figure 1).
The objective of the BBWA process is to build a common understanding of both the fish and
wildlife resources of Bristol Bay and the potential impacts to those resources from large-scale
development and to identify possible options for protecting these resources. The overall
assessment represents an integration of several types of evaluations. The first component is an
assessment of the resources themselves, also called a characterization, which synthesizes current
conditions within the Nushagak and Kvichak watersheds and compares those resources to other
regional or reference areas. It's focus is determining if the resources in question, the wildlife of the
11
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Nushagak and Kvichak watersheds in the case of this report, represent an exceptional resource that
might be worthy of special protection. For exceptional resources, the characterization identifies the
environmental factors that contribute to the extraordinary nature of the resource. The
characterization thus identifies what must be protected to retain an exceptional status. The second
component of the evaluation is a predictive risk assessment, devoted, in this case, to estimating the
effects of reasonably foreseeable large-scale development in the area on wildlife. It is organized
on the established EPA frameworks for ecological and cumulative risk assessments. The potential
development scenarios and the results of the predictive risk assessment include inherent
uncertainties and cumulative risks.
Nushagak-Mulcnatna
Watershed
14,077 sq. miles
Kvichak
Watershed
9,467 sq. miles
Cape Constantine
Bristol Bay
Figure 1. Nushagak and Kvichak River watersheds and inner Bristol Bay.
Miles
10 20
METHODOLOGY
There are two phases of the EPA BBWA: characterization and predictive risk assessment.
Characterization is the review and documentation of relevant literature, interviews with
knowledgeable agency staff and other experts on the ecological and economic significance of the
fish and wildlife resources in the Nushagak and Kvichak watersheds. The goal of this review and
documentation process is to describe the level of current scientific knowledge offish and wildlife
resources of Bristol Bay, and to answer the question "What is the ecological and economic
significance of Bristol Bay fish and wildlife resources locally and around the North Pacific
Ocean?" This USFWS report is primarily intended to provide information to EPA about wildlife
resources in the Nushagak and Kvichak watersheds for their use in developing the BBWA but, it
12
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could also be useful for other land use planning or project environmental review and permitting
activities. EPA is preparing a separate report which characterizes fish in the Bristol Bay region.
Predictive risk assessment is the review and documentation of relevant literature, interviews with
knowledgeable agency staff and other experts on the risks, threats and stressors associated with
current and foreseeable human activity on the health, productivity, ecological integrity, and long-
term sustainability of fish and wildlife resources of Bristol Bay. It includes review and
documentation of mitigation practices used to abate threats and risks to fish and wildlife resources.
Based on the scope of the BBWA, USFWS formulated three objectives for this report to EPA.
These are:
1. Describe significant, representative wildlife species in Bristol Bay and their importance to
humans;
2. Describe the importance of marine derived nutrients to these wildlife species; and
3. Describe the role of these wildlife species in distributing marine derived nutrients throughout
the ecosystem.
Geographic Scope of USFWS Evaluation
The primary geographic scope of the BBWA, and therefore this report, is the Nushagak and
Kvichak watersheds. In addition to terrestrial mammal species, the USFWS is characterizing and
assessing migratory birds, many of which use marine waters during some portion of the year.
Therefore, we needed to determine the extent of freshwater influence from these river systems in
the Bristol Bay marine environment. Data on the hydrography of the Bay is limited; however, an
evaluation based on salinity differences, drift cards and fluorescent dye (Straty, 1977), indicates
that the net seaward flow of lighter and less saline river runoff water from the Kvichak River (and
the adjacent Naknek River to the south) moves in a counterclockwise direction along the
northwest side of inner Bristol Bay, where it mixes with water from the Nushagak River. Data
indicate that the mixture of Kvichak and Naknek river water remains relatively distinct as far as
Cape Constantine. On the southeast side of the bay, the answer is less clear. However, Egegik
River water does not appear to travel much further north than Middle Bluff Light. North of Middle
Bluff Light water immediately along the shoreline is probably mostly Naknek River water, but by
around 8 km off shore, Naknek River water is mixed with Kvichak River water. Therefore, to
facilitate evaluation of migratory birds in marine waters, we are defining the area of freshwater
influence as the area north of a line drawn from Cape Constantine east to Middle Bluff Light. We
call this area inner Bristol Bay for purposes of this report. This area is marked with the yellow
dashed line on Figure 1.
Selection of Wildlife Species for Characterization
EPA asked the USFWS for assistance in evaluating wildlife species that can be directly linked to
salmon, due either to their direct dependence on salmon for survival (food) or to their role in
distributing marine derived nutrients throughout freshwater aquatic or terrestrial ecosystems. The
vast number of wildlife species in the Nushagak and Kvichak watersheds made it impractical to
characterize each individual species. Additionally, staff resources available to the USFWS and
time constraints imposed by the EPA BBWA process, also meant that there would be no new
primary wildlife data collection and that our analysis would be based primarily on previously-
collected data.
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The first step, therefore, was to identify a subset of wildlife species that represent major
components of biodiversity in the region. Initially, the USFWS decided to use portions of an
approach developed by The Nature Conservancy for ecoregional planning (Groves et al., 2000).
This process was modified for use by the Mat-Su Salmon Habitat Partnership to develop a
Strategic Habitat Action Plan and also by The Nature Conservancy in preparing the Alaska
Peninsula and Bristol Bay Basin Ecoregional Assessment (The Nature Conservancy in Alaska,
2004). A simplified version was also used by the Southwest Alaska Salmon Habitat Partnership to
develop a draft Strategic Habitat Conservation Action Plan. The USFWS is a member of both fish
habitat partnerships and USFWS biologists were also involved with the Ecoregional Assessment,
so we are familiar with ecoregional planning concepts.
Ecoregional planning, as described by Groves et al. (2000) is a complex, step-by-step approach
with the goal of selecting and designing networks of areas of high biodiversity significance
(conservation sites) that will conserve the diversity of species, communities, and ecological
systems in an ecoregion. Since this is not the goal of the USFWS's work as part of the EPA
BBWA, we used selected portions of the approach to help us identify wildlife species targets.
Species-level targets facilitate identification of threats and development of strategies and actions to
abate threats.
The USFWS selected key wildlife species for our contribution to the EPA BBWA based largely
on professional judgment, and consultation with EPA, and members of the BBWA
Intergovernmental Technical Team. Key species regulate energy flow and nutrient dynamics or
they may be ecosystem engineers that modulate habitat structure (Davic, 2003). We define key
species as being those we know from experience, have a direct link to salmon or are of special
interest to local or Alaska residents. Additionally, migratory birds and bald eagles are considered
key species because the USFWS has direct statutory authority for them under the Migratory Bird
Treaty Act and the Bald and Golden Eagle Protection Act.
The term "key species" should not be confused with the term "keystone species" which are those
whose impact on a community or ecological system is disproportionately large for their abundance
and biomass (Paine, 1995). Keystone species have spatial, compositional and functional
requirements that may encompass those of other species in the region and they may play a critical
role in maintaining the structure of an ecological community. They affect many other organisms in
an ecosystem and help to determine the types and numbers of various other species in the
community. They contribute to ecosystem function in a unique and significant manner through
their activities. Their removal may initiate changes in ecosystem structure and often a loss of
diversity (e.g., beaver, bison, prairie dog). Keystone species may also be wide-ranging and depend
on vast areas for survival. These species include top-level predators (e.g., wolves, grizzly bear) as
well as migratory mammals (e.g., caribou), anadromous fish and birds (The Nature Conservancy
in Alaska, 2004). Keystone species regulate local species diversity in lower trophic levels (Davic,
2003).
The key species the USFWS selected for inclusion in the wildlife component of the BBWA are:
brown bear;
moose;
barren ground caribou;
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wolf;
waterfowl (as a guild);
bald eagle;
shorebirds (as a guild); and
landbirds (as a guild).
The USFWS also considered including species listed under the Endangered Species Act and
seabirds but a preliminary assessment revealed there are no significant occurrences of either listed
species or significant assemblages of seabirds in the Nushagak and Kvichak watersheds or inner
Bristol Bay.
The Characterization Process
Staff from the Anchorage Fish and Wildlife Field Office (AFWFO) prepared initial drafts of all
but two of the species accounts. The waterfowl species account was developed by a contractor and
the shorebird account was drafted by a USFWS National Wildlife Refuge biologist. Species
accounts were drafted based on a suggested outline provided by AFWFO. We circulated for
review and comment initial draft species accounts to numerous experts from the USFWS, other
federal and State agencies, and other knowledgeable individuals, many of whom are retired federal
or State wildlife biologists. The USFWS then obtained assistance from expert wildlife biologists
to revise the draft species accounts on brown bear, caribou, moose, and bald eagles. Finally, we
circulated the revised draft species accounts for review and comment. A list of authors and
reviewers for each species account is provided in Appendix 1. All completed species accounts are
incorporated in this report.
Species accounts were largely developed using available published data and information, but in
some instances we used unpublished information from species experts. Our preference was to use
data specific to the Nushagak and Kvichak watersheds. However, we also used relevant
information from other regions of Alaska or other parts North America.
CHARACTERIZATION OF WILDLIFE
Overview of Wildlife Species
Alaska- Many Alaskans depend on the State's diverse wildlife resources1 for food and enjoyment.
Traditional subsistence harvest and personal use, sport and guided hunting, as well as
nonconsumptive uses of wildlife, such as wildlife viewing, are critical components of the Alaska
economy and lifestyle (ADF&G, 2006). According to the Alaska Department of Fish and Game
(ADF&G), the value of game species such as moose, caribou, and deer are well understood by
most Alaskans(ADF&G, 2006). Historically, species not taken for subsistence, sport, or
commercial use, were perceived as having little direct economic value. However, the contribution
of nongame species to Alaska's economy is substantial, although difficult to quantify. According
to ADF&G, basic biological information on life history, population levels, and other parameters is
lacking for many species, but the majority of Alaska's wildlife resources are considered healthy.
1 ADF&G includes fish, reptiles, amphibians, and marine mammals (including whales) in the definition of wildlife in
the Comprehensive Wildlife Strategy. Of the 1,073 vertebrate species known to occur in Alaska, there are 469 species
of birds and 64 species of terrestrial mammals (ADF&G 2006; Jarrell et al 2004).
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Southwest Alaska Region- Southwest Alaska, including the Nushagak and Kvichak watersheds,
possesses intact, naturally functioning terrestrial ecosystems that still support their historic
complement of species, including large carnivores. Such ecosystems, containing historic levels of
biodiversity, are becoming extremely rare globally. Large terrestrial herbivore-predator
interactions are an intrinsic property of intact functioning ecosystems and are a flagship ecological
feature of southwest Alaska (Bennett et al., 2006; Jarrell et al., 2004; Nushagak-Mulchatna
Watershed Council, 2007).
Based on a compilation of vertebrate species lists for Togiak National Wildlife Refuge (NWR)
(USFWS, 2009b), Alaska Peninsula and Becharof NWR Bird List (USFWS, 201 Ob), and
southwest Alaska national parks (Cook and MacDonald, 2005), there are 192 species of birds and
41 species of terrestrial mammals in southwest Alaska (Appendix 2). There is no comprehensive
species list available for the Nushagak and Kvichak watersheds, but it is reasonable to assume that
species in these watersheds are nearly identical to those in the larger region.
Of all the species described in this report, caribou, moose and waterfowl are probably the most
important subsistence species to local residents. These species also provide significant recreational
hunting opportunities for both nonlocal resident and nonresident hunters. Other wildlife species
which provide subsistence food for local residents include black bears, beaver, ptarmigan, and
porcupine. River otter, red fox, marten and wolverine are also important in the region. We have
not identified any of these other wildlife species as key species for purposes of this report.
The Importance of Marine-Derived Nutrients to Bristol Bay Watershed
Ecosystems
One of the most important ecological functions of salmon is to transfer large quantities of nutrients
from the marine environment into terrestrial and freshwater ecosystems within the watersheds
where adults return to spawn. Pacific salmon spend most of their lives feeding and growing at sea
before returning to spawn and die in their natal streams or lakes, and approximately 95 to 99% of
the carbon, nitrogen, and phosphorus in the adult salmon body are derived from the marine
environment (Larkin and Slaney, 1997; Schindler et al., 2005). These nutrients are transported
inland when salmon return to their natal streams or lakes to spawn. For oligotrophic Lake Diamna,
the annual nitrogen pool associated with the annual spawning migration is comparable in size with
the dissolved nitrogen pool in the lake, indicating the importance of adult salmon to the lake's
nitrogen budget (Kline et al., 1993). Marine-derived nutrients (MDN), in combination with other
ecosystem features such as suitable spawning habitat and oceanic carrying capacity (Schindler et
al., 2005), are essential for the survival and growth of the next generation of salmon, and also
greatly benefit a number of other fish and wildlife species.
Brown bears within the Nushagak and Kvichak watersheds depend on the annual salmon runs as
an excellent source of lipids. In several Alaskan brown bear populations, lipids obtained through
the consumption of salmon contributed approximately 80% of the mass gained by bears prior to
winter hibernation (Hilderbrand et al., 1999c). Accumulation of fat reserves is important for
successful hibernation, and for female reproductive success (Farley and Robbins, 1995;
Hilderbrand et al., 1999c).
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Wolves also consume salmon seasonally when available, and this marine-derived protein source
can be a major component of the lifetime total diet of wolves in areas where the two species co-
exist (Adams et al., 2010; Darimont et al., 2003; Szepanski et al., 1999). Salmon are not solely a
food resource for coastal wolves, as some Pacific salmon spawning grounds are hundreds of miles
from the ocean. A study in Interior Alaska documented the seasonal salmon consumption among
wolves who lived more than 1,200 river miles from the coast (Adams et al., 2010).
Both brown bears and wolves play an important role in distributing MDN from streams to the
terrestrial environment, both by transporting salmon carcasses prior to consumption and through
excretion of wastes rich in salmon nutrients (Darimont et al., 2003; Helfield and Naiman, 2006).
As far as we can determine, there has been no research conducted on the MDN links between
salmon and moose or caribou. However, we believe that while it is reasonable to assume that
MDN transported to the Nushagak and Kvichak watersheds by salmon may be detectable in tissue
of animals, especially those preferentially feeding in riparian areas (e.g. moose), this does not
necessarily mean there would be any detectable benefit to these herbivores. In order for there to be
a direct effect, salmon would have to have a strong fertilizing effect on forage, resulting in
additional biomass on the landscape, and ungulates would have to be forage limited in order for
the increased biomass to matter (Adams, personal communication). There have been anecdotal
reports of ungulates (both deer and moose) feeding directly on salmon carcasses (Hilderbrand,
personal communication).
Waterfowl benefit from salmon as both direct sources of prey and carrion and indirect nutrient
drivers of aquatic systems. The large influx of nutrients to riverine and terrestrial systems strongly
benefits waterfowl (Holtgrieve, 2009; Willson et al., 1998; Willson and Halupka, 1995). Of the 24
duck species that regularly occur in Bristol Bay, at least eleven species are known to prey on
salmon eggs, parr, smolts, and scavenge on carcasses (Table 9). Of these, greater and lesser scaup,
harlequin duck, buffiehead, common and Barrow's goldeneyes, and common and red-breasted
mergansers exhibit directed foraging on salmon. Among dabbling ducks, mallards feed most on
salmon because they are distributed across a diversity of summer habitats in spawning areas, and
they are the principal wintering dabbling duck on the North Pacific coast where fall-winter salmon
runs occur. Fish predators like mergansers feed extensively on salmon fry and smolts (Munro and
Clemens, 1932; Munro and Clemens, 1937; Munro and Clemens, 1939; Salyer and Lagler, 1940;
White, 1957; Wood, 1987a). Other duck species may prey on smolt incidentally. Salmon eggs are
a seasonally rich food source for harlequin ducks, goldeneyes and scaup that frequent rivers and
streams (Cottam, 1939; Dzinbal and Jarvis, 1984; Munro, 1923) and probably for other
opportunistic ducks. Species ranging from dabbling ducks (mallard, green-winged teal) and diving
ducks to sea ducks that inhabit spawning waters probably opportunistically scavenge easy protein-
rich meals from salmon carcasses.
Spawned-out salmon carcasses provide an ideal food resource to bald eagles as they accumulate
on stream banks, river bars, lake and ocean shores, and tidal flats (Armstrong, 2010). Salmon
carcasses may provide food to eagles beyond the spawning season, as large numbers of carcasses
may be frozen into river ice during the winter and become available for consumption by eagles the
following spring (Hansen et al., 1984). The abundance of salmon affects bald eagle population
size, distribution, breeding and behavior (Armstrong, 2010). In one studied population, salmon
availability in spring was tightly correlated with whether adult bald eagles laid eggs in a given
year, and also influenced the timing of egg-laying (Hansen et al., 1984). As with other salmon
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carcass consumers, bald eagles affect the ecosystems within their range by distributing MDNs in
their excretions (Gende et al., 2002).
Direct or indirect interactions between shorebirds and salmon are not well-documented. Some
shorebird species are observed to consume dead salmon and salmon eggs, but it is unlikely that
shorebirds have an impact on salmon populations. No studies have been conducted to deduce the
contribution of salmon to the energetics of shorebird populations; however, the abundance of
invertebrates in the intertidal zone is very likely due in part to MDN from salmon that die on the
coast and in the rivers feeding Bristol Bay. Shorebirds play a role in distributing MDNs into the
terrestrial system, especially during the migratory period, but this has not been quantified. They
frequently feed in the intertidal zone, but roost in the terrestrial zone, where they deposit their
waste. In the spring, shorebirds need to acquire critical food resources, not only to fuel their
migration, but also, for some species, to assure that they arrive on the breeding grounds with
sufficient reserves to initiate nesting and egg production (Klaassen et al., 2006; Yohannes et al.,
2010). Some of the longest migrations known to birds involve shorebird species (bar-tailed
godwit) that use Bristol Bay intertidal areas in autumn (Battley et al., 2011; Gill et al., 2009). Such
flights are possible not only due to the extreme abundance of intertidal invertebrates (polychaetes,
crustaceans, gastropods, and bivalves) in the region, but also because the adjacent uplands are
usually rich in fruits of ericaceous plants or tubers on which birds like plovers, whimbrels, and
godwits, regularly feed (Elphick and Klima, 2002; Johnson and Connors, 1996; Paulson, 1995;
Skeel and Mallory, 1996). Some species like whimbrel, Hudsonian godwit, and black and ruddy
turnstones feed on herring roe, carrion, including salmon carcasses, and salmon eggs (Elphick and
Tibbitts, 1998; Gill et al., 2002; Handel and Gill, 2001; Nettleship, 2000; Norton et al., 1990).
Many shorebird species make use of freshwater invertebrates; small fish may be consumed by
yellowlegs (Elphick and Tibbitts, 1998) and phalaropes (Rubega et al., 2000).
Landbirds also benefit from salmon carcasses when available. Aquatic invertebrate larvae feed on
salmon carcasses (Wipfli et al., 1999), overwinter in the soil, and emerge in the spring as adults.
These invertebrate adults become aerial prey for a variety of landbird species in the spring, and
serve as an important seasonal subsidy during a period when terrestrial invertebrate biomass is low
(Nakano and Murakami, 2001). Salmon also benefit landbirds by increasing plant productivity due
to MDN inputs (Helfield and Naiman, 2001), potentially resulting in an abundance of berries and
seeds for avian consumption (Christie and Reimchen, 2008). Some birds, such as the American
Dipper, directly consume salmon eggs, fry, and small bits of carcasses when available. In one
study of dippers, consumption of salmon fry was related to higher fledgling mass and lower brood
reduction (Obermeyer et al., 2006).
In summary, MDN from salmon cycle throughout the ecosystem of watersheds with healthy
salmon runs, benefitting wildlife, increasing vegetation productivity, and promoting the
production of periphyton, aquatic macroinvertebrates, resident freshwater fish, and juvenile
salmon (Helfield and Naiman, 2006). This nutrient cycling is in turn dependent on interactions
with wildlife, which distribute MDN into the terrestrial environment through both transport of
carcasses and excretion of wastes. The loss of either salmon or key wildlife species may result in
significant changes to the productivity, diversity and physical structure of the ecosystem, via
mechanisms that extend beyond simple "food chain" interactions (Helfield and Naiman, 2006).
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Overview of Land Cover and Habitat Types
The National Land Cover Database (NLCD) 2001 is a Landsat-derived, 30-meter spatial
resolution map that illustrates land cover for the United States, including Alaska (Homer et al.,
2004) (http://alaska.usgs.gov/science/geography/nlcd.html). The NLCD dataset is a fairly broad
level characterization and is the only dataset that covers the entire Nushagak and Kvichak
watersheds. Therefore, we use the NLCD as a surrogate for a wildlife habitat-type map. Nineteen
land cover types are described for Alaska: open water; perennial ice/ snow; developed, open
space; developed, low intensity; developed, medium intensity; developed, high intensity; barren
land; deciduous forest; evergreen forest; mixed forest; dwarf scrub; shrub/scrub;
grassland/herbaceous; sedge/herbaceous; moss; pasture/hay; cultivated crops; woody wetlands;
and emergent herbaceous wetlands (Selkowitz and Stehman, 2011; Talbot, 2010).
Figure 2 shows the land cover types present in the Nushagak and Kvichak watersheds. The NLCD
does not describe wetlands in great detail and wetlands likely extend into other land cover types
not specifically identified as such (e.g., sedge/herbaceous land cover, which may include wet
tussock tundra, and evergreen forest, which may include wet black spruce bogs). Table 1 shows
the total size of each watershed and sub-watershed, as well as the percentage of each NLCD land
cover type in each.
Nushagak-Mulcnatna
Watershed
14,077 sq. miles
Port
sworth
Kvichak
Watershed
9,467 sq. miles
Stuyahp
!n
Ekwok
V
Iliumna
Igiugig
National Land Cover Dataset (NLCD)
USGS - using year 2000 LANDSAT imagery
Open Water
Perennial Ice/Snow
Developed, Open Space
Developed, Low Intensity
Developed, Medium
Barren Land
Deciduous Forest
Evergreen Forest
Mixed Forest
Dwarf Scrub
Grassland/Herbaceous
Sedge / Herbaceous
Moss
Woody Wetlands
Emergent Herb. Wetlands
Shrub / Scrub
Developed, High
Figure 2. National Land Cover Dataset, land cover types for the Nushagak River watershed and
Kvichak River watershed.
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Currently, there is only one statewide ecosystem map available for Alaska (Nowacki et al., 2001).
This map describes 32 ecoregional landscapes, but it is very coarse and not intended to provide
specific habitat classifications for wildlife. This ecosystem map does not provide sufficient detail
to be used for habitat type classification as part of this project.
1, mi,) the
or Cover
Size
(Square
Miles)
Land Cover
Type
Open-Water
Perennial
Ice & Snow
Developed,
Low
Intensity
Barren
Land
Deciduous
Forest
Evergreen
Forest
Mixed
Forrest
Dwarf
Scrub
Shrub Scrub
Sedge
Herbaceous
Moss
Woody
Wetlands
Emergent
Herbaceous
Kvichak River Watershed
Lake
Clark
Subbasin
3,532
Lake
Iliamna
Subbasin
5,935
Sub-total
9,467
Nushagak River Watershed
Mulchatna
River
Subbasin
4,291
Upper
Nushagak
River
Subbasin
5,026
Lower
Nushagak
River
Subbasin
3,386
Wood
River
Subbasin
1,367
Sub-total
14,077
Total
23,537
Percent of Watershed
5.96
6.51
0.02
23.55
2.92
15.51
4.45
10.85
28.27
0.01
0.01
0.64
0.81
23.89
0.38
0.01
5.88
2.79
5.34
3.20
22.57
31.79
1.67
0.02
0.19
2.28
17.20
2.67
0.01
12.47
2.84
9.13
3.67
18.19
30.66
1.05
0.01
0.36
1.73
2.31
0.39
0.00
4.61
1.94
10.04
2.30
24.50
51.08
0.48
0.00
1.28
1.53
5.34
1.61
0.00
3.31
3.63
11.86
5.08
12.77
54.97
0.37
0.02
0.16
1.05
5.36
0.00
0.00
0.09
2.15
9.63
2.99
20.06
44.86
3.86
0.09
0.74
10.17
14.28
1.32
0.00
.36
4.36
13.81
5.47
5.26
44.12
0.13
0.34
1.24
6.32
5.29
0.82
0.00
2.94
2.83
10.96
3.77
17.24
50.23
1.22
0.06
0.75
3.90
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BROWN BEARS
Introduction
The purpose of this section is to characterize the brown bears that spend most of their lives in the
Nushagak and Kvichak watersheds. Some of the general information comes from decades of
research on this species in its entire cosmopolitan ranges. If variable facts are derived from study
areas other than the Nushagak and Kvichak watersheds, it is so noted. However, owing to their
interconnected home ranges and close genetic relationships, brown bears' habitat selection and
behavior show a great degree of overlap wherever they occur (Schwartz et al., 2003).
In this section, we use the term "brown bears" to refer to all North American bears of the
classification Ursus arctos, although bears in interior parts of North America traditionally have
been referred to as grizzly bears and those on the coast, in salmon-rich areas, as brown bears.
Ursus arctos on the Alaska Peninsula are commonly called brown bears.
Habitat
Brown bears are a wide-ranging species that, throughout the course of a year, use many different
plant and animal communities. Habitat is defined here as the location or environment where an
organism is most likely to be found. Habitats provide the food, water, and cover that a species
needs to survive. Biologists describe individual bear's habitats as "home ranges." A home range is
the normal area that an animal uses to carry out the activities of securing food, mating, and caring
for young. Brown bears are generally solitary, food-maximizing individuals whose home ranges
vary with the availability of their seasonal foods. When food is abundant, as is the case during
salmon runs, home ranges of female bears may be smaller because of their ability to obtain
sufficient energy to meet their nutritional needs. Conversely, home ranges may increase in order to
take advantage of more widely dispersed food resources (McLoughlin et al., 1999).
On the Alaska Peninsula, brown bears emerge from their dens between early April and early June.
Male brown bears tend to emerge before females. Females with cubs of the year are often the last
to leave the den (Miller, 1990). Brown bears often spend June to mid-August in lowland and
coastal areas, though probably not all bears in the Nushagak and Kvichak watersheds include
coastal plains in their home ranges. A study conducted on Admiralty Island found that brown
bears did not necessarily use rich coastal habitats as part of their home ranges, though the reason
for these different patterns of habitat selection are not known (Schoen et al., 1986).
Brown bears typically spend July through mid-September near streams that support salmon runs
(Schoen et al., 1986). The Nushagak and Kvichak watersheds contain at least 8,286 linear miles
(13,335 linear kilometers) of anadromous fish habitat, not including lakes (Johnson and Blanche,
2011). As salmon begin to appear in the streams, bears move closer to them, sometimes
congregating where shallow streams make preying on fish more efficient (Aumiller and Matt,
1994). Studies of bear predation on salmon in a series of streams in the Wood River system
demonstrated that the probability a fish is killed by bear increases with decreasing stream size
(Quinnetal., 2001).
Bears move to higher elevations in the fall, presumably to feed on berries and other food items
(Collins et al., 2005). Some brown bears will continue to feed on fish until October, especially at
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shallow streams where dead and dying sockeye salmon are washing out of lakes after spawning
(Fortin et al., 2007).
Dens are typically dug at higher elevations within bears' home ranges (Van Daele et al., 1990), on
20- to 50-degree slopes at between 300 and 1,600 meters in elevation. Most brown bears enter
dens by early November, with a mean entrance date of October 14. In a study from the Nelchina
area in south-central Alaska, brown bears spent an average of 201 days in winter dens (Miller,
1990).
Food Habits
Alaska Peninsula brown bears move within their home ranges in order to exploit seasonally
available food sources (Glenn and Miller, 1980). During late summer and fall, brown bears gain
weight rapidly, primarily stored as fat; peak body mass generally occurs in fall, just prior to
hibernation (Hilderbrand et al., 2000). It is essential for bears to maximize weight gain prior to
hibernation, since they metabolize only fat and muscle during that time and must rely on those
stored energy reserves for reproduction and survival.
In the spring, after den emergence, bears commonly graze on early season herbaceous vegetation,
such as cow parsnip (Heracleum lanatum\ horsetails (Equisetaceae), lupine (Lupinus spp.), false
hellebore (Veratrum viride) and grasses (Gramineae) (ADF&G, 1985). They also search for and
scavenge winter-killed carrion, as well as moose and caribou calves (Glenn and Miller, 1980).
Bears with access to salt marshes commonly graze on sedges (Carex spp.), grasses (e.g., Elymus
spp.), sea-coast angelica (Angelica lucida), and forbs (e.g., Plantago spp. and Triglochin spp.).
Brown bears are also known to dig soft-shelled clams (Mya arenarid) and Pacific razor clams
(Siliqua patuld) on intertidal beaches (Smith and Partridge, 2004). Brown bears on the coast may
also scavenge for dead marine life (Glenn and Miller, 1980).
In July through October, brown bears move to streams to take advantage of the predictable runs of
salmon. The Nushagak and Kvichak Rivers provide natal homes to five species of Pacific salmon
(Oncorhynchus spp.) and are part of the world's largest of run of salmon in Bristol Bay (Salomone
et al., 2011). The abundance of salmon in the Nushagak and Kvichak watersheds makes them
prime habitat for brown bears because salmon are an excellent source of lipids.
Lipids obtained through the consumption of salmon account for approximately 80% of the mass
gained by bears (Hilderbrand et al., 1999b). More than other factors, the accumulation of fat
determines whether brown bears will hibernate successfully, or in the case of females, produce
cubs (Farley and Robbins, 1995). In addition, bigger, fatter adult females produce faster growing
cubs that survive better than do cubs produced by smaller, leaner females (Ramsay and Stirling,
1988). Larger, fatter males also receive an advantage from their size. Dominance in males is
necessary to win breeding opportunities and defend estrus females, and larger males tend to
compete better for these opportunities than smaller males (Robbins et al., 2007).
Research conducted in Bristol Bay and Southeastern Alaska to determine factors that influenced
consumption choice of salmon made by brown bears found that the availability, as well as the age
and spawning status of the salmon, played a significant role in consumption choice (Gende et al.,
2004). When salmon were not abundant, bears consumed more biomass of each fish, rather than
just consuming body parts high in energy content. When salmon were abundant, bears ate parts of
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the fish considered to be higher in energy such as roe and the brain. In addition, bears were found
to consume ripe fish before fish that had spawned-out. Some bears were even observed catching
and releasing fish that had spawned-out in order to consume only ripe fish with higher energy
content (Gende et al., 2004).
Brown bears are also known to feed extensively on wild fruits, including crowberry (Empetrum
nigrum), lowbush cranberry (Vaccinium vitis-idaea), and bog blueberry {Vaccinium uliginosum)
(Fortin et al., 2007; Rode et al., 2006a; Rode et al., 2006b). The simultaneous ingestion of salmon
and berries appears to benefit bears, in terms of growth rate, compared to the ingestion of one or
the other alone (Robbins et al., 2007).
Fall foraging is especially important for brown bears. In the fall, brown bears seek out available
meat sources including salmon, ungulates, rodents, and mice, as well as berries, in order to store as
much fat as possible. The more efficiently bears forage, the more vital lipids they can store,
thereby improving their ability to live long and reproduce often (Robbins et al., 2007).
Behavior
Brown bears have generalist life history strategies, extended periods of maternal care, and
omnivorous diets. A generalist species is able to thrive in a wide variety of environmental
conditions and can make use of a variety of different resources such as a varied diet. Brown bears
travel comparatively long distances to find the amount and variety of food they need to flourish,
despite six months of hibernation.
Movement patterns that define home ranges are influenced by important food resources, breeding,
reproductive status, individual dominance status, security, and human disturbance (Schwartz et al.,
2003). Differences in home range size between study areas are attributed to differences in habitat
quality and distribution (McLoughlin et al., 2000). The larger ranges of adult males overlap
several females (Schwartz et al., 2003). Female brown bears are generally faithful to their home
ranges. Sub-adult females tend to stay close to or within the home range of their mothers.
However, sub-adult males tend to disperse longer distances (Glenn and Miller, 1980). Brown
bears searching for alternative foods outside their usual home ranges, in particular dispersing
bears, often run into more conflicts with humans, and human-caused bear mortality increases
(Schwartz et al., 2003).
Different study areas illustrate the range in size of brown bear home ranges relative to habitat
quality. For example, in the Nelchina River basin of south-central Alaska, adult female home
ranges averaged 408 km2, while those of adult males averaged 769 km2 (Ballard et al., 1982). In
contrast, Collins et al. (2005) estimated 356 km2 for adult females in the southwest Kuskokwim
Mountains, west of the Nushagak watershed. On the relatively productive Alaska Peninsula, Glenn
and Miller (1980) found an average seasonal range of 293 km2 for adult females and 262 km2 for
adult males. The authors were unable to gather data from a comparable sample size for adult males
(n=4 males vs. n=30 females), and therefore do not support a range size comparison based on
these results. They mention other parameters that seem to contradict the small male seasonal range
estimated from their small sample size. For example, the cumulative 6-year movements of 13 adult
males were greater than those of 49 females. In addition, they found that seasonal range movement
of subadult males (744 km2) was three times that of subadult females (249 km2).
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During winter months brown bears hibernate in dens and rely on stored energy reserves for
survival. In hibernation, bears can go up to seven months without eating, drinking, defecating, or
urinating (Schwartz et al., 2003). Generally, brown bears seek out remote, isolated areas and sites
that will accumulate enough snow to insulate them from cold winter temperatures, often on steep
slopes. Bears may prefer steeper slopes for denning sites, due to reduced potential for disturbance
(Goldstein etal., 2010).
Female brown bears enter estrus beginning around late spring and, depending upon male
availability, can still breed into August. However because males are rarely limiting in a
population, most breeding occurs in May and June. After the eggs are fertilized development
proceeds to the blastocyst stage and then halts. Embryo implantation is delayed until hibernation
begins. It is possible that a litter could have multiple sires. Female brown bears that have
successfully bred and have implanted embryos have an obligate denning requirement as the
newborns are completely helpless at birth and remain so for several months. Most births occur in
January and February after 6 to 8 weeks of gestation. All maternal care from fetal development
through the first four months of lactation occur in the den and all nutrient reserves for the
developing young are drawn from maternal body stores accumulated the previous summer/fall.
Litters can range from one to four cubs; however, twins are the most common (Schwartz et al.,
2003).
Brown bears in the Nushagak and Kvichak watersheds interact with humans around residences,
recreational sites, and in the backcountry. The behavior of bears during these interactions depends
on many variables (Herrero et al., 2005). Brown bears that have received food or garbage
"rewards" while near humans or their habitations can become food-conditioned. Food-
conditioning is a form of operant conditioning in which bears learn to associate sources of food
with humans or their infrastructure (Matt, 2010). Food-conditioned bears are more likely to
encounter humans in an aggressive manner, perhaps because they assertively seek foods where
humans are found. Preventing access to anthropogenic foods keeps bears from being positively
rewarded for close association with people (Herrero, 2002).
Habituation describes behavior that is different from food-conditioning (Aumiller and Matt, 1994;
Herrero, 2002). A habituated bear has been repeatedly exposed to a neutral situation, such as a
person observing it from a close distance. Bears will conserve energy by muting their reaction.
Consequently, habituation often is assumed to have occurred when bears tolerate people at close
distances. Habituation is not an all-or-none response and may vary widely among individual bears.
Some bears habituate to certain human artifacts such as roads and other structures (Follman and
Hechtel, 1990).
Today most brown bear attacks are associated with defensive behavior, such as females protecting
cubs or incidents involving protection of a food cache such as an ungulate carcass (Herrero, 2002;
Herrero and Higgins, 1999; Herrero and Higgins, 2003). Whether or not a bear charges or attacks
a human in a defensive manner is dependent on many factors in the immediate environment, as
well as the prior experience and individual behavior of both the human and the bear (Herrero et
al., 2005). Historical records strongly suggest that brown bears have not been important predators
on people, although rare incidences of predation may have occurred, as they still do today
(Herrero, 2002).
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Interspecies Interactions
Pacific salmon (Oncorhynchus spp.) and brown bears meet the basic criteria of "keystone
species." The loss of either species results in significant changes in the productivity, diversity and
physical structure of their communities, far beyond just their "food chain" interactions. By both
consuming and transporting partially consumed salmon, brown bears distribute MDNs via
decaying salmon carcasses and through excretion of wastes rich in salmon nutrients (Helfield and
Naiman, 2006).
Pacific salmon spend most of their lives feeding and growing at sea before returning to spawn and
die in their natal streams. Approximately 99% of the carbon, nitrogen, and phosphorus in their
bodies when they return to freshwater is from the marine environment (Hilderbrand et al., 1999c).
These nutrients are transported inland when salmon return to their natal streams to spawn. For
example, sockeye salmon (Oncorhynchus nerka) are estimated to import approximately 12,700 kg
of phosphorus and 101,000 kg of nitrogen back to the Wood River system each year (Holtgrieve,
2009).
Nitrogen, phosphorus, and carbon are among the most important MDNs. These nutrients cycle
through the ecosystem, benefitting other forms of wildlife and vegetation, promoting the
production of periphyton (e.g., algae), aquatic macroinvertebrates, resident freshwater fish, and
juvenile salmon (Helfield and Naiman, 2006). Insects that directly benefit from decaying salmon
include stoneflies (Plecoptera spp.), caddis flies (Trichoptera spp.), mayflies (Ephemeroptera
spp.), midges (Diptera spp.: Chironmids (Chironomidae spp.), blackflies (Diptera: Simuliidae\
and carrion beetles (Coleoptera: Silphidae) (Helfield, 2001).
As salmon enter freshwater streams in late July, and throughout August, brown bears become
hyperphagic (consume abnormally large quantities of food) as they store energy for winter by
consuming salmon which contain protein and fat (Hilderbrand et al., 1999b). Female brown bears
are each estimated to consume 1,003 kg (SD Vi 489 kg) of salmon each year and transport
approximately 37.3 kg of MDNs to terrestrial ecosystems on the Kenai Peninsula (Hilderbrand et
al., 2004; Hilderbrand et al., 1999a).
In one study, brown bears delivered 83 to 84% of marine-derived nitrogen found in white spruce
trees near two Kenai Peninsula creeks (Hilderbrand et al., 1999a). In addition, Helfield and
Naiman (2006) found Sitka spruce growth rates to be three times greater near salmon spawning
sites than in areas lacking spawning sites. Other studies also show that "bears feeding on salmon
increased soil ammonium concentrations three-fold and nitrous oxide (NOs) flux by 32 fold"
(Holtgrieve, 2009).
The level of MDNs transported by brown bear shows the significance of the abundance of salmon
in areas with brown bears. It also shows that loss of either population would change the nitrogen
budget "and, by extension, the productivity and structure of the riparian forest" (Helfield and
Naiman, 2006). The potential loss of this nitrogen source would also greatly affect other
organisms throughout the food chain and ecosystem (Helfield, 2001).
Mortality, Productivity, and Survivorship
Brown bears have one of the lowest reproductive rates among mammals. This low rate is mainly
due to their relatively late age of first reproduction, small average litter size, and long interval
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between litters. Their low reproductive rate makes brown bears particularly susceptible to impacts
from humans (Schwartz et al., 2003). On the Alaska Peninsula, the mean age is 4.4 years at first
reproduction for female brown bears (Miller and Sellers, 1992). The earliest that a brown bear
female might breed is in the late spring at 4.4 years. If the female is of adequate size and
nutritionally healthy, the fertilized ova develop into a blastocysts that stop developing until they
implant in the uterus in late November. Following 6 to 8 weeks of gestation, tiny cubs are born
within a den. The female would then emerge from her den with her surviving cubs in the late
spring at 5.4 years. If cubs are raised until the typical age of weaning (2.4 years), the female's age
of second breeding likely would not occur until she was 7.4 years (Schwartz et al., 2003).
Therefore, during the first ten years of her life, a female grizzly bear is capable of producing only
two litters.
Age at first reproduction, litter size and intervals between litters vary among populations and
individuals. Litter sizes vary from one to four cubs, with twins being the most common. Females
are capable of breeding in the same spring that they wean their cubs; however, they do not always
wean them at 2.4 years and may keep them until the cubs are 3.4 years. Evidence shows that the
average interval between litters is generally between three and six years (Schwartz et al., 2003). In
a ten-year study in southwest Alaska, there was great variation in reproductive intervals (Kovach
et al., 2006), although when compared to other studies in Alaska, the southwest population had
one of the highest production rates of offspring, yet the lowest number of female offspring weaned
per year.
Adult female brown bears that eat meat (mostly salmon) in the fall gain approximately 80% fat
mass (Hilderbrand et al., 1999b). Deposited fat is more important than lean body mass in
producing cubs during winter dormancy (Farley and Robbins, 1995). While late summer and fall
salmon are a critical resource on their own, it is the likely the availability of both fall berries and
salmon, together, enable brown bears to accumulate body reserves important for reproduction and
hibernation (Hilderbrand et al., 1998; Robbins et al., 2007). While it seems logical that
productivity (independent of mortality) would be higher in populations with access to salmon in
the fall, Kovach et al. (2006) found that, in southwest Alaska, females had only slightly higher
productivity than the figures reported for Yellowstone National Park and the Selkirk Mountains,
where salmon are not present (Kovach et al., 2006).
The age and sex structure of brown bear populations is dynamic. Many variables, such as habitat
conditions, time of year of observations, and hunting make generalizations difficult. However, in
one study of a hunted population on Kodiak Island, the population structure of brown bears was
26% cubs, 22% yearlings, 27% sub-adults, and 25% adults (Troyer and Hensel, 1964). The
number of cubs born varies from year to year, and the proportion of cubs in any brown bear
population is a function of both reproductive rate and early mortality. Cub survival rates, based on
observation of den emergence from the first year to the second year, were estimated for two areas
on the Alaska Peninsula. At Black Lake, cub survival was an estimated 57%, while cub survival in
Katmai National Park was an estimated 34% (Miller et al., 2003).
Survival of adult males varies among populations, but is generally lower in hunted populations. In
1985, in the middle Susitna River basin study area in south-central Alaska, there was an estimated
82% annual survival for adult males (Miller et al., 2003). Ten years later, in 1995, Miller et al.
estimated only 71% annual survival for adult males, due to increased hunting pressure. Female
survival rates are generally higher than males. In the middle Susitna study, Miller et al. (2003)
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estimated 90% annual survival for adult females in both 1985 and 1995. In their study of female
survivorship in a hunted population in the southwest Kuskokwim Mountains, midway between
Dillingham and Bethel, Alaska, Kovach et al. (2006) found mean annual survival estimates of 90.1
to 97.2% for radio collared females aged five years or older.
Due to the difficulty of observing entire life histories of brown bears, the causes of natural
mortality are not well known. While it is known that adult males kill juveniles and that adults kill
other adults, there are insufficient data to fully assess the effects of predation on younger bears by
adult bears (Schwartz et al., 2003). Brown bears are exposed to more dangers during some life
stages than others. Survival rate estimates for cubs during their first year of life shows particular
vulnerability. In the middle Susitna study, survival for cubs of the year was estimated at 67%
(Miller et al., 2003). Kovach et al. (2006) reported survival rates of 48.2 to 61.7% for cubs of the
year.
In the latter study, researchers estimated survival rates of 73.3 to 83.8% for one- and two-year-old
offspring, combined. Dispersing sub-adults may be forced to choose marginal home ranges or
areas near human habitation that are dangerous to their survival (Servheen, 1996). Brown bears
can be afflicted with parasites and diseases that may contribute to their demise from other causes
such as starvation. However, there are no known instances of parasites or diseases causing major
die-offs within populations (Schwartz et al., 2003).
In many brown bear populations, human-caused mortality is higher than natural mortality. The
rate of human-caused mortality varies greatly in Alaska, where contact between bears and humans
is a function of human population density, activities of both species, and hunting regulations.
Servheen (1996) lists the following categories of human-caused mortality, in order of frequency:
direct human/bear confrontations (hikers, backpackers, photographers, hunters, etc.); attraction of
grizzly bears to improperly stored food and garbage associated with human habitations and other
sources: careless livestock husbandry, including the failure to properly dispose of dead livestock;
inadequate protection of livestock; loss of bear habitat; and hunting, both lawful and illegal.
Population
Brown bear population abundance is usually measured in terms of density, since it is widely
considered to be the most biologically meaningful measure of abundance. Two methods for
estimating bear population density, capture-mark-recapture, and aerial line-transect survey, have
been used recently in Alaska. Both estimates survey in the early spring, prior to leaf growth on
deciduous shrubs and trees.
Alaskan brown bear populations vary significantly in density depending on the availability and
distribution of food (Miller et al., 1997; Schwartz et al., 2003). Brown bear populations achieve
maximum densities in areas where the populations have access to multiple runs of Pacific salmon
(Hilderbrand et al., 1999c; Miller et al., 1997). Based on a modified capture-mark-recapture
method for estimation, the Pacific Coast of the Alaska Peninsula has the highest documented
brown bear density in North America, at 551 bears (representing all ages) per 1,000 km2 (Miller et
al., 1997). The North Slope of Alaska has the lowest estimated density at 3.9 bears of all
ages/1,000 km2 (Reynolds, 1976).
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While population density estimates for the entire Nushagak and Kvichak watersheds are not
currently available, recent aerial line-transect studies in portions of the watersheds and in nearby
watersheds can give some idea about brown bear densities. Using the double observer aerial line-
transect method, brown bear density in the Lake Clark National Park and Preserve portion of the
Kvichak watershed was estimated at 39 bears/1000 km (1999 E. Becker personal
communication). Recent aerial line transects in the remainder of the inland Kvichak watershed
reveal an estimated 47.7 bears of all ages/1000 km2 (Becker, 2010). In the nearby northern Bristol
Bay area (Togiak NWR and Bureau of Land Management Goodnews Block), brown bear
population density was 40.4 bears/1,000 km (Walsh et al., 2010). The later study included both
coastal and inland habitats.
As expected, surveys that include coastal habitat have higher population density estimates during
spring. Researchers using the same line-transect survey method as in the above studies, estimated
brown bear density at 124 bears/1,000 km2 in Katmai National Park, an area that included both
coastal and inland brown bear habitat. Along the coast of Lake Clark National Park and Preserve
(NPP), brown bear density was estimated at 150 bears/1,000 km2.
When inferring the distribution of brown bears from density estimates, it should be noted that
brown bears move long distances across the landscape. Bears that are counted in coastal areas in
the spring may move inland and upstream in the summer and fall to take advantage of pre- and
post-spawning salmon. It is common to see brown bears in interior Lake Clark NPP feeding on
post-spawning salmon in September and October, and less commonly, in December (Mangipane,
personal communication). Fortin et al. (2007) recorded brown bears feeding on salmon into
October on the Kenai Peninsula.
Human Use/Interaction/Management
Major human "uses" of brown bears in the Nushagak and Kvichak watersheds include sport
hunting, subsistence hunting, and bear viewing. Besides these uses, there are many other
interactions between brown bears and humans, including both residents and non-residents visiting
the area for purposes other than seeking encounters with brown bears.
Sport Hunting for Brown Bears
Game Management Unit 9- The Alaska Board of Game has placed a high priority on maintaining
a quality hunting experience for the large brown bears of the Alaska Peninsula in Game
Management Unit (GMU) 9, which includes the Kvichak watershed (Figure 3). Due to relatively
easy aircraft access and the high quality of bear trophies in the unit, an active guiding industry for
the area developed during the 1960s. Non-resident sport hunters are required to use a guide for
brown bear hunting throughout Alaska, and to have their harvest inspected and sealed by ADF&G.
The ADF&G management program strives to maintain stable guide and client numbers over time.
As of 2007, approximately 75% of the GMU 9 brown bear harvest came from guided hunts
(ADF&G, 2009).
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Game Management Unit - ADF&G
Watersheds
Kvichak
Nushagak-Mulchatna
V#*&*
Newhalen* iiiamna
^.l»Xi>A%/y i
Figure 3. Nushagak and Kvichak River watersheds and associated Alaska Department of Fish and
Game, game management units.
The current brown bear management objective for GMU 9 is to maintain a high bear density with
a sex and age structure that will sustain a harvest composed of 60% males, with 50 males aged
eight years or older taken during the combined fall and spring seasons. In GMU 9, in the 2007
Regulatory Year (July 1, 2006 through June 30, 2007), results reported to ADF&G revealed a
harvest of 621 bears (72% male and 28% female).
Game Management Unit 17- In GMU 17, which includes the most of the Nushagak River
watershed, brown bears are neither as abundant nor, usually, as large as those found on the Alaska
Peninsula. GMU 17 has not received much hunting pressure in the past. However, interest in
hunting brown bears in many parts of Alaska is increasing, and bear hunting in GMU 17 has
increased substantially since the mid-1990s. As of 2007, the brown bear population objective for
GMU 17 was to maintain a brown bear population that will sustain an annual harvest of 50 bears
composed of at least 50% males (ADF&G, 2009).
Subsistence Hunting for Brown Bears-
On non-federal lands in GMUs 9b and 17, subsistence brown bear hunters must obtain a
subsistence registration permit for bears to be taken for food. In addition to requiring a registration
permit, the subsistence brown bear hunting regulations establish a hunting season of September 1
to May 31, limit take to one bear/regulatory year, and prohibit take of cubs and sows with cubs
(ADF&G, 2011 a). Salvage of the hide and skull is optional and the hide and skull need not be
sealed, unless they are removed from the Western Alaska Brown Bear Management Area, in
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which case they must be sealed by an ADF&G representative and their trophy value destroyed. All
edible meat must be salvaged for human consumption. On federal lands within GMUs 9 and 17,
residents must consult the Federal Subsistence Management Program Regulations, available at
http://alaska.fws.gov/asm/law.cfml?law=2&wildyr=2010. There are some differences between
federal and State subsistence hunting regulations. For example, on federal lands in GMU 9B, there
is a federal registration permit harvest quota of four females or ten bears and the season closes as
soon as the first quota is reached.
Bear Viewins-
Within the Kvichak watershed there are specific destinations for recreational visitors to view
brown bears. Lodges on Lake Clark, Kukakluk Lake, Nonvianuk Lake, and Battle Lake offer
brown bear viewing in addition to fishing expeditions. Several guides and air taxis take brown
bear viewers to Funnel Creek and Moraine Creek on day trips.
The 2006 National Survey of Fishing, Hunting, and Wildlife-Associated Recreation reported that
491,000 Alaskan residents and non-residents participated in wildlife watching as a primary
activity (USFWS and US Department of Commerce Census Bureau, 2006). Bear viewing is now
the leading recreational activity in Lake Clark NPP. The incidental "use" of bears has
spatiotemporal impacts to the bear use (Rode et al., 2007; Tollefson et al., 2005).
Other Human-Bear Interactions
Local Residents and Bear-Human Conflicts- In both GMU 9 and 17, the number of bears killed
annually includes both legally harvested bears and reported non-hunting mortalities. Villages with
open landfills attract bears during the spring, summer, and fall. Residential garbage, dog food, and
fish drying racks also bring bears close to humans. Incidences of local residents killing bears in
defense of life and property near villages and fishing sites are not uncommon. Although reporting
rates seem to have improved in recent years, most non-hunting mortalities of bears are reported
either indirectly or not at all; as such, any conclusions based solely on harvest data or reported
non-hunting mortalities should be viewed with caution (ADF&G, 2009).
During the 2007 regulatory year, in GMU 9, ADF&G received 17 reports of bears killed by people
in defense of life and property; however, wildlife managers estimated that the number of
unreported brown bear killings in the unit might be over 50 when considering unreported data.
During the same period in GMU 17, ADF&G received 5 reports of defense of life and property
mortalities; however wildlife managers assumed there were more unreported brown bear killings
(ADF&G, 2009).
Other Recreational Users and Bear-Human Conflicts- In Lake Clark NPP, park managers
analyzed 171 bear-human incidents over 24 years. They found that, in 46% of the incidents, brown
bears received food as a result of the encounter. Bears were killed in 23% of the incidents (Wilder
et al., 2007). Managers were concerned about the large number of food-conditioning incidents,
given that food-conditioned bears are responsible for the majority of human injuries from bears in
national parks (Herrero, 2002; Wilder et al., 2007). Food-conditioned bears have been found to be
3 to 4 times more likely to be killed by humans than non-food conditioned bears (Mattson et al.,
1992). Wilder et. al. (2007) also noted that casual bear photographers at private recreational cabins
at Telequana Lake, may have contributed to the high number of bear-human incidents, saying
"that individuals repeatedly fed bears in this area to facilitate photography."
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MOOSE
Introduction
Moose, the largest member of the deer family (Cervidae\ have a circumpolar distribution in the
northern hemisphere (ADF&G, 201 Id; Telfer and Kelsall, 1984), occupying a broad band of
boreal forest dominated by spruce, fir, or pine trees. Fire is a major force that shapes these
vegetative communities (Odum, 1983; Telfer and Kelsall, 1984). Moose occur in northern forests
of North America, Europe, Russia, and China (Karns, 2007). The association between moose and
the northern boreal forest is unique, as there are no counterparts in the southern hemisphere
(Shelford, 1963). There are four recognized subspecies of moose in North America (Hundertmark
et al., 2003; Peterson, 1952; Peterson, 1955); the one found in the study area (Nushagak and
Kvichak watersheds) is the Alaska-Yukon subspecies (Alces alces gigas) (Miller, 1899). This
subspecies is often referred to as the "tundra moose," since it is often found in and near tundra
areas (Bubenik, 2007). However, it also inhabits the boreal forest, low elevation riparian and delta
habitats, and mixed deciduous forest areas. The Alaska-Yukon subspecies is generally larger than
the other subspecies in body size and antler development. In Alaska, moose are found in suitable
habitats throughout the mainland portion of the State, except on the northernmost coastal plain
(LeResche et al., 1974), ranging from the Stikine River in southeast Alaska to the Colville River
on the Arctic Slope (ADF&G, 201 Id). Moose are ruminants, with a four-chambered stomach, and
are classified as browsers based on their foraging strategy (Hoffman, 1973). These herbivores
consume mainly tree and shrub leaves and woody twigs during winter (Renecker and Schwartz,
2007). Due to their large body size and the volume of vegetation consumed, moose play an
important role in plant productivity and nutrient cycling in ecosystems where they occur (Molvar
etal., 1993).
Here we characterize moose that inhabit the Nushagak and Kvichak watersheds, but we provide
substantial information accumulated from decades of research on this species from across North
America. Since moose are a generalist species, much of this information is relevant to moose in
the Nushagak and Kvichak areas. We borrow heavily from the work of a distinguished group of
moose biologists, which provides a near complete summary of moose biology and management
(Franzmann and Schwartz, 1998; Franzmann and Schwartz, 2007) and have attempted to cite them
when appropriate. We also relied on a separate, condensed version of these data with more recent
updates (Bowyer et al., 2003).
Habitat
Both stable and transitory habitats are important in the evolution of moose (Geist, 1971).
Permanent habitats are those that persist through time without alteration in character or condition,
such as riparian willow/poplar communities and high-elevation shrub/scrub communities that do
not succeed to different kinds of vegetation (Peek, 2007). Telfer (1984) characterized the full
range of moose habitats to consist of boreal forest, mixed forest, large delta floodplains, tundra
subalpine shrubs, and stream valley/riparian zones. According to Peek (2007), boreal forest
habitats are considered fire-controlled and likely represent the primary environments in which
moose evolved (Kelsall and Telfer, 1974; Peterson, 1955). As noted by Peek (2007), delta
floodplains are expected to have the highest density of moose, followed by shrub/shrub habitats,
boreal forests, mixed forests, and stream valley/riparian zones. A study in the Copper River Delta
supported the finding that large deltas and floodplains are the most productive of these five major
habitat types for moose (MacCracken et al., 1997b). Boreal forest habitats are the least stable
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through time, whereas stream valley/riparian zones are the most stable. In Alaska and the
Northwest Territories, the climax tundra and subalpine shrub communities at higher latitudes and
elevations are more stable in time and space than the alluvial and riverine habitats (Viereck and
Dyrness, 1980).
Transitory habitats of moose include boreal forests where fire creates successional shrub
communities that provide extensive forage. Geist (1971) hypothesized that islands of permanent
habitat found along water courses and deltas, and in the high elevation dwarf shrub communities,
serve as refugia where moose populations persist and from which they disperse into transient
habitats created by fire. The common occurrence of fire in boreal forests is considered sufficient to
promote adaptations favoring dispersal of yearling moose to newly created habitats (Peek, 2007).
The dominant land cover types in the study area consist of high elevation dwarf shrub, shrub/scrub
and tundra habitats, with lower elevation boreal forests (deciduous, evergreen and mixed) and
riparian habitats (woody wetlands) along water courses (Table 1). All of these cover classes
represent high-quality moose habitat.
Habitat selection by moose is influenced by availability of food, predator avoidance, and snow
depth (Dussault et al., 2005). Peek (2007) advanced the view that moose select habitat primarily
for the most abundant and highest quality forage. Since these resources are unequally distributed
in space and time, moose habitat may be considered as a series of patches of different kinds and
sizes, with the value of each patch varying through the year. However, the total year-round value
of a diverse habitat should be emphasized even if each part is only critical at one season or
another. Peak (2007) further stated that the caveat to this general proposition was that sufficient
size of both overall habitat, and possibly each patch of any given habitat, must be accessible to
make an area most suitable for occupation by moose. As a corollary, if a certain critically
important community, such as shrub/scrub vegetation type, is unavailable in sufficient quantity,
then the ability of an overall habitat to support moose may be reduced even if it contains a highly
diverse set of other plant communities.
The typical annual pattern of moose habitat selection includes open upland and aquatic areas that
provide the best forage in early summer, followed by more closed canopy areas that provide the
best forage as summer progresses and plant quality changes. In autumn, after the rut and into
winter, moose intensify use of open areas with the highest biomass of dormant shrubs, where the
remaining major source of palatable forage occurs. Closed canopy areas are used in late winter
when forage is naturally at its lowest value and quantity for the year. The nature of the cover used
at this time will provide the best protection available from wind and cold, and may range from tall
shrub communities to tall closed canopy conifer stands (Peek, 2007). Metabolic activity in moose
generally corresponds to this pattern, being highest in summer and lowest in winter (Regelin et al.,
1985). Alaskan moose generally do not use areas > 1,220 m in elevation (Ballard et al., 1991).
Also, in Alaska, because forage quantity and quality (nutritional value) in summer and winter can
differ by orders of magnitude, winter habitat availability is often the ultimate limit on moose
abundance (Stephenson et al., 2006). Spatial heterogeneity of habitat on a relatively small scale (<
34 km2) enhances habitat quality for moose (Maier et al., 2005), probably because it enables
moose to respond to rapidly changing conditions such as climate (Stephenson et al., 2006).
Moose benefit from early successional stages of vegetation, which provide the woody browse
biomass that moose require (Schwartz, 1992). A disturbance regime that provides persistent shrub
communities, distributed in a diverse mosaic on the landscape, is essential to high moose density
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(Stephenson et al., 2006). In Alaska, this disturbance can be provided by fire (LeResche et al.,
1974; Maier et al., 2005), glacial outwash, and earthquakes (Stephenson et al., 2006). On the
Kenai Peninsula, forest succession following fire provided the most abundant forage for moose 20
years post-burn (Bangs et al., 1985; Schwartz and Franzmann, 1989; Spencer and Chatelaine,
1953; Weixelman et al., 1998). Schwartz and Franzmann (1989) reported that, after the 1969 fire,
moose abundance peaked about 15 years after the burn, when browse plants reached maximum
productivity. The increase was attributed to high production and low mortality, with some initial
shifting of home ranges from adjacent high-density populations. Where fire was absent for 25
years, moose densities on the Kenai Peninsula were sufficiently high to cause the forage base to
shift from a multispecies complex to a much less diverse community dominated primarily by
white birch (Oldemeyer et al., 1977). In the boreal forest, the optimum success!onal stages for
moose are 11-30 years after burning (K el sail et al., 1977).
In Game Management Unit (GMU) 17 in northern Bristol Bay, moose habitat is enhanced
primarily by the scouring of gravel bars and low-lying riparian areas by ice and water during the
spring thaw (Woolington, 2008). Willows and other plants quickly regenerate after bank scouring
and subsequent deposit of river silt (Woolington, 2004). This disturbance mechanism is
particularly important for the Nushagak and Mulchatna Rivers and for the lower reaches of the
major tributaries to those rivers (Woolington, 2008). Major river systems with large riparian
zones, like the Nushagak and Mulchatna Rivers, represent alluvial habitats that support an
abundant moose population, because they feature an abundance of nutritious food, primarily in the
form of regenerating willow stands. Deciduous shrubs proliferate in these areas because of the
annual influx of nutrients from waterways, sufficient soil moisture, and changing river channels.
Lightning-caused wildfires also occur occasionally in GMU 17 (Woolington, 2008), and provide
disturbance that enhances moose habitat. Moose habitat has not been formally assessed for GMUs
17B and 17C. Much of GMU 17 is wet or alpine tundra, and moose are located mostly along the
riparian areas (Woolington, 2008).
In interior Alaska, habitat characteristics and female moose densities were evaluated using a
spatial linear model (Maier et al., 2005). The densest moose populations occurred closer to towns,
at moderate elevations, in areas with greatest amounts of riparian habitat, and in areas where fire
occurred between 11 and 30 years prior. Moose avoided non-vegetated areas. Female moose
preferred areas with patch richness, indicating their need for a diverse habitat with both food and
availability of concealment. It was postulated (Maier et al., 2005) that moose might have preferred
to be near towns either because human disturbance of vegetation provides high-quality foods for
moose, or because predators such as wolves and grizzly bears tend to avoid human-inhabited
areas.
On the Copper River Delta, open tall alder-willow and low sweetgale-willow habitats were used
most by moose, and use of closed tall alder-willow habitat was intermediate (MacCracken et al.,
1997b). Aquatic and woodland spruce habitats were used the least by Copper River Delta moose.
Aquatic plants were used seasonally by Copper River Delta moose during the period from April
through August, and were used primarily for foraging by both sexes (MacCracken et al., 1993).
In northwest Alaska during March and April, moose occurred at stands of felt-leaf willow (Salix
alaxensis) 85.0 % of the time, followed by other willow (Salix) (6.4 %), riparian areas (3.9 %),
gravel bars (2.5 %), and upland areas (1.3 % of the time) (Gillingham and Klein, 1992).
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Winter Habitat- The influence of snow on moose habitat use patterns has received considerable
attention. In severe winters, snow depth can be a limiting factor for moose populations. Deep
snows can reduce browse availability by burying it, and travel through deep snow increases energy
expenditure (Ballard et al., 1991). Snow characteristics of ecological significance include
temperature, density, hardness and depth (Peek, 1986). Since the temperature of snow fluctuates
less than ambient temperatures and never gets as low as the air temperature, snow provides
insulation for moose against temperature extremes (Peek, 2007). Density and hardness influence
the ability of an animal to travel across or through the snowpack. Under some conditions, snow
density can be sufficient to support the mass of a wolf, but not a moose. Under these
circumstances wolf predation on moose tends to be high (Ballard and Van Ballenberghe, 2007). It
has been shown for other cervids (mule deer and elk) that energy expenditure while moving
through snow increases exponentially with increasing snowpack maturation through the winter.
Hardness and density affect sinking depth, and snow level at front knee height has been suggested
as a threshold parameter (Parker et al., 1984). Applying the same principals and relationship to
moose, a snow depth beyond 50-60 cm would result in a relatively large increase in energy
expenditure for movement (MacCracken et al., 1997a). Snow depths ranging from 70-100 cm
have been shown to limit the travel of moose (Des Meules, 1964; Kelsall, 1969; Kelsall and
Prescott, 1971). When snow depths approach 97 cm, moose have been confined to areas where
forest canopies are dense (Kelsall and Prescott, 1971).
The distribution of snow within the forest influences moose habitat use patterns. Snow depth is
nearly always greater in open areas until late winter, when snow exposed directly to the sun melts
more rapidly than snow protected by tree canopies. Snow falling on tree branches of fine-needled
conifers, such as spruce, tends to be retained in the canopy and produces a lower snow depth
immediately beneath the tree canopy in areas called a tree well (Pruitt, 1959). When this snow
sheds from the tree branches to the ground, it tends to produce a hard dense surface, which
provides more support for moose traveling beneath the canopy (Peek, 2007). In some geographic
locations with deep winter snow, mature coniferous forests can provide zones of shallow snow
accumulation that benefit moose survival (Balsom et al., 1996). In deep snow habitats where
conifers are absent, such as in shrub/scrub tundra or riparian communities, moose still use the best
microsites offered by combinations of shrub canopies and topographical situations that reduce
snow depths. However, the principal adaptation simply is to reduce energy expenditure (Peek,
2007).
Severe winters have been associated with high moose calf mortality from starvation (Ballard et al.,
1991). In Quebec, females with calves had a greater preference for habitats providing protection
from predators, while solitary adult females preferred habitats with moderate food abundance,
moderate protection from predators, and substantial shelter against deep snow (Dussault et al.,
2005). In Denali National Park and Preserve (DNPP) during the severe winter of 1986, large males
were the only moose able to remain in the Jenny Creek unit, which had a higher forage biomass
but deeper snow than other units (Miquelle et al., 1992). Moose are very tolerant of cold
temperatures, but are susceptible to heat stress. The upper critical temperature range for moose
during winter is -5 to 0°C; during the summer upper thermal limits are 14 to 20°C (Renecker and
Hudson, 1986).
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Food Habits
The moose diet is comprised mainly of leaves, twigs and bark of woody plants (Schwartz, 1992;
Van Ballenberghe et al., 1989). Renecker and Schwartz (2007) reviewed the diet items consumed
by moose. They list more than 221 plant genera/species, with willow (Salix), birch (Betula), and
alder (Alnus) predominating across North America. Daily patterns of moose use of time and space
explain how the animal satisfies hunger, remains fit, avoids thermal stress, maintains security from
predators, and reproduces. Since many of an individual moose's life cycle needs interact daily,
tradeoffs often occur, because most requirements are more critical at certain times than at others.
The day-to-day needs of moose for food and thermoregulation are most often preempted in favor
of other activities that accommodate fitness and mating. However, the survival instinct is satisfied
most on a daily basis by optimizing food consumption at minimal risk and effort. In this regard, a
basic constraint for most moose is an abundant food supply of low quality (Renecker and
Schwartz, 2007).
Digestive strategies of wildlife species vary significantly. Moose are ruminants, with a four-
chambered stomach. In ruminants, browse, forbs, and grass are held in the large-chambered rumen
of the stomach until adequate nutrients are extracted from the fibrous materials and the plant
particles are small enough to pass through to the omasum and true stomach. On the basis of
feeding habits, specialization and design of the digestive anatomy, ruminants are classified into
three main groups (Hoffman, 1973): browsers that eat mainly shrubs and trees, grazers that eat
mainly grass, and mixed or intermediate feeders that eat a mixture of grass, forbs, and browse.
The moose is a browser and has been classified as a seasonally adaptive concentrate selector
(Hofmann, 1989). Concentrate selectors have a relatively small ruminoreticular chamber and must
search for high-quality foods that will pass rapidly through the digestive system. Moose consume
plant species and parts (twigs and foliage) high in cell-soluble sugars that ferment rapidly in the
rumen. They generally avoid plants that are fibrous and require extensive breakdown in size
before passage from the rumen. Moose have a relatively narrow muzzle, prehensile lips and
tongue that allow them to select high-quality plant parts (Renecker and Schwartz, 2007). Moose
ferment (digest) mostly the soluble components of their food, and propel digesta rapidly through
their digestive system (Schwartz, 1992). Their digestive efficiency is regulated by forage
selection, rumination, gut morphology, and mechanisms controlling the rate of passage of food
(Schwartz, 1992). Moose can ingest and process high-quality foods more rapidly, (e.g., aquatic
plants eaten in summer), because both passage and digestion rates are enhanced (McArt et al.,
2009).
In the range of moose, plant species respond to seasonality by growing during the short summers
and entering a state of dormancy during long the winters. As plants change seasonally so does
their nutrient quality. Plants begin their growth phase in early spring, long before actual green-up
occurs. In general, spring and summer foods are 1.5 to 3 times more nutritious than winter foods,
depending on which constituent is examined (Schwartz and Renecker, 2007). Summer diets
contain excess digestible energy and protein, whereas winter diets generally are insufficient to
meet maintenance requirements (Renecker and Hudson, 1989; Schwartz et al., 1988; Schwartz et
al., 1987). As a result, feeding habits of moose vary by season, via a complex interaction of
internal physiological regulators and the external environment. There is an annual cycle of food
selection and intake, fat metabolism, metabolic rate, and body mass dynamics that is not driven
simply by food quality and availability (Schwartz, 1992). The gastrocentric hypothesis predicts
that large male moose will eat large amounts of low quality, fibrous foods, while smaller-bodied
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females will consume smaller amounts of higher-quality forage to meet the demands of
reproduction and lactation (Oehlers et al., 2011). Both sexes reduce food consumption and
metabolic rates in the winter, and operate at a net energy deficit by utilizing fat reserves (Miquelle
etal., 1992; Schwartz, 1992).
Protein and energy are considered the major limiting nutrients within the environment (Schwartz,
1992). Summer protein intake is critical for lactating female moose (McArt et al., 2009). Tannins
have a negative influence on forage quality, because they quantitatively reduce protein availability
(Robbins et al., 1987). In two areas of Alaska, browse quality differences were consistent with
observed differences in moose reproductive success (McArt et al., 2009). In recent years, the
productivity of DNPP moose has been significantly higher than that of moose in the Nelchina
Basin. A study of browse quality in the two areas found that, on average, nitrogen levels were 9%
lower and tannin levels 15% higher in Nelchina than in DNPP, resulting in a digestible protein
differential of 23%. The researchers concluded that the Nelchina moose population was nitrogen-
limited. In both systems, browse quality declined significantly as summer progressed, with
nitrogen levels decreasing and tannin levels increasing in all species of browse studied. In
comparison with early-summer forage, digestible protein had decreased by an average of 35% by
mid-summer and 70% by late summer (McArt et al., 2009).
High-quality summer forage, particularly near wetlands, allows nursing cows to regain body
condition and calves to grow so they can better escape predators and survive their first winter
(ADF&G, 20lid). During the spring-summer period, moose feed in aquatic habitats. In the
Copper River Delta, aquatic habitats produced about four times more forage than terrestrial
habitats, and the forage was more digestible (MacCracken et al., 1993). While some researchers
have linked the summer consumption of aquatic plants by moose to a craving for sodium (Jordan,
1987), the Copper River Delta data did not support that hypothesis. Those data suggested that
moose selecting aquatic forage were switching from an energy-maximizing to a time-minimizing
strategy (MacCracken et al., 1993). This is because aquatic plants are high in water content and
although a moose can fill it's rumen quickly, the relative quantity of dry matter consumed is less
than when eating the same amount of terrestrial vegetation, such as leaves. Although a relatively
small part of the wild moose diet, another forage type selected seasonally by moose is bark. Bark
stripping occurs mostly in winter, when there are a lesser number of twigs available due to snow
depth (MacCracken et al., 1997b). In DNPP, female moose also stripped bark in aspen-spruce
forests in May and June, coincident with birth and lactation (Miquelle and Van Ballenberghe,
1989). Studies have also identified certain plant species that moose avoid because of low
palatability due to chemical defenses in plants, such as black cottonwood (Populus balsamifera
trichocarpd) on the Kenai Peninsula (Weixelman et al., 1998) and white spruce (Picea glaucd)
both on the Kenai and in other parts of Alaska (Weixelman et al., 1998).
Moose in the Copper River Delta consume three different diets that vary among the seasons of
winter, spring/early summer, and late summer/fall (MacCracken et al., 1997b). Willow dominated
all three diets; the differences were related to the amounts of sweetgale (Myrica gale), marsh five-
finger (Potentilla palustris) and graminoids in the diet. Winter diets included sweetgale and alder
(Alnus spp.), which are both nitrogen-fixers, leading to relatively higher protein content.
Spring/early summer diets were most diverse, due to the increased use of emergent aquatic plants
such as marsh five-finger. Late summer/fall diets were least diverse, consisting almost entirely of
willow leaves and twigs (MacCracken et al., 1997b).
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Moose diets in DNPP were also found to vary by season. In the summer, seven species of willow
comprised a total of 81.5% of the diet (Van Ballenberghe et al., 1989). In that study, diamond-leaf
willow (Salixplamfolia) was eaten more than any other plant species in summer (45.7% of diet).
In contrast, willow comprised over 94% of the total diet of DNPP moose in winter (Risenhoover,
1989).
Moose can influence the composition and productivity of the terrestrial plant community through
browsing (Bedard et al., 1978). In DNPP, moose initiated positive feedback loops on their
environment through browsing (Molvar et al., 1993). Willows exhibited a growth response to
moose herbivory; specifically, leaf area was significantly greater at the site with high moose
density than at the site with low moose density. Annual biomass productivity per growing point on
willow stems increased with increasing browsing intensity on the plant as a whole, via release
from apical dominance. Moose also increase rates of nutrient cycling, as their urine and feces
transfer nutrients to soil. The organic content of soil can also be enhanced by moose, in turn
benefitting microbiota such as decomposers (Molvar et al., 1993). In a study in interior Alaska,
twigs re-growing from two-year old willow stems that had been browsed by moose had larger
diameters than those that had not been browsed in the previous year (Bowyer and Neville, 2003).
Browsing on felt-leaf willow did not have an effect on nitrogen content, digestibility, or tannin
content, which indicated that the willow did not exhibit a tannin-mediated inducible defense
system in response to herbivory (Bowyer and Neville, 2003).
Marine derived nutrients (MDNs) carried upstream by spawning salmon have implications for
nutrient flow into riparian habitats, and are thought to enhance growth and productivity therein
(Quinn et al., 2009). While it is plausible that MDNs might contribute to increased plant
productivity and thus benefit moose, evidence of this direct impact was not located in the scientific
literature.
Moose density is often associated with food abundance (Eastman and Ritcey, 1987; Joyal, 1987;
Oldemeyer and Regelin, 1987; Schwartz and Franzmann, 1989; Thompson and Euler, 1987). As
reviewed by Renecker and Schwartz (2007), forage biomass varies with successional age of
forests. In Newfoundland, available woody biomass increased from about 200 kg/ha in two-year-
old clear cuts to more than 2,000 kg/ha by eight years, at which time it peaked and subsequently
declined gradually (Parker and Morton, 1978). On the Kenai Peninsula, important browse species
peaked about 15 years after fire (Spencer and Hakala, 1964). The biomass of important browse
species in successional stands of forest has been estimated on the Kenai NWR; browse production
measured at 3, 10, 30, and 90 years post-burn was 37, 1,399, 397, and 4 kg/ha, respectively
(Oldemeyer and Regelin, 1987).
Behavior
Movements and Home Ranges- The ways in which moose use their environment both spatially
and temporally are of great interest to resource managers (Hundertmark, 2007). The dynamics of
animal movements and distribution in space and time are integral to behavioral, ecological,
genetic, and population processes. Thus, the attributes of the space occupied by individual
animals, both annually and seasonally (home ranges), patterns of movement within home ranges,
establishment of new home ranges by young moose and colonization of new habitats ^dispersal)
and movements between seasonal ranges (migration) must be considered in comprehensive
management programs.
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The size of a moose home range varies with the sex and age of the animal, season, habitat quality,
and weather. Two studies from Alaska generated the largest estimates of home range size,
although one of these (Grauvogel, 1984) included migratory locations in the estimate of seasonal
ranges, which can increase home range size significantly (Hundertmark, 2007). Moose in south-
central (Ballard et al., 1991) and northwest Alaska had mean seasonal ranges > 92 km2. With the
exception of home ranges of non-migratory adults in the later study, total home range sizes
exceeded 259 km . In contrast, estimates of annual ranges for moose in northwest Minnesota were
< 3.6 km2 (Phillips et al., 1973).
Seasonal ranges, when they exist, represent partitioning of the environment based on behavioral
and energetic constraints. Migratory moose (those that use separate winter and summer ranges)
use distinct seasonal ranges because they attempt to optimize their nutrient intake on summer
range, but winter conditions on these ranges may preclude occupation during some or all winters.
Moose that remain on the same range during winter and summer are termed non-migratory, and do
so because the environmental conditions permit their residence. A third seasonal range associated
with mating, occurs in autumn, but many investigators define this as part of the summer range
(Hundertmark, 2007). Breeding areas for tundra moose are typically in open habitats where
visibility is good. This is likely for behavioral purposes so bulls and cows can see each other as
they display. It may also afford some predator protection.
In several moose populations studied in Alaska, some individuals were non-migratory residents
while other individuals migrated seasonally. In the Copper River Delta, 8 of 15 collared females
were migratory, while two of five collared males were migratory (MacCracken et al., 1997b).
Moose in that area exhibited greater fidelity to their summer range than their winter range
(MacCracken et al., 1997a); winter severity influenced winter migratory behavior from year to
year in the Copper River Delta (Stephenson et al., 2006). Moose populations in south-central
Alaska (GMU 13, comprising the Nelchina and upper Susitna basins) also included both migratory
and non-migratory individuals (Ballard et al., 1991). Those moose exhibited three seasonal periods
of movement - autumn migration to wintering areas, spring migrations to calving areas or summer
feeding grounds, and early fall migrations to rutting areas (Ballard et al., 1991). In the Togiak
River drainage of the northern Bristol Bay area (GMU 17A, the), some collared moose were
resident whereas others migrated seasonally (Woolington, 2008). During a population estimation
survey in February 1995, 29 moose were documented moving westward from the upper Sunshine
Valley in GMU 17C (the lower Nushagak watershed) into GMU 17A (Woolington, 2008).
Cows with new-born calves restrict their movements for the first few weeks, after which they
gradually expanded their home range to approximate home range size of other adults (LeResche,
1974). In one study, cow-calf pairs had smaller summer home ranges than did other moose, and
calf movements increased exponentially with age during the first six weeks of life (Ballard et al.,
1980).
When differences in annual home range sizes are attributed to sex, males always are found to
occupy larger areas. In south-central Alaska males had significantly larger home ranges than did
females (Ballard et al., 1991). In northwestern Alberta researchers noted no difference between the
sexes, but did note the tendency of bulls to occupy larger winter and spring home ranges (Lynch
and Morgantini, 1984).
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The timing of seasonal migration has been observed to vary significantly among individuals from
several moose populations in Alaska. In the Nelchina Basin of south-central Alaska, individual
moose movements varied by months both in the initiation and the duration of winter migration
(Van Ballenberghe, 1977). Snow depth was an important factor that influenced winter migratory
behavior in that population. Cows with calves tended to migrate to wintering grounds earlier than
did males and cows without calves (Van Ballenberghe, 1977). During spring in the Nelchina
Basin, the initiation of migration varied substantially between individuals, but all individuals
migrated quickly once they started moving (Van Ballenberghe, 1977). Individual moose in south-
central Alaska (GMU 13) also initiated migration to wintering areas at very different times,
ranging from mid-August to mid-February (Ballard et al., 1991). Moose in GMU 13 exhibited
more variation in spring migration than Nelchina Basin moose did (Ballard et al., 1991). Dates of
spring migration ranged from March through mid-July; during some years, moose remained on the
winter range for calving. Subsequent movement to the summer range in mid-summer seemed
related to plant development (Ballard et al., 1991).
Moose in different areas of Alaska were found to migrate different distances seasonally and have
different annual home range sizes (Ballard et al., 1991; Gillingham and Klein, 1992; MacCracken
et al., 1997b) (Table 2). Moose on the Seward Peninsula of northwest Alaska migrate up to 80 km
seasonally (Gillingham and Klein, 1992). In south-central Alaska, the distance between winter and
summer ranges of migratory moose averaged 48 km, and ranged from 16 to 93 km (Ballard et al.,
1991).
Moose use of seasonal home ranges is traditional (Ballard et al., 1991). In south-central Alaska
GMU 13, only one of 101 radio-collared female adults dispersed from their traditional home range
during a 10-yr study period (Ballard et al., 1991). During the fall of 1978, that female relocated
177 km from her previous location (Ballard et al., 1991).
In the northern Bristol Bay region, some moose collared in GMU 17A since 2000 have moved
westward within GMU 17A and into the southern part of GMU 18 (Woolington, 2008). This is
thought to be part of a continued westward expansion into previously unpopulated moose habitat
(Woolington, 2008).
2. for In
Study area
Kenai
Peninsula
Migratory
status
M
N
N
Adult/M
Adult/M
Adult/F
Mean home range size (km2) Reference
Total Winter
137(56-185)
52 (34-64)
127(25^140) 63(13-184)
Summer
(Bangs etal., 1984)
36(2-152) (Bangs and Bailey,
Seward
Peninsula
M
Adult
1980)
938(236-1,932) 311(36-1,393) 324(41-1,323) (Grauvogel, 1984)
South-central
Southeast
N
I
N
M
N
Adult
Adult
F
F
Adult/F
218(91-350)
339 (205-593)
290(111-787)
427 (274-580)
28(9-51)
98 (36-223)
122 (21-334)
113(10^130)
147(15-375)
11(3-30)
93 (44-150)
210 (60-559)
103 (23^156)
263 (60-622)
14 (2-30)
(Ballard etal., 1991)
(Doerr, 1983)
Migratory status: M = migratory, N= nonmigratory, and I = intermediate. Data from Hundertmark (2007).
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Sexual Segregation and Grouping Behaviors- Bowyer et al. (2003) provide a succinct discussion
of sexual segregation in moose, and we paraphrase it here. Sexual segregation is the differential
use of space by the sexes outside the mating season (Barboza and Bowyer, 2000) and often
includes differential use of habitats and forages. Sexual segregation is especially pronounced in
moose and plays a crucial role in their ecology (Bowyer et al., 2001; Miller and Litvatitis, 1992;
Miquelle et al., 1992). In Alaska, male and female moose select habitats differently, leading to
their spatial segregation throughout most of the year (Oehlers et al., 2011)). Spatial segregation
occurs because adult males select habitats with greater forage abundance and females select areas
with more concealment cover during winter (Bowyer et al., 2001; Miquelle et al., 1992), while
cows with calves select denser cover and are more secretive than other age groups (Peek, 2007). In
southeast Alaska, female moose selected habitat that maximized high-quality forage while
minimizing predation risk, while male moose selected habitat that provided the highest forage
intake (Oehlers et al., 2011). In DNPP, summer habitat selection by adult females also varied,
depending on whether or not they had a calf (Miquelle et al., 1992). Females with calves remained
solitary and preferred forested habitats, which provided better cover from predators. Such
differences in habitat use between the sexes have implications for sampling of moose populations,
because it can affect the accuracy of sex and age ratio information obtained by direct observation
(Bowyer et al., 2002; Peek, 2007; Peterson, 1955; Pimlott, 1959).
The degree to which Alaskan moose segregate by gender varies by season. In DNPP, sexual
segregation is most common in winter, when only 19% of all observed groups had both large
males and females (Miquelle et al., 1992). Spatial segregation in that study was most extreme
during a deep-snow winter, when only large males could access forage at higher-elevation Jenny
Creek due to their larger body size (Miquelle et al., 1992).
The effect of habitat enhancement on sexual segregation was studied in interior Alaska after
mechanical crushing of willow stands (Bowyer et al., 2001). In that study, males occurred
predominantly on the more open, disturbed area during winter, whereas females and young used
older stands of willow, where dense vegetation offered substantial concealment from wolves.
Females and young faced a tradeoff between foraging on a greater abundance of food in the
disturbed area and a reduced risk of predation in the mature stand (Bowyer et al., 2001).
The way in which moose, either individually or in groups, partition their habitats and associate
with other moose can be informative in determining the needs of the various segments of the
population (Hundertmark, 2007). Moose have been referred to as "quasi-solitary," and large
groups are uncommon (Houston, 1968). The tendency of moose to lead a solitary life or to occur
in groups depends on their age, sex, and reproductive status, and varies by season. Molvar and
Bowyer (1994) note that Alaskan moose are more gregarious than moose from Eurasia and
suggest that the formation of social groups is a recently evolved adaptation in Alaska moose in
response to a relative abundance of predators and to relatively open terrain. In DNPP, larger
groups ventured farther from cover but were less efficient at foraging due to inter-individual
aggression (Molvar and Bowyer, 1994).
Cows with calves are consistently the most solitary members of the population, probably because
of predator avoidance (Hauge and Keith, 1981; Hundertmark, 2007; Miquelle et al., 1992).
Alaskan female moose with calves are nearly always solitary at the time of birth (Miquelle et al.,
1992; Molvar and Bowyer, 1994). Females without calves are more likely than males to be
40
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solitary during the early summer, but they become more gregarious as the summer progresses
(Miquelle et al., 1992). In DNPP, during the summer, females without calves were seen alone only
23% of the time (Miquelle et al., 1992). During the months of June through August, male moose
in DNPP were consistently gregarious (Miquelle et al., 1992). When in a group, small males were
more likely than large males to be in a group that included females, at all times except the rut and
post-rut (Miquelle et al., 1992).
In south-central Alaska GMU 13, calves separated from their parents at an average age of 14
months (Ballard et al., 1991). In that study, 33% of yearlings and one two-year old were observed
in one to six temporary re-associations with their mothers after their original dispersal. Calves
were more likely to re-associate with their mother if she was not caring for a new calf (Ballard et
al., 1991).
Matins and Maternal Behaviors- Moose in North America have two general forms of mating
behavior. In the taiga of Canada, moose have a serial mating system, in which bulls search for
cows in heat by traveling widely, while calling and thrashing their antlers (Bubenik, 2007). The
bull digs shallow pit holes in which he urinates, but they are randomly located and seldom found
on the same spot in successive years. For all cows to be bred during the short three-week mating
season, the serial mating strategy requires a relatively high number of bulls. Bubenik (2007)
concluded that due to differences in climatic conditions of the periglacial tundra, the tactic of
serial mating was replaced by communal or harem mating in tundra moose. Toward the end of
August, a prime tundra bull settles in a mating area of about 10 km2. Rutting areas appear to be
traditional (Bubenik, 2007). In early September, bulls begin scent-urinating on trails and in pit
holes. Two prime bulls may share a harem when it contains eight or more cows. During the eight
to ten days of breeding in the harem, a tundra bull probably can fertilize as many or more cows
than the taiga bull does during the entire three-week rut because the tundra bull can mate with
each female in his harem without traveling long distances to locate a new female.
Many mammals have evolved seasonal reproductive patterns that ensure adaptation to predictable
annual changes in the environment. Moose exhibit marked seasonal changes in reproductive
behavior that reflect adaptations to yearly fluctuations in food availability to ensure favorable
conditions for rearing young (Schwartz, 2007). This means that moose do not reproduce all year
long, but only during one season (autumn). By breeding in the fall, it insures calves are born in
spring when forage is high in nutrient quality and the cow has a high probability of producing
enough milk to successfully raise the calf. Day length may provide the clue to annual timing of the
breeding season. Length of the breeding season is relatively short for moose. Since it is difficult to
determine the exact date of breeding under natural conditions, few studies provide detailed
information. Researchers with the most robust data sets each concluded that moose exhibit a very
well-defined breeding season, as judged by conception dates and the spread of observed breeding
(Crichton, 1992; Schwartz and Hundertmark, 1993; Thomson, 1991). The mean date of breeding
in British Columbia ranged from October 5 to 10, with a standard deviation of five days
(Thomson, 1991). The average day of breeding in Manitoba was 29 September and 93% of all
females were bred by 12 October (Crichton, 1992). The average breeding date in Alaska was 5
October, with a range from 28 September to 12 October. There was very little difference among
years in all studies, suggesting that photoperiod, rather than weather influences rut timing.
Synchrony of the rut has also been observed in DNPP. Over a twelve-year interval, rutting
consistently occurred during the brief period from 24 September through 5 October (Van
Ballenberghe and Miquelle, 1993).
41
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Moose cows across North America give birth during a relatively short period of time. The peak
birthing period occurs from about 15 May through 7 June (Schwartz, 2007). In DNPP, timing of
birth in moose was consistent from year to year, despite variation in climate between years
(Bowyer et al., 1998). Birth timing exhibited "extreme synchrony" and Bowyer et al. (1998)
hypothesized that moose were tracking long-term patterns of climate in the area to time
reproduction. Hence, there is concern that moose will be vulnerable to climate change even before
extensive changes to vegetation occur (Bowyer et al., 1998).
In DNPP, the primary drivers influencing birth site selection were microclimate, forage abundance
and quality, and risk of predation (Bowyer et al., 1999). Birth sites were not re-used, and some
females appeared to behave unpredictably shortly before giving birth, perhaps in an attempt to
thwart predators (Bowyer et al., 1999). Proximity to human development did not influence birth
site selection. Moose preferred birth sites with abundant willow, high visibility to detect predators,
and a southeasterly exposure that would be warmer and drier (Bowyer et al., 1999). Bark stripping
was common around birth sites, because females seldom travelled more than 100 m from their
young and hence rapidly depleted the birth site's forage (Bowyer et al., 1999).
Activity Budgets- Moose spend most of their active life foraging. Seasonal rates of forage intake
tend to mimic the cyclic nature of energy metabolism in moose (Regelin et al., 1985), with higher
rates of activity and intake in spring and summer and reduced rates during winter. Activity
budgets tend to follow a similar pattern. Activity budgets have been studied for DNPP moose
during winter (Risenhoover, 1986) and spring/summer (Van Ballenberghe and Miquelle, 1990).
DNPP moose exhibited low activity levels from January through April, when they were active, on
average, only 27.3% of the time (6.5 h/d) (Risenhoover, 1986). Risenhoover (1986) found that
activities associated with resting and foraging constituted 99.3% of the time of DNPP moose in
winter. In contrast, Miquelle et al. (1992) found that small males in DNPP spent some of their
active time engaged in social behavior during winter. In winter and early spring in DNPP, moose
exhibited a polyphasic pattern alternating between foraging and bedding, with about six cycles per
24 hours (Risenhoover, 1986).
Following their relative inactivity in winter, DNPP moose increased their metabolic rate in April,
as evidenced by the onset of antler development in males and increased mobility of cows
(Risenhoover, 1986). Activity increased during May to a peak in early June, then began to decline
until mid-August (Van Ballenberghe and Miquelle, 1990). DNPP moose were active 12.8 h/d at
the peak, and activity had declined to 9 h/d by late summer (Van Ballenberghe and Miquelle,
1990). In summer DNPP moose spent about equal amounts of time feeding, resting and
ruminating during each 24-h period (Van Ballenberghe and Miquelle, 1990). When comparing
winter activity budgets (Risenhoover, 1986) to summer activity budgets (Van Ballenberghe and
Miquelle, 1990) in DNPP, large differences were observed (Table 3).
42
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Table 3. Moose activity in spring/summer' In
i
Activity Parameter
Total active time/day (hr)
Total resting time/day (hr)
# Activity bouts/day
Duration of activity bouts (min)
Duration of resting bouts (min)
Foraging time/day (hr)
Rumination time/day (hr)
Winter
6.5
17.5
5.7
68
178
4.9
11.7
Spring/Summer
10.1
13.9
8.2
73
97
7.5
6.7
1 = "Winter" is January through April; data from Risenhoo ver 1986
2="Spring/Summer" is 1 May through 15 August; data from Van Ballenberghe
and Miquelle (1990)
On the Seward Peninsula of northwestern Alaska, moose winter activity time allotments were
43.2% feeding, 42.8% bedding, 8.4% walking, 4.4% standing, and 1.2% other (Gillingham and
Klein, 1992). Walking time was far greater than reported during winter in DNPP (< 1%)
(Risenhoover, 1986). Gillingham and Klein (1992) attributed this difference to Seward Peninsula
moose using the Kuzitrin River as a feeding and movement corridor during winter. The use of a
narrow, linear feature, such as a river bottom, means that a moose needs to travel farther up and
down the river to obtain food, as opposed to feeding in a large, non-linear area. There are at least
two other differences between DNPP moose and Seward Peninsula moose. While there is an
abundance of predators in DNPP (wolves and bears), there are no predators of moose on the
Seward Peninsula in winter (Gillingham and Klein, 1992). Also, moose activity was highly
synchronized during mid-afternoon in late April on the Seward Peninsula; presumably due to heat
stress (Gillingham and Klein, 1992). In contrast, there was no significant correlation between
mean daily temperature and daily activity level in DNPP during winter (Risenhoover, 1986).
On the Copper River Delta, inactive bout duration of moose was shortest on the west delta, which
had the highest estimates of forage abundance and quality among the three areas studied
(MacCracken et al., 1997a). It was suggested that the relative duration of inactive bouts might be
useful as an index of habitat quality for moose.
Interspecies Interactions
Boer (2007) provided an excellent review of the interspecific relationships between moose and
other species. Interspecific interactions between moose and other species take on one or more of
the following general forms: competition, parasite-mediated competition, predation, and
commensalism (Boer, 2007). Due to the diversity of habitats, species combinations, and
abundance of sympatric species, a variety of competition mechanisms operate throughout moose
range. Of the interspecific interactions possible, competition is the most obvious one influencing
moose habitat use and distribution (Boer, 2007). Throughout their North American range, moose
compete with an array of other ungulate species. However, in the Nushagak and Kvichak
watersheds, caribou are the only other ungulate species abundant enough to consider. Direct
competition between moose and caribou appears limited and insignificant (Davis and Franzmann,
1979). Food preferences of moose and caribou coincide to some degree, but the diet of caribou
appears to be more specialized. Caribou consume forbs and deciduous vegetation and lichens in
43
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winter (Darby and Pruitt, 1984; Servheen and Lyon, 1989). Moose primarily consume browse, but
also use forbs and deciduous vegetation during summer (Dodds, 1960; Eastman and Ritcey, 1987).
As reviewed by Boer (2007), in multi-prey systems, moose and caribou populations may influence
each other indirectly. Increasing moose numbers in western and central portions of DNPP have
resulted from increased availability of caribou as alternate prey for wolves (Singer and Dalle-
Molle, 1985). In the eastern section of that park, migrating caribou were available as prey for only
a brief period of time, and therefore, they were not a particularly important factor of the area's
prey base. In that area, moose populations have declined. Moose are the primary prey of wolves in
other areas of Alaska as well (Gasaway et al., 1983), although others have attributed an increase in
moose numbers in northern Alaska to a preference by wolves for caribou (Coady, 1980).
Interspecies population dynamics have been studied in several areas of Alaska with multiple
predators and multiple prey species. These relationships can be quite complex and can vary based
on both abiotic and biotic factors within the ecosystem. None of the formal scientific interspecies
studies reviewed here were conducted in the Nushagak and Kvichak watersheds.
In Alaskan ecosystems with multiple predators, bears were responsible for more moose calf kills
than were wolves. Black bears can be significant predators of moose calves (Franzmann et al.,
1980). Of 47 radio-collared neonatal calves on the Kenai Peninsula, black bears killed 34%,
whereas brown bears and wolves each killed 6% (Franzmann et al., 1980). In the western Interior,
near McGrath, black bears were also the dominant source of predation mortality of calves during
six out of seven years studied; wolves and brown bears were secondary predators in that system
(Keech et al., 2011). In contrast, brown bears were the primary predators of moose calves in a
south-central Alaska study, causing 73% of calf mortality (Ballard et al., 1991). Brown bears were
also the primary predator in east-central Alaska (GMU 20E), where 79 to 82% of radio-collared
moose calves died by the age of eleven months (Gasaway et al., 1992). In that study, 52% of
moose calves were killed by brown bears, 12 to 15% of calves by wolves, and 3% by black bears.
Several studies have compared the causes of calf mortality in the nutritionally unproductive 1947
burn and in the productive high-quality habitat of the 1969 burn on the Kenai Peninsula, Alaska
(Franzmann and Schwartz, 1986; Schwartz and Franzmann, 1989; Schwartz and Franzmann,
1991). Black bears killed 34 and 35% of the calves, respectively, whereas wolves and brown bears
killed 5 to 13%, respectively. Total calf mortality from all causes ranged from 51 to 55%. Moose
densities were four times greater in the 1969 burn area (370/100 km2) and the population was
increasing, whereas the population in the 1947 burn was about 100/100 km and declining. The
investigators concluded that habitat quality had a significant impact on reproductive rate and
population growth. The moose population in high-quality habitat (1969 area) was capable of
sustaining this level of predation and continued to grow, whereas the population in poor habitat
(1947 area) was not.
Wolves appeared to select for moose calves in some areas and seasons in Alaska, but not in others.
In south-central Alaska, moose calves were taken in proportion to their abundance in the overall
population during May through October (Ballard et al., 1987). In contrast, during November
through April in that study, wolves preyed on moose calves selectively. During those months
calves were only 12 to 20% of the moose population, but they consisted of 40% of moose kills by
wolves (Ballard et al., 1987). During autumn in northwest Alaska, wolves displayed a lack of
selectivity for moose calves, which were killed in proportion to their relative abundance within the
44
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population (Ballard et al., 1997). On the Kenai Peninsula, wolves killed mostly moose calves
(47%), yearlings, and older adults (Peterson et al., 1984). Half of moose adults killed by wolves
during that study were more than twelve years of age. Wolf predation on moose calves was
highest during the winter with deepest snow, and calves killed after 1 January were commonly
malnourished, with bone marrow fat content <10%.
In east-central Alaska (GMU 20E), predation was the primary cause of non-hunting deaths for
yearling and adult moose (Gasaway et al., 1992). Of 46 such moose that died from 1981 through
1987, 89% were killed by brown bears or wolves, 9% died from antler wounds or locked antlers,
and 2% drowned. Peterson et al. (1984) examined the incidence of debilitating conditions among
109 wolf-killed adult moose on the Kenai Peninsula. They found that 20 such moose had moderate
or severe periodontitis, 14 had arthritis, one had a broken leg, and one had a leg wedged between
trees. Of 40 wolf-killed adult moose assessed for bone marrow fat content, four had levels <20%,
indicating severe malnutrition (Peterson et al., 1984).
Wolves in different regions in Alaska displayed different relative preferences for moose and
caribou as prey. In south-central Alaska, moose were the primary prey of wolves, constituting
38% of observed kills, while caribou was the second most important prey at 21% (Ballard et al.,
1987). In northwest Alaska, caribou was the preferred prey of wolves (Ballard et al., 1997). In
January through April 1988, when caribou were abundant, 92% of observed ungulates killed by
wolves were caribou. In contrast, in 1989 and 1990 when caribou were less abundant, they
constituted 11% and 48% of observed ungulate kills, respectively (Ballard et al., 1997).
Estimated kill rates for wolf packs on the Kenai Peninsula varied from one moose/3.1 days to one
moose/21.4 days (Peterson et al., 1984). The average kill interval in winter for Kenai wolf packs
with more than two members was 4.7 days. In 38 wolf-moose encounters observed on the Kenai
Peninsula, wolves succeeded in killing only two of the moose (Peterson et al., 1984).
Ballard et al. (1997) speculates that the recent occurrence of moose (since the 1940s) in northwest
Alaska has altered the historical migratory patterns of wolves in that area. There is evidence that
wolves in northwest Alaska used to migrate with the caribou herds, but now they do that only
when alternate prey (moose) numbers are insufficient.
Mortality, Productivity, and Survivorship
Understanding the dynamics of a population requires knowledge of how many individuals it
contains, how fast it is increasing or decreasing, its rate of production of young, and its rate of loss
through mortality (Van Ballenberghe and Ballard, 2007). Moose populations increase by the
addition of calves born to the population each year and decrease by the loss of animals. Death can
occur from the moment of birth. Moose die from a variety of causes including hunting, predation,
starvation, accident, drowning, vehicle collision, parasites, and disease. Mortalities are generally
divided into two major categories: human-caused or natural. Moose populations are adaptable to
artificially disturbed habitats, and therefore are often found in close proximity to roads, major
highways, and railways. But this association is far from compatible (Child, 2007). In populated
areas of Alaska a large number of moose are killed each year by collisions with motor vehicles
and trains (Bowyer et al., 2003; Child, 2007). For example, from 1963 to 1990, 3,054 moose were
killed on the Alaska Railroad, with annual losses ranging from seven to 725 (Modafferi, 1991). In
the severe winter of 1989-1990, deep snow caused more moose to travel on roads and railroads
45
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and fatalities thereon exceeded the previous record by more than 100 animals. In the Willow-
Talkeetna area during the same winter, the number of railroad kills represented a 70% loss from
the resident population (Schwartz and Bartley, 1991). Other sources of mortality include sport and
subsistence hunting and poaching (Woolington, 2004).
Prime adult moose tend to have very high rates of survival because they are not as vulnerable to
natural causes of mortality when compared to younger (calves) and older age classes. Survival
rates are generally estimated by radio collaring individuals and following them for some period of
time (Van Ballenberghe and Ballard, 2007). Ballard et al. (1991) provided data on mean annual
survival rate from a sample of radio-collared adult female moose during a ten-year period. From
25 to 80 moose per year were followed in a study area where hunting of cows was prohibited.
Annual survival rates were estimated at 94.8%. Data from yearling females spanned four years
with two to 22 individuals per year collared, and annual survival rates averaged 95.1%. Annual
survival of yearling and adult males averaged 75.4 and 90.9% respectively, with hunting the major
mortality factor. On the Kenai Peninsula, researchers followed 51 radio-collared females for six
years and reported a 92% annual survival rate (Bangs et al., 1989). Survival of cows aged 1 to 5
years was estimated at 97% and 84% for females aged 16 to 21 years. Hunting was not a
significant cause of mortality of the study population. As reported by Van Ballenberghe and
Ballard (2007), various other studies using radio-collared moose have reported annual survival
rates of adults ranging from 75 to 94%, depending upon the extent of human hunting. In general,
starvation and wolf predation during severe winters causes the greatest mortality in older age
classes (Ballard et al., 1991; Bowyer et al., 2003); moose weakened from starvation are
particularly vulnerable to wolf predation. Bull moose occasionally wound each other during the rut
and die from these wounds (ADF&G, 201 Id).
As reviewed by Van Ballenberghe and Ballard (2007), hunting is a major limiting factor of many
moose populations throughout the world. In fact, hunting pressure can reduce moose population
density (Crete et al., 1981). In Quebec, where natural mortality apparently was low, harvest rates
as high as 25% were reported (Crete, 1987). Moose harvest rates ranging from 2 to 17% have also
been reported for various other parts of North America (Crete, 1987). In concert with other factors,
including severe winters, high harvest rates have contributed to moose population declines in
Alaska (Gasaway et al., 1983). In addition, hunting can significantly reduce the number of bulls,
perhaps sufficiently to reduce the level of first-estrus conception (Bishop and Rausch, 1974).
When, due to heavy hunting pressure, there are fewer than ten bulls per 100 cows, some cows
simply may not encounter a bull early in the mating season. Breeding early in the mating season
means the rut would be synchronous and calving would therefore be synchronous. If a cow mates
late in the breeding season, the result would be later calving in the spring. In some European
environments, where severe winters, predation and nutritional stress are absent, moose harvest as
high as 50% of the winter population is sustainable (Cederlund and Sand, 1991). Most North
American moose populations harvested at this rate would decline sharply. In nearly all areas
where hunting is legal, harvest is managed under sustainable principals, so hunting mortality
seldom results in unintended population declines (Timmerman and Buss, 2007).
Within the Nushagak and Kvichak watersheds both hunter numbers and moose harvest have
increased. Correlated to a four-fold increase in moose hunters in GMU 17 from 1983 to 2006
(from 293 to 1,182), reported moose harvest tripled (from 127 to 380). In GMU 17B (the upper
Nushagak watershed), the reported harvests for the past five years, when data were available,
ranged from 113 to 183, with a mean annual harvest of 149 moose. In GMU 17C (the Togiak
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watershed), the five-year mean annual harvest was 224, with a range of 193 to 251 (Woolington,
2008). Hunters must harvest moose with antler spreads of no less than 50 inches. The largest
antlers reported exceeded 69 inches.
Juvenile moose tend to die at higher rates than adults. Calves are typically the most vulnerable age
class. Calf moose mortality can be divided into two general time periods when mortality is
highest: from birth to about six months of age, and from about nine months to one year of age
(Van Ballenberghe and Ballard, 2007). These periods correspond roughly to particular
vulnerabilities, specifically, to bear and wolf predation in the first period and hunting (in some
areas), wolf predation, and winter starvation in the second period. According to Van Ballenberghe
and Ballard (2007), neonatal mortality varies greatly, depending on several factors, most notably
the extent of predation. Several studies of radio-collared moose calves have documented that
predators may account for up to 79% of newborn deaths and that survival during the first eight
weeks of life may be as low as 17% (Ballard et al., 1981; Ballard et al., 1991; Franzmann et al.,
1980; Gasaway et al., 1992; Larsen et al., 1989; Osborne et al., 1991). Further losses during the
first year of life may result in annual survival rates as low as 10% (Van Ballenberghe and Ballard,
2007). In south-central Alaska, Ballard et al. (1991) observed that brown bears caused the majority
of natural death of calves younger than five months of age, whereas, on the Kenai Peninsula,
Franzmann et al. (1980) documented that black bears were the greatest cause of moose calf
mortality.
Moose breed in late September to early October (Van Ballenberghe and Miquelle, 1993) and adult
females give birth to one or two (twin) calves in late May-early June each year (Schwartz, 2007;
Testa et al., 2000). Production of moose calves is the result of a complex chain of biological
processes including estrus cycles, rutting behavior, fertilization, gestation, pre-partum events and
birth (Boer, 1992; Schwartz, 2007; Van Ballenberghe and Ballard, 2007). Fecundity, or
productivity of individual moose, is related to sexual maturation and a broad array of ecological
factors affecting food supply, forage quality, and weather that affect the physiological status of
females. These factors influence ovulation, pregnancy rates, litter size, and fetal sex ratios.
Ultimately, fecundity and subsequent survival of young determines recruitment rates and
population trends, which are important factors in moose population dynamics.
Reproductive tract studies have shown that female moose do not ovulate during the mating season
in the first year of life and therefore do not produce calves as yearlings. Cows may or may not
breed in their second year, depending on body mass (Saether, 1987). Most cows are sexually
mature at around 28 months of age and females continue to breed to the end of their life span at
around 18 years of age (ADF&G, 201 Id; Schwartz, 2007). Litter size in moose ranges from one to
three (Peterson, 1955), but litters greater than two are rare (Coady, 1982). Body condition of
female moose (as influenced by habitat quality) has correlated strongly with twinning rates in
several diverse moose populations (Franzmann and Schwartz, 1985). In an area known to contain
abundant high-quality food resources on the Kenai Peninsula, up to 70% of cows with calves one
year had twins the next year. This contrasts to other populations, in which twinning rates as low as
5% were reported (Houston, 1968; Markgren, 1969; Pimlott, 1959), but some of the estimates may
have considered post-natal mortality. The data suggest that twinning rates exceeding 40 to 50%
are uncommon for moose populations strongly limited by nutrition. Twinning frequency is a good
indicator of cow health condition and habitat quality (Dodge et al., 2004). Calves that survive
predation from bears in the summer wean in August, but will remain with their mother until the
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next calf is born the following spring (Schwartz, 2007), or for an additional year if no new calf is
born (Testa, 2004).
Population, Subpopulations, and Genetics
The number of animals in a population is one measure of abundance, but is only useful when the
geographic boundaries of an area are well defined, because that allows biologists to estimate
density (the number of individuals per unit area), which is a more robust parameter. Moose
population densities were compared by Gasaway et al. (1992) over very large areas (>2000 km2)
of generally continuous moose habitat across a broad area of Alaska and the Yukon Territories.
They noted that smaller sites exhibited high variability in prey and predator densities and in
habitat quality, making it more difficult for realistic comparisons. They focused their comparisons
on the post-hunt, early winter season thereby enhancing comparability further. The mean density
of moose from 20 populations was estimated at 0.148/km (range 0.045-0.4177 km ) in areas
where predation was thought to be a major limiting factor of moose. Densities of 16 other
populations in the same area, where predation was not limiting, averaged 0.66 moose/km2 (Van
Ballenberghe and Ballard, 2007). Ballard et al. (1991) provided 29 moose density estimates from
Alaska, including some populations studied by Gasaway et al. (1992); they ranged from 0.05 to
1.247 km2.
ADF&G estimates the total population of moose in Alaska at 175,000 to 200,000 animals
(ADF&G, 20 lid). The 2004 population estimate for the study area of the Nushagak and Kvichak
watersheds was 8,100 to 9,500 moose (Butler, 2004; Woolington, 2004). This estimate was based
on population data from ADF&G GMUs 17B, 17C, 9B and less than half the area of GMU 9C,
outside the Katmai National Park (Figure 3).
Moose are relatively new inhabitants in the Bristol Bay area, possibly having migrated into the
area from middle Kuskokwim River drainages during the last century (Woolington, 2004). Moose
were either not present or were sparse in the northern Bristol Bay area until the turn of the
twentieth century, and even then the moose population did not increase until three decades ago
(Butler, 2004; Woolington, 2004). Suspected reasons for low moose populations in the Bristol Bay
region are heavy hunting pressure, particularly on female moose in the western part, and bear
predation in the eastern part (Butler, 2004; Woolington, 2004). Over the last 25 years, managed
harvesting, predator control, an increase in caribou herds as an alternative predator food source,
and consecutive mild winters have led to an increase and expansion of the moose population
westward (Butler, 2004; Woolington, 2004).
The largest moose population in the study area is in the Nushagak drainage; the upper watershed
(GMU 17B) has an estimated 2,800 to 3,500 moose, while the lower drainage (GMU 17C) has an
estimated 2,900 to 3,600 moose (ADF&G, 201 Id). These moose comprise about 73% of the total
moose in the Nushagak and Kvichak watersheds. The Nushagak drainage has large, healthy areas
of riparian habitat, a major component of which is felt-leaf willow, a preferred browse species
(Bartz and Naiman, 2005). The number of moose in the Kvichak watershed was estimated at 2,000
in GMU 9B, and less than 400 moose in the portion of GMU 9C outside Katmai National Park
(Butler, 2008).
Fall trend counts have been notoriously unreliable in providing consistent data on moose
populations in GMU 17 (Woolington, 2008). Suitable survey conditions, including complete snow
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coverage, light winds and moose movements onto winter range rarely occur before antler drop.
Regular population estimation surveys of portions of the unit during late winter provide the best
population information; unfortunately they do not provide reliable information on sex and age
composition.
Moose population estimates in the northern Bristol Bay area are produced by a spatial statistics
stratification model, which uses harvest ticket data from sport and subsistence hunters (Butler,
2004; Woolington, 2004). The ADF&G, Division of Subsistence suspects there is a considerable
unreported subsistence harvest as well as illegal harvest occurring in some regions of Alaska
(ADF&G, 201 Id). Illegal harvest of moose in Unit 17 was probably more of a problem in the past
than during recent years. Unit residents used to actively pursue moose with snow machines during
the winter and spring, when both male and female moose were taken. Attitudes have changed
following considerable efforts by State and federal management agencies, working with local
communities to help hunters see the benefits of reducing illegal moose kills. It is now common to
see moose near local villages throughout the winter (Woolington, 2008).
Human Use (Subsistence, Recreation)/Interaction/Management)
In Alaska 7,400 moose were harvested in 2007. Residents harvested 6,750 moose and 685 were
taken by non-resident hunters (ADF&G, 201 Id). The harvest of 7,400 moose yields approximately
3.5 million pounds (1,587,573 kg) of meat.
Harvest records from ADF&G for 1983 to 2002 indicate that GMU 9 and 17 provided 7% of the
total moose harvest in Alaska (BLM 2007). According to ADF&G, Division of Subsistence
(http://www.adfg.alaska.gov/sb/CSIS/) local subsistence hunters from King Salmon, Naknek, and
South Naknek harvested 19 moose in GMU 9B in 2007; total meat harvested was estimated at
10,206 pounds (4,629 kg). In unit 17B, local residents from Igugig, Koliganek, and New Stuyahok
harvested 88 moose in 2005 (last year of available data); total meat harvested was estimated at
48,208 pounds (21,867 kg). Likewise, residents from Naknek and South Naknek harvested 4
moose from unit 17C with a total of 5,357 pounds (2,430 kg) of meat. In total, subsistence moose
meat accounted for 63,771 pounds (28,987 kg) of meat with an average of 128 pounds (158 kg)
harvested per household (Table 4).
Moose are an important subsistence food species for people residing in the area served by the
Bristol Bay Area Health Corporation (Ballew et al., 2004). In a survey about traditional food
consumption conducted in 2002, 86% of respondents from that region reported consumption of
moose meat within the past year, at a median per capita consumption rate of five pounds (2.3 kg)
per year (Ballew et al., 2004). Moose was the third greatest subsistence source of meat to residents
of that region; residents reported eating more salmon and caribou than moose. Subsistence
statistics (Table 4) also suggest that, on average, a high percentage of individuals from villages in
the area (38%) attempted to harvest a moose, with about 20% succeeding. Additionally, about
24% of individuals reported sharing their moose with others, while 44% received meat from
others.
In addition to being a source of subsistence meat, moose also contribute to the local economy,
through jobs created as a result of non-resident hunters seeking a remote fly-in or boat-in
experience to take a trophy moose.
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Table 4. Subsistence statistics for moose harvest in Alaska Dept. of Fish and Game, game management units (GMU) 9B, 17B, and 17C.
GMU
09B
09B
09B
17B
17B
17B
17C
17C
Total
Mean
Community
name
King Salmon
Naknek
South Naknek
Igiugig
Koliganek
New Stuyahok
Naknek
South Naknek
Study
year
2007
2007
2007
2005
2005
2005
2007
2007
Using
(%)
33
48
29
100
86
94
48
29
58
Attempted
harvest
(%)
31
23
24
50
68
65
23
24
38
Successfully
harvested
(%)
10
5
0
42
50
51
5
0
20
Shared
meat
(%)
10
5
0
75
54
43
5
0
24
Received Reported
meat harvest
(%)
24
47
29
67
46
65
47
29
44
(#)
5
4
0
6
16
30
4
0
Estimated
harvest
(#)
9
10
0
6
24
58
10
0
Reported
harvest
Ibs
2,700
2,160
0
3,240
8,640
16,200
2,160
0
35100
Mean
Estimated harvest/
harvest household
Ibs
4,849
5,357
0
3,510
12,960
31,738
5,357
0
63771
(#)
55
29
0
270
309
331
29
0
128
(Data are from 2005 or 2007 and represent the most recent information available.) (http://www.adfg.alaska.gov/sb/CSIS/)
50
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BARREN GROUND CARIBOU
Introduction
Alaska is currently home to 31 herds of wild caribou (Rangifer tarandus granti\ with a
combined population of approximately 760,000. Caribou herds are defined by their traditional
and predictable use of calving areas that are separate and distinct from the calving grounds of
other herds (Skoog, 1968). Use of other seasonal ranges is variable and less traditional. Caribou
from different herds may overlap on seasonal ranges other than calving areas (Cameron et al.,
1986). Historically, most caribou herds have fluctuated widely in numbers and in use of range
(Skoog, 1968).
Adult bull caribou in southwestern Alaska usually weigh between 350 and 450 Ibs (159 to 182
kg), while females weigh between 175 and 225 Ibs (80 to 120 kg) (ADF&G, 1985). Body weight
can vary with environmental and nutritional conditions (Cameron, 1994; Valkenburg et al.,
2003). Caribou are the only members of the deer family in which both males and females grow
antlers. Bulls begin to shed the velvet on their antlers between late August and early September,
marking the start of breeding season. The largest bulls begin shedding their antlers in late
October, with smaller bulls losing their antlers later in the winter. Females shed velvet in
September (Skoog, 1968). Pregnant females usually keep their antlers until the calving season in
the spring, while non-pregnant females lose their antlers about a month before calving begins.
Some females never grow antlers (Whitten, 1995). Caribou populations throughout the Bristol
Bay region have declined recently and body weights and antler sizes are now relatively low. In
the past the area produced large-bodied animals with record book antlers (Valkenburg et al.,
2003).
Population History of Caribou in the Upper Bristol Bay Region
Historical accounts from the early 1800s indicate that caribou were plentiful in the Bristol Bay
region. There may have been a large herd that ranged from Bristol Bay across the Kuskokwim
and Yukon deltas all the way to Norton Sound. By the late 1800s caribou throughout this area
had declined dramatically. Caribou numbers may have increased in the early 1930s, but were
declining again by the late 1930s. Domestic reindeer were brought to the Bristol Bay region in
the early 1900s, but by the 1940s, reindeer herds were widely neglected and either died out or
were assimilated into wild caribou populations (Skoog, 1968; Woolington, 2009a). Caribou in
the Nushagak drainage remained relatively scarce into the 1970s, at about 10,000-15,000 animals
(Woolington, 2009a).
Over the past thirty years, caribou herds in southwest Alaska have continued to undergo
significant changes in population numbers. The Nushagak and Kvichak drainage basins are now
used primarily by caribou from the Mulchatna herd. The Mulchatna herd grew rapidly during the
1980s and 1990s, from a population of about 18,600 animals in 1981 to a peak of approximately
200,000 in 1997. By 1999 the Mulchatna herd had declined to 175,000 and it continued to
decline, to approximately 30,000 in 2008 (Valkenburg et al., 2003; Woolington, 2009a). As the
Mulchatna herd grew, it overlapped with and eventually assimilated the much smaller Kilbuck
(or Qavilnguut) herd that formerly ranged infrequently into the western part of the Nushagak
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drainage. By the late 1990s the Kilbuck herd had ceased to function as a distinct population
(Woolington, 2009a).
The Northern Alaska Peninsula herd recovered from a population low of about 2,000 in the late
1940s to about 20,000 in 1984. The population remained at about 15,000 to 19,000 through
1993, but has since declined steadily to about 2,000 to 2,500 today (Butler, 2009a). For the most
part, caribou of the Northern Alaska Peninsula herd remain well south of the Kvichak drainage.
However, from 1986 to 2000 many caribou from the Northern Alaska Peninsula herd wintered in
the Kvichak drainage, south of Lake Iliamna (Butler, 2009a). In the late 1980s and early 1990s,
the Kvichak drainage was also used by far greater numbers (up to 50,000) of Mulchatna caribou
(Woolington, 2009a). The two herds always returned to their traditional calving and summer
ranges and remained distinct (Butler, 2009a; Hinkes et al., 2005; Woolington, 2009a).
The Nushagak Peninsula herd is a small population that was established in 1988, when caribou
from the Northern Alaska Peninsula herd were translocated to the Nushagak Peninsula south of
the Nushagak River delta, on the west side of upper Bristol Bay. The Nushagak Peninsula had
been unoccupied by caribou for approximately 100 years (Hotchkiss, 1989; Paul, 2009). The
Nushagak Peninsula herd grew rapidly after its introduction, from 146 caribou to over 1,000
caribou in 1994. Growth continued at a slower rate to about 1,400 caribou in 1997. Population
density peaked at approximately 1.2 caribou per km . During the next decade, calf recruitment
and adult female survival decreased and the population declined to 546 caribou in 2006
(Aderman, 2009). The population remained at about 550 caribou until 2009 and then increased to
801 by 2011 (Aderman and Lowe, 2011).
Habitat
Seasonal Preference- Spring calving grounds tend to be in open tundra areas or high and rugged
mountains. Predator densities are often lower in such areas, but large caribou herds can also
calve at high densities in sparsely forested terrain, where their sheer numbers and synchronized
timing of births can swamp the effects of predators (ADF&G, 1985; Skoog, 1968).
During summer (mid-June to mid-August), caribou feed in open tundra, mountain, or sparsely
forested areas. To avoid harassment from mosquitoes and other insects, caribou often gather on
windswept ridges, glaciers, lingering snow drifts, gravel bars, elevated terrain, cinder patches,
and beaches. Caribou near the coast may also avoid insects by standing head down and
motionless on mudflats (ADF&G, 1985; Skoog, 1968).
In winter caribou often feed in forested areas, especially where there are spruce-lichen
associations. In addition to forested areas, caribou can also be found along ridge tops, on frozen
lakes and in bogs during winter (ADF&G, 1985; Skoog, 1968).
Food Habits
Spring- From mid-April to mid-June, caribou usually eat catkins of willow (Salix alaxensis, S.
planifolia spp. S. pulchra, and S. glauca), grasses and sedges (Carex bigelowii, C.
membranacea, C. podocarpa, and Eriphorum vaginatuni). They also consume fruticose lichens,
resin birch (Betula glandulosa), dwarf birch (B. nana), and horsetails (Equisetum spp.)(ADF&G,
1985; Skoog, 1968).
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Summer- From mid-June to mid-August, caribou typically consume willow leaves, resin birch,
and dwarf birch, as well as sedges and grasses, especially grasses from the genera Alopecurus,
Arctagrostis, Dupontia, Festuca, Poa, Puccinellia, Calamagrostis, and Hierochloe. They also eat
horsetails, legumes (Astragalus umbellatus, Lupinus arcticus, Hedysarum alpinum, and
Oxytropis nigresens), and forbs such as Gentiana glauca, Swertia perennis, Sedum roseum,
Antennaria monocephala, Artemisia arctica, Epilobium latifolia, Pedicular is spp., Petasites
frigidus, Polygonum bistorta, Rumex arcticus, and Saxifraga spp. (ADF&G, 1985; Skoog, 1968).
Fall- Caribou feed on grasses, sedges, and lichens throughout the fall. They also feed on willow
and water sedge (Carex aquatilis\ if they are available (ADF&G, 1985). Caribou also feed on
mushrooms, when available (Skoog, 1968).
Winter- Caribou winter diets consist primarily of lichens (especially Cladonia spp. and Cetraria
spp.), with smaller amounts of sedges and grasses, as well as horsetails, and the tips and buds of
willows and dwarf shrubs (e.g., Vaccinium uliginosuni). They may consume vegetation in
muskrat pushups during winter, as well as aquatic vegetation in poorly drained coastal plains
(ADF&G, 1985; Skoog, 1968).
Behavior
Seasonal Range Use and Migrations- Some caribou herds travel long distances between
summer and winter ranges, in order to find adequate sources of food and bear their calves in
areas relatively free of predators (Bergerud, 1996; Griffith et al., 2002; Skoog, 1968; Whitten et
al., 1992). Physical features on the landscape influence caribou migration routes. Caribou must
negotiate around open seawater, large lakes, swift rivers, rivers with floating ice, rocky regions
in high mountains, volcanic cinder patches, glaciers, and burns. Features such as frozen lakes and
rivers, as well as ridge tops, eskers, streambeds, and hard-surfaced snow drifts aid caribou during
winter migration (ADF&G, 1985). Since the 1980s, calving areas, other seasonal ranges, and
migration routes of the Mulchatna herd have varied widely. The Mulchatna herd has ranged
extensively throughout most of the Nushagak and Kvichak watersheds, but caribou from this
herd also spend much of their time to the north in the Kuskokwim River drainage (Woolington,
2009a).
In contrast to most other migratory caribou herds, the Mulchatna herd does not use the same
traditional calving ground annually, although its calving areas have remained distinct from those
of any other herds. The Mulchatna herd calved in the Bonanza Hills area of the upper Mulchatna
River watershed during the 1980s. In 1992, calving shifted west to the Mosquito River drainage
in the upper Mulchatna watershed. From 1994 to 1999 calving generally occurred in the upper
Nushagak River watershed. From 2000 to 2002 calving was split between the lower Nushagak
watershed and the South Fork of the Hoholitna River, in the Kuskokwim drainage. In 2003 and
from 2005 to 2008, calving occurred near Kemuk Mountain in the Nushagak drainage, as well as
near the South Fork of the Hoholitna in the Kuskokwim drainage. In 2004 calving was
widespread, from Dillingham in the Nushagak watershed, north to the Holitna and Hoholitna
Rivers in the Kuskokwim drainage (Woolington, 2009a).
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The Mulchatna herd often ranges widely across the Nushagak drainage during summer and fall,
but also frequently uses areas to the north and west, in the Kuskokwim Mountains. During the
1980s much of the Mulchatna herd wintered north and west of Lake Iliamna, in the Kvichak
drainage. In the 1990s most wintering shifted to the Kuskokwim Mountains. For the past decade
part of the herd has wintered in the Nushagak and Mulchatna watersheds while part of the herd
has wintered in the Kuskokwim watershed. In 2006/2007 and 2007/2008, up to 20,000
Mulchatna caribou wintered in the lower Nushagak and Kvichak valleys, with some going as far
south as the Naknek valley in 2006/2007 (Woolington, 2009a).
Mulchatna caribou are often widely dispersed during movements between seasonal ranges. In
accordance with the highly variable locations of seasonal ranges, migration tends not to follow
the same routes from year to year (Woolington, 2009a).
Historically, the Northern Alaska Peninsula herd has spent most of its time in areas from the
Naknek drainage to the south, far removed from the Kvichak and Nushagak watersheds. From
about 1986 through 2000 many caribou from the Northern Alaska Peninsula herd did winter in
the Kvichak drainage, south of Lake Iliamna, but since 2001, only a single radio-collared caribou
from this herd has wintered north of the Naknek River (Butler, 2009a).
Like many small caribou herds, the Nushagak Peninsula herd is sedentary and spends the entire
year on the Nushagak Peninsula, although a few caribou from this herd have made short forays
off the Peninsula (< 20 km and for < 1 month) (Aderman and Woolington, 2001). So far, there
has been no overlap between the Nushagak Peninsula herd and much larger migratory Mulchatna
herd.
Response to Disturbance- Industrial activities impact caribou by hindering or altering their
movements or displacing them from preferred habitats. Barren-ground caribou on the North
Slope of Alaska have avoided development such as exploration wells (Fancy, 1983) and linear
developments such as roads and pipelines (Dau and Cameron, 1986) by distances of 2 to 5 km.
Establishment of extensive, densely packed development with interconnecting road networks,
high levels of traffic, aircraft activity, and ongoing construction or production activity around the
Prudhoe Bay oilfields has resulted in general displacement of caribou from some areas (Griffith
et al., 2002). Avoidance and displacement are most prevalent among females with young calves
(Cameron and Whitten, 1980; Cameron et al., 1979; Griffith et al., 2002). Similarly, woodland
caribou in Canada typically avoided areas near mining sites by 1 to 5 km (Weir et al., 2007).
Mining activities had the highest impact on caribou during calving season. Larger groups and
females with young typically avoided mining sites more often than smaller groups and caribou
without young (Weir et al., 2007). Weir et al. (2007) identified corridors such as roads and
seismic lines as the greatest development impact on caribou because they increase the chance of
encounters with humans and predators. The large Red Dog Mine in northwestern Alaska has had
only limited and localized effects on caribou movements and distribution, in part because the
mine occupies only a tiny fraction of the Western Arctic Caribou herd's otherwise pristine range.
Also, mine operators and workers have implemented policies to minimize conflicts between
traffic and caribou along the road from the mine to the port site (Dau, 2009). In Norway
movement patterns and range use by wild reindeer have been disrupted by combinations of
highways and railroads, as well as by large hydroelectric developments (Nellemann et al., 2001;
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Nellemann et al., 2003). Impacts of any development tend to be less when they occur on non-
critical seasonal range, in areas or at times when caribou are at low density relative to available
range, or when similar habitats are available nearby (Griffith et al., 2002).
Interspecies Interactions
The interrelationships of wolves, caribou, and moose populations have been studied extensively
in Alaska (Gasaway et al., 1983; Mech et al., 1998; National Research Council, 1997). In large
areas of interior Alaska, moose tend to persist for long time periods at low densities, with
population size regulated by high rates of predation by wolves and bears (Gasaway et al., 1992).
In contrast, caribou are able to periodically escape regulation by predators and at least
temporarily achieve high densities (Davis and Valkenburg, 1991; Valkenburg, 2001). Such a
pattern is consistent with caribou population dynamics in the Nushagak and Kvichak watersheds.
Predation by wolves does not appear to be a major factor in regulating the Mulchatna herd,
possibly due to rabies outbreaks that periodically reduce the wolf population (Valkenburg et al.,
2003; Woolington, 2009a). Large migratory caribou herds like the Mulchatna may also avoid
predation by moving seasonally to areas with few resident predators or by erratic and
unpredictable use of seasonal ranges. Wolves in the Nushagak and Kvichak drainages are not
known to follow migratory caribou (Woolington, 2009a).
Mortality, Productivity, Survivorship
Mortality- Caribou populations are influenced by the availability and quality of forage plants,
predation, weather, climate, disease, and hunting (Valkenburg, 2001). Winter severity, accidents,
and insect harassment can also affect caribou numbers (Hinkes et al., 2005). Rapid growth of the
Mulchatna herd from 1980 to 1995 indicated that predation pressure was not a limiting factor on
the herd at that time. During its continued decline from 1997 to the present, this herd has been
strongly limited by nutrition. Poor nutrition has also been associated with high levels of bacterial
pneumonia, hoof rot (Spherophorous necrophorous) and high parasite loads (Valkenburg et al.,
2003; Woolington, 2009a).
Predation by wolves and bears is now limiting calf survival and recruitment in the Northern
Alaska Peninsula herd, but lowered productivity due to nutritional stress is also a problem
(Butler, 2009a). Calf recruitment in the Nushagak Peninsula Herd has been lower, as the herd
declined in recent years, but causes are not well known. Depletion of lichens on winter range
may have contributed to poor nutrition (Valkenburg et al., 2003).
Breeding- Rutting occurs during fall migration and on wintering grounds. Females tend to breed
at 28 months of age, but age at first breeding can vary from 16 to 41 months, depending on
health (Hinkes and Van Daele, 1996). Females in good nutritional condition have a pregnancy
rate of 80% or more, but pregnancy rates may drop dramatically when cows are in poor
condition, due to severe weather effects on grazing, or in some cases, due to overgrazing of
range when caribou are at high densities. Gestation typically takes 225 to 235 days. Calving
occurs in late May or early June and females usually have one calf per year (Skoog, 1968). Birth
rates in the Mulchatna, Northern Alaska Peninsula, and Nushagak Peninsula herds all dropped
after these herds reached peak population levels and then began to decline (Valkenburg et al.,
2003).
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Human Use/Interaction/Management
Nearly all caribou harvested in the Nushagak and Kvichak watersheds are from the Mulchatna
herd. Caribou are an important subsistence food species for people residing in the area served by
the Bristol Bay Area Health Corporation (Ballew et al., 2004). In a survey about traditional food
consumption conducted in 2002, 88% of respondents from that region reported consumption of
caribou meat within the past year (Ballew et al., 2004). Caribou was second only to salmon as a
subsistence source of meat for residents of the region. Caribou are also harvested by non-local
residents, who fly into the area to hunt. Harvest levels for all hunters are highly dependent on
caribou distribution during the fall and winter, as well as weather and snow cover conditions that
affect hunter access to caribou (Woolington, 2009a). Harvest is also generally correlated with
population size, with historically high harvests occurring when caribou have been most
abundant. Reported harvest of Mulchatna herd caribou from 1991 to 1999 ranged from 1,573 to
4,770 (Table 5); although those totals include Mulchatna caribou taken in areas outside the
Nushagak and Kvichak drainages (Woolington, 2009a). Estimates of total harvest from the
Mulchatna herd during this period were roughly twice as high (3,770 to 9,770)(Valkenburg et al.,
2003). However, harvest probably never exceeded 5% of the annual population and did not limit
herd growth or range expansion or cause the decline of the population (Woolington, 2009a). As
the Mulchatna herd declined in numbers after 1999, reported harvest steadily dropped to a low of
767 in 2007/2008 (Table 5). Lower harvests reflect generally reduced availability of caribou
(Woolington, 2009a). Also, long hunting seasons, high bag limits (five caribou), and same-day-
airborne hunting that were allowed during the 1990s and early 2000s have since been replaced
by more restrictive regulations.
5,
Regulatory Year
1991-1992
1992-1993
1993-1994
1994-1995
1995-1996
1996-1997
1997-1998
1998-1999
1999-2000
2000-2001
2001-2002
2002-2003
2003-2004
2004-2005
2005-2006
2006-2007
2007-2008
Estimated Herd Size
90,000
115,000
150,000
180,000
190,000
200,000
N/A
N/A
175,000
N/A
N/A
147,000
N/A
85,000
N/A
45,000
N/A
Reported Harvest
1,573
1,602
2,804
3,301
4,449
2,366
2,704
4,770
4,467
4,096
3,830
2,537
3,182
2,236
2,175
921
767
(Woolington, 2009)
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Hunting for Nushagak Peninsula Herd caribou is managed under regulations set by the Federal
Subsistence Board. From 1995 to 2011, a total of 673 caribou were reported harvested from this
herd. Reported harvests were < 12.3% of the population annually during this period (Aderman
and Lowe, 2011), but there may have been additional unreported harvest (Valkenburg et al.,
2003). Factors other than hunting (e.g., depletion of lichens on winter range) may have been
involved in the decline of the Nushagak Peninsula Herd after 1999 (Valkenburg et al., 2003).
Nevertheless, it was clear that the herd could no longer support the high levels of harvest seen
during the 1990s. Harvest quotas were reduced and the herd is now increasing again.
With the Northern Alaska Peninsula herd now at very low population levels, the herd no longer
extends as far north as the Kvichak drainage. Overall harvest of the herd is greatly restricted, and
none occurs in the_Nushagak and Kvichak watersheds.
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WOLF
Introduction
The gray wolf (Canis lupus) is the largest wild extant canid (Paquet and Carbyn, 2003). The
historic distribution of wolves once covered most of North America, but as the contiguous
United States were settled during the past two and one-half centuries, the wolf was widely
persecuted due to its tendency to prey on livestock and pets (Mech, 1995). By the 1970s, wolf
populations in the contiguous United States were decimated, which led to their protection under
the Endangered Species Act. The gray wolf is currently listed as "endangered" in most of the
Lower 48 states, except in Minnesota, where it is "threatened," in the northern Rocky Mountains
where the species was recently de-listed as "recovered," and for several experimental
populations in Wyoming and the southwest United States (USFWS, 2011).
Habitat
Wolves are habitat generalists and their home ranges can encompass a variety of diverse habitats
(Mech, 1970; Mladenoff et al., 1995; Paquet and Carbyn, 2003). Historically, gray wolves were
distributed throughout the northern hemisphere in every habitat where large ungulates were
found (Mech, 1995). Prey abundance and availability strongly influence habitat use by wolves
(Paquet and Carbyn, 2003). Male and female wolves do not differ in habitat selection, and the
pack maintains their territory throughout the year.
Wolf pups are born, protected, fed, and raised in natal and secondary den sites, a series of
rendezvous sites, and surrounding areas (Paquet and Carbyn, 2003). Dens provide shelter and are
often located in a hole, rock crevice, hollow log, overturned stump, abandoned beaver lodge, or
expanded mammal burrow (Paquet and Carbyn, 2003). Rendezvous sites are areas where pups
are left while pack members forage (Theberge, 1969).
Decades ago, it was commonly thought that wolves needed wilderness to survive. More recent
studies have shown that wolves do not need wilderness, but they do require adequate prey and a
relatively low rate of mortality caused by humans (Mech, 1995; Mladenoff et al., 1999). The
presence of roads has a complex impact on habitat selection by wolves. Roads benefit wolves by
easing their travel and access to prey, but conversely roads are associated with human contact
and increased wolf mortality through either intentional or accidental killing (Houle et al., 2010;
Mladenoff et al., 1999). Near the Kenai NWR in Alaska, wolves preferred a gated pipeline road,
presumably because it offered an easy travel corridor with little human use (Thurber et al., 1994).
In that study, wolf absence from human-settled areas and heavily travelled roads seemed to be
caused by wolf behavioral avoidance rather than direct human-caused mortality of wolves in
those areas.
Food Habits
Diet- Wolves are obligate carnivores whose use of prey depends largely on the availability and
vulnerability of ungulates (Weaver, 1994). Dietary habits such as preferred prey species and prey
switching tactics vary substantially among wolf packs in different locations, in response to local
ecological relationships.
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Wolves can be flexible and shift to non-ungulate prey species when ungulate prey are scarce
(Forbes and Theberge, 1996) or to take advantage of seasonally abundant, nutritious alternate
prey species such as salmon (Darimont et al., 2008). Some wolf packs require dietary
supplementation during the summer, in order to meet the high energetic demands of
reproduction (Paquet and Carbyn, 2003). Dietary supplementation by alternate prey species is
also important for wolf packs in northwestern Alaska that rely on migratory caribou, which move
seasonally to calving grounds inaccessible to wolves (Ballard et al., 1997).
Beavers and snowshoe hare are important to the winter diet of wolves in Algonquin Park in
Ontario, as are scavenged moose carcasses (Forbes and Theberge, 1992; Forbes and Theberge,
1996). Other animals such as lemmings, voles, muskrat, and a variety of birds (especially
waterfowl) and their eggs also supplement the wolf diet (Kuyt et al., 1981), while fish and
berries are consumed seasonally, where available (Darimont and Paquet, 2000; Kohira and
Rexstad, 1995).
Coastal wolves also consume marine mammal carcasses, mussels, crabs, and even barnacles
(Darimont and Paquet, 2000). The Ilnik wolf pack on the Alaska Peninsula was found to
preferentially utilize coastal habitat along Bristol Bay, where it was frequently observed
consuming marine mammal carcasses that had washed ashore (Watts et al., 2010). In the winter,
when Bristol Bay was frozen, the pack was documented using offshore sea ice, and wolves killed
sea otters (Enhydra lutris kenyonf) near the coastline when the otters were trapped above the sea
ice (Watts etal., 2010).
Wolves on the Kenai Peninsula of Alaska were found to rely heavily on moose during the
summer (Peterson et al., 1984). Moose comprised an estimated 97% of ingested prey biomass in
summer, which was largely scavenged from old kills; only 16% of moose carcasses found with
summer wolves were fresh kills. In contrast, 80% of moose consumed during the winter were
fresh wolf-kills (Peterson et al., 1984). Kenai Peninsula wolves also ate snowshoe hare and
beaver during the summer, and minor quantities of small rodents, birds, vegetation and other
prey (Peterson et al., 1984). Scat from wolves in south-central Alaska confirmed reliance on
moose; beaver and snowshoe hare were also commonly consumed (Ballard et al., 1987). Wolves
in south-central Alaska also eat caribou, muskrat, squirrel, voles, vegetation, and a variety of
other dietary items (Ballard et al., 1987).
Salmon as a Food Source2- Preying on salmon may have considerable adaptive value for
wolves regardless of ungulate density. Foraging theory predicts the avoidance of dangerous
ungulate prey in favor of less dangerous alternatives such as salmon (Stephens and Krebs, 1986).
Salmon also offers superior nutritive value; in one study pink salmon contained more than four
times as much energy per 100 g of meat than raw black-tailed deer (Darimont et al., 2008).
Behavioral observations suggest that wolves may have a broad history of seasonal consumption
of salmon in areas where the two species co-exist (Darimont et al., 2003).
2 Ongoing research in LCNPP and the Alaska Peninsula and Becharof NWRs is providing new information on the
relationship of wolves to salmon. The results of this research have not yet been analyzed or published, but
preliminary results show that wolves rely on salmon, when available, for a significant portion of their diets. This
information is cited as a personal communication in this section.
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Wolves have often been observed consuming only the head of salmon instead of the whole fish
(Darimont et al., 2003). There are several possible explanations for this behavior. Wolves may be
consuming only the most energetically valuable part of the prey item, or they may be targeting
specific micronutrients such as omega-3 fatty acids (Gende et al., 2001). Wolves may also be
selecting head tissue to minimize their exposure to parasites such as Neorickettsia helminthoeca,
which can infect salmon and can be fatal to canids (Darimont et al., 2003).
Wolf packs that seasonally utilize salmon can reap benefits during the fall and winter seasons.
The consumption of salmon in the fall may improve pup survivorship during weaning (Person,
2001). Winter snow can preserve salmon carcasses buried underneath, enabling use by wolves
and other scavengers for the rest of the winter (Carnes 2004). Carnes (2004) compared scat
across different packs in the Copper and Bering River deltas, and noted that wolf consumption of
salmon increased in winter relative to other seasons; he hypothesized this might be related to the
relative lack of seasonal availability of moose in those areas.
Several studies in Alaska have examined the importance of salmon to wolves from various parts
of the State. In the Copper and Bering River Deltas, late summer rendezvous sites for wolves
were typically located alongside shallow spots in spawning areas or at bends where gravel bars
extended out into streams (Carnes 2004). Researchers observed wolves, especially pups at these
spots, waiting for spent salmon carcasses to float by (Carnes 2004). In southeast Alaska marine
protein composed 18% of the lifetime total diet of Alexander Archipelago wolves; most of the
marine contribution was likely salmon, but other marine organisms were probably also
consumed (Szepanski et al., 1999). In southwest Alaska's Togiak NWR, wolves have been
observed delivering intact salmon carcasses to their pups at rendezvous sites (Walsh, 2011).
Similar foraging behaviors have been observed among wolves on the Alaska Peninsula (GMUs
9C and 9E, extending from the Naknek River drainage through Port Moller), where wolves often
transport captured salmon to den or rendezvous sites (Watts, personal communication).
Salmon are not solely a food resource for coastal wolves. Some Pacific salmon migrate long
distances inland, returning to spawning grounds that may be hundreds of miles from the ocean
(Quinn, 2004). A study within DNPP, in Interior Alaska, documented substantial seasonal
salmon consumption among wolves who lived more than 1,200 river km from the coast (Adams
et al., 2010). Wolves with ranges in areas where salmon were seasonally abundant and ungulates
occurred at low densities ate the most salmon; salmon averaged 17% of their total long-term diet
(Adams et al., 2010). Preliminary data from LCNPP indicates that wolves use salmon from the
time the fish enter streams, through the fall, and then again after late-winter ice out (Mangipane,
personal communication).
Dispersal of Marine-Derived Nutrients (MDNs) by Wolves- The influences of salmon on
terrestrial systems are largely dependent on predators that remove salmon from streams,
consume a portion, and leave the remains behind (Hilderbrand et al., 1999a; Reimchen, 2000).
Abandoned salmon carcasses contribute to ecosystem processes, as scavenging, decomposition,
and fecal-urinary deposition provide MDNs to terrestrial systems that are typically nitrogen- and
phosphorus-limited (Ben-David et al., 1998; Hilderbrand et al., 1999a; Reimchen, 2000; Willson
etal., 1998).
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Wolf behavior further influences the pattern of distribution of MDNs to terrestrial ecosystems, as
wolves often transport caught salmon some distance rather than consuming it in the stream or
immediate vicinity. In British Columbia wolves were observed to consume caught salmon on
grass next to the river 70% of the time (Darimont et al., 2003). However, in LCNPP, preliminary
data indicate that wolves move considerable distances over several days to feed on salmon. In
2009, in LCNPP, an individual wolf was documented travelling up to 64 km from a den site to
feed on salmon and carry ingested remains back to feed pups. In 2010 and 2011, the same
individual travelled up to 24 km and 40 km to feed on salmon (Mangipane, personal
communication). In some locations, wolves consume only the salmon head, leaving the
remainder of the carcass behind (Darimont et al., 2003), whereas bears often consume eggs,
muscle and other body parts of salmon, especially when salmon density is not high (Gende et al.,
2001). In LCNPP wolves have been observed feeding on fish carcasses frozen into lake ice, and
the backbones and heads left from human subsistence fishing (Mangipane, personal
communication; Spencer, personal communication).
Behavior
Wolf Packs- Gray wolves are territorial and social carnivores that typically live in packs of about
six to eight animals but packs may include >20 wolves (Mech and Boitani, 2003). Wolf packs
typically consist of a single breeding pair, pups of the year, and their older siblings (Mech and
Boitani, 2003). Mating occurs during late January to March and gestation is usually 63 days. In
each pack, a single litter of pups is born in a den during late April to May. Nonetheless, multiple
litters have been observed within some packs occurring in Alaska (Ballard et al., 1987; Meier et
al., 1995). Litter sizes range from one to 12 pups but usually four to six pups are born (Fuller et
al., 2003). Dens in coastal temperate rainforests are located within the root wads of living or
dead trees (Person and Russell, 2009). In boreal forest or tundra, dens are located in sandy areas
or gravel eskers (Ballard and Dau, 1983; McLoughlin et al., 2004). Wolves and their pups
occupy dens between late April and early July, and then move to rendezvous sites where
sequestered pups are fed by pack members until September or early October when they are
sufficiently large to move with the pack (Mech et al., 1998; Packard, 2003; Person and Russell,
2009). Pup mortality during summer is affected strongly by availability of food (Fuller et al.,
2003). Wolves usually remain within their natal packs until they reach sexual maturity at 22 to
24 months. At that age, some may disperse from their packs to find mates and establish their own
packs. However, researchers reported dispersers ranging in age from 10 months to 5 years (Mech
and Boitani, 2003). Abundant prey may induce some wolves to defer dispersal until they are
older and thus packs may grow to large size (Fuller et al., 2003; Mech and Boitani, 2003).
Dispersing wolves may travel hundreds of kilometers and traverse very difficult terrain before
settling (Mech and Boitani, 2003) and they are able to cross large bodies of water. For example,
in southeastern Alaska, dispersing wolves were documented swimming 3 to 4 km in open ocean
to move between islands (Person and Russell, 2008).
Range- Resident wolf packs occupy extensive territories that they attempt to defend from other
wolves. Wolf territories tend to be smaller in summer, when packs remain closer to dens and
home sites (Mech, 1977), and are larger in winter, when the pack resumes nomadic travelling as
pups mature. In south-central Alaska, the average distance between dens of neighboring packs
was 45 km (Ballard et al., 1987).
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Territories of wolf packs tend to be much larger in Alaska than in the remainder of the United
States, due to the relatively low density of prey in Alaska. Home ranges and pack sizes largely
are influenced by availability of prey (Mech and Boitani, 2003). For example, when prey is
abundant, home ranges tend to be small and pack sizes large and the opposite is true when prey
are scarce. In Alaska, ungulates such as moose, caribou, and Sitka black-tailed deer (Odocoileus
hemionus sitkensis) are the most important prey (Gasaway et al., 1992; Kohira and Rexstad,
1997). In general, wolf territory size is inversely related to the density of available prey (Fuller et
al., 2003). Average territory sizes of wolf packs from different regions of Alaska were
consistently larger in winter than in summer (Table 6) (Adams et al., 2008; Ballard et al., 1997;
Ballard et al., 1998; Ballard et al., 1987; Burch et al., 2005; Peterson et al., 1984). Each of these
studies of Alaska wolf packs documented territory sizes much larger than those of wolves from
north-central Minnesota, where small winter territory size (average 116 km2) and high wolf
density (39/1,000 km2 in mid-winter) were attributed to an abundant white-tailed deer population
(Fuller, 1989). Preliminary analysis of recent data from LCNPP has shown annual ranges from
1,155 km2 to over 5,000 km2. One pack in this study had ranges of 2,214 km2, 2,189 km2, and
1,834 km2 from 2009 to 2011 (Mangipane, personal communication).
6, of In
Region
Northwest
Denali National Park
Central Brooks Range
Kenai Peninsula
South-central
Summer Winter Annual
621 1,372 1,868
871
358-2,3 15a
466-864b
1,644
Reference
Ballard etal., 1997
Burch et al., 2005
Adams etal., 2008
Peterson etal., 1984
Ballard etal., 1987
a - Range of territory sizes estimated for wolf packs over the four-yr study period
b - Range of average annual territory sizes during study period
~ not determined
* This table does not include data from an ongoing study in LCNPP.
Dispersal (Emigration)- Several studies in Alaska have documented emigration as a vital factor
influencing the population dynamics of wolves (Adams et al., 2008; Ballard et al., 1997; Ballard
et al., 1987; Peterson et al., 1984). Individuals may leave a pack and strike out on their own in
response to low prey densities (Messier, 1985). High rates of infectious disease (Ballard et al.,
1997), social stress within the pack, or a lack of opportunity to achieve the high social status
needed to successfully breed (Peterson et al., 1984) may also cause wolves to disperse to new
territories.
Dispersal is a key mechanism that wolves use to colonize new habitats that become available.
Dispersing wolves experience a high rate of mortality, but when successful, they are able to
establish a new pack (Peterson et al., 1984) or join an existing pack (Mangipane, personal
communication). Successful colonization of new territory requires both a vacancy of suitable
habitat and bonding with a mate (Rothman and Mech, 1979).
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Seasonal Movements- Wolves in south-central Alaska do not follow migratory movements of
moose or caribou outside their pack areas, but do follow elevation movements of moose within
their pack areas (Ballard et al., 1987). In a study in northwest Alaska, wolf packs usually did not
follow migratory caribou from the Western Arctic herd, but rather, switched to moose for prey
during the winter and maintained year-round resident territories (Ballard et al., 1997). However,
in years when moose densities were low, up to 17% of radio-collared wolf packs in northwest
Alaska followed migratory caribou and then returned to their original territory for denning
(Ballard et al., 1997).
In Bristol Bay, there is no evidence that wolf packs follow the Mulchatna caribou herd, although
wolves are occasionally seen with the herd as it moves throughout the region (Woolington,
2009b). However, recent information in LCNPP shows that wolves (in most cases, lone yearling
wolves) were following caribou herds for all or a portion of the year (Mangipane, personal
communication). Packs are more likely to have established territories and take advantage of
caribou when they move through those territories.
Daily distances traveled within a pack's territory range from a few kilometers up to 200 km
(Mech, 1970). On Ellesmere Island, Northwest Territories, mean travel speed of wolves during
summer, on barren ground, was 8.7 km/hr for regular travel and 10.0 km/hr when returning to a
den (Mech, 1994). In south-central Alaska, a wolf pack followed for 15 days in the spring moved
an average of 24 km per day (Burkholder, 1959). In LCNPP, all packs and age classes of wolves
have been documented travelling up to 34 km in 15 hours (Mangipane, personal
communication).
Wolves in general are good swimmers; coastal wolves are particularly adept at swimming and
are able to swim distances as far as 13 km between islands (Darimont and Paquet, 2002).
However, wolves may be unwilling to swim in pursuit of large ungulates. On the Kenai
Peninsula, wolves ceased pursuit of moose that entered ponds or lakes and swam away from
shore (Peterson et al., 1984). However, as soon as waterbodies freeze, wolves travel across them
freely (Spencer, personal communication).
Interspecies Interactions; Response to Change in Salmon Populations/
Distribution
In coastal regions and along major river systems salmon are important seasonal prey for wolves
(Adams et al., 2010; Kohira and Rexstad, 1997). Indeed, in some coastal areas, salmon may
seasonally decouple the dependence of wolves on ungulate prey (Darimont et al., 2008). Salmon
is a particularly important food for Alaskan wolf packs in some areas and changes in salmon
abundance may have effects on the alternate prey species of these wolves. In DNPP, salmon
were found to be a particularly important food item for wolves in areas with low ungulate density
but high salmon abundance (Adams et al., 2010). The availability of salmon had a strong impact
on the numerical abundance of wolves in the northwestern flats area of DNPP; wolves were only
17% less abundant in that area compared to the rest of the study area, even as ungulate densities
were 78% lower. The higher wolf population density facilitated by the availability of salmon was
thought to result in increased overall predation pressure on ungulates in that system (Adams et
al., 2010). Moose were the predominant ungulate in the northwestern flats, occurred at densities
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approaching the lowest in North America (Gasaway et al., 1992), and appeared to be limited by
predation rather than nutritional constraint (Adams et al., 2010).
Some wolves in Alaska eat salmon carcasses throughout the winter, as cold winter temperatures
and snowfall effectively preserve this food resource. In one study of wolf scat, salmon
consumption was observed to increase in the winter relative to other seasons (Carnes, 2004). A
shortage of available salmon might result in a reduced winter food supply for such wolves, which
could lead to either increased predation pressure on alternate prey species or reduced wolf
survival if alternate foods were not available.
In Alaska, ungulates such as moose, caribou, and Sitka black-tailed deer are the most important
prey (Gasaway et al., 1992; Kohira and Rexstad, 1997). Under some circumstances, predation by
wolves may limit or regulate ungulate populations, sometimes suppressing their numbers at very
low densities (top-down forcing) (Gasaway et al., 1992). In other cases, ungulate populations are
influenced mostly by carrying capacity of the range regardless of wolf predation (bottom-up
effects) (Ballard et al., 2001). Nonetheless, the relative effects of top-down and bottom-up
factors can shift over time depending on habitat changes, weather conditions, and
anthropomorphic disturbances (Bowyer et al., 2005).
The interrelationships of wolves, caribou and moose populations have been characterized in
several ecosystems. Alterations in moose densities can have a major influence on caribou
populations, through their effect on wolf predation rates. In southeastern British Columbia, wolf
population numbers can be suppressed due to a lack of available food in winter, when moose
numbers are low, particularly when caribou over-winter in areas inaccessible to wolves (Seip,
1992). Elevated moose densities may cause a concomitant rise in wolf numbers. If moose
numbers later decline, wolves in the area will turn to caribou as an alternative food source,
potentially causing a profound effect on the caribou population. Industrial development can
exacerbate this effect by increasing wolves' access to caribou, via the creation of new linear
corridors, such as roads and pipelines (James et al., 2004). These interrelationships may not be
applicable to the southwest Alaska ecosystem, which has barren-land caribou herds, in contrast
to the low-density woodland caribou herds from the Canadian studies. In LCNPP, when the
Mulchatna caribou herd was at high numbers, wolves fed on caribou and moose numbers were
high. When the herd size declined, wolves fed more on moose and moose numbers declined by
50 percent (Mangipane, personal communication).
Wolves are coursing predators that actively pursue prey rather than passively ambushing them
(Mech, 1970; Mech et al., 1998). Consequently, in much of Alaska, they typically select open or
sparsely forested habitats that enable detection and pursuit of prey. Deep snow that hinders
movement of ungulate prey or restricts them to small, forested patches often facilitates predation
by wolves (Mech and Peterson, 2003). For large ungulate prey such as moose, wolves often
focus predation on calves, which tend to be the most vulnerable. In interior Alaska, wolves are
most effective hunting in flat or rolling terrain covered with sparse boreal forest or tundra. In
coastal rainforests, prey tend to be most vulnerable to predation by wolves in open muskeg
heaths at low elevations (Farmer et al., 2006).
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Mortality, Productivity, and Survivorship
Annual mortality in unexploited wolf populations in Alaska and Yukon ranges from 16 to 27%
and is linked strongly with abundance of prey (Fuller et al., 2003). Mortality results from
accidents, disease, and intra- and inter-pack strife. Where wolves are hunted and trapped, human
exploitation often is the overwhelming source of mortality (Fuller et al., 2003; Person and
Russell, 2008). In a heavily exploited population in south-central Alaska average annual
mortality was 45% of which most (36%) was human caused (Ballard et al., 1987). Human-
caused mortality can be compensatory with respect to other sources of natural mortality (Fuller et
al., 2003). Resident pack members usually have higher survival than nonresident dispersing
wolves because dispersers may be traveling through unfamiliar or unsuitable terrain, and are
subject to attacks by resident wolves (Fuller et al., 2003).
Wolves can live for up to 13 years in the wild (Mech, 1988), typically dying as a result of
starvation, accidents (Mech, 1977), intra-specific fights (Ballard et al., 1987), disease (Ballard et
al., 1997; Woolington, 2009b), or human-related causes (Paquet and Carbyn, 2003; Woolington,
2009b). Starvation and disease are often co-occurring, but the nature of that relationship has not
been fully established (Paquet and Carbyn, 2003). When wolves attempt to take down large prey,
such as moose or caribou, they risk injury or death (Mech, 1970; Paquet and Carbyn, 2003).
Human-related causes of wolf mortality include legal hunting for sport, subsistence or predation
control (Ballard et al., 1997; Ballard et al., 1987; Woolington, 2009b).
Wolves have prolific reproductive potential (Ballard et al., 1987; Boertje and Stephenson, 1992),
and pups generally experience high survival rates through their first autumn (Adams et al., 2008;
Ballard et al., 1987). These pulsed increases in pack size each year must be compensated for by a
combination of mortality and emigration, if the population size is to remain roughly constant
over time (Adams et al., 2008). The relative contribution of emigration, natural mortality, and
human-caused mortality in wolf packs has varied substantially in different parts of Alaska.
In the central Brooks Range in northern Alaska, during the period 1987 to 1991, the resident
wolf population increased by 5% per year while experiencing a 12% annual harvest rate (Adams
et al., 2008). Harvest and natural causes were each responsible for half the annual mortality in
radio-collared wolves. Causes of natural deaths in those wolves, when distinguishable, were
wolves killing other wolves (n=6), avalanche (n=l), and old age (n=l) (Adams et al., 2008).
Pups constituted about half the wolf population each autumn, and young wolves emigrated from
the study area at high rates as yearlings (47%) and two-year-olds (27%).
In northwest Alaska, during the period 1987 to 1992, the annual survival rate averaged 55.2% for
radio-collared wolves (Ballard et al., 1997). Hunting was responsible for 69% of known
mortalities. Rabies was also a significant cause of death in this population (21% of mortalities)
during the period 1989 to 1991. Twenty-one wolves (25%) dispersed from their original territory
during the study, with the highest rates of dispersal occurring during the rabies outbreak.
In south-central Alaska, in GMU 13 (the Nelchina and upper Susitna basins), wolf population
levels were highest at the beginning of one study period (1975) and declined each year through
the end of the study (1982), due to aircraft-assisted ground shooting and state-managed wolf
control (Ballard et al., 1987). Litter sizes in this wolf population ranged from two to nine pups,
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with an average of six pups. Natural mortality accounted for only 20% of total wolf mortality in
that study. That wolf population could sustain a mortality rate of 50% from all sources before
experiencing a population decline. Human harvests in excess of 40% of autumn wolf numbers
caused population declines. Wolf control practices were effective at reducing wolf numbers, but
when wolf control ceased the wolf population rebounded quickly.
Wolves re-established on the Kenai Peninsula by natural immigration in the 1960s, after being
absent since the early 1900s. That wolf population was studied during the period 1976 to 1981,
and population changes were documented (Peterson et al., 1984). Wolf density in the winter of
1976/1977 was 11 wolves/1,000 km2; density increased to 16 in 1978/1979, dropped to 11 to 12
in 1980/1981, and increased back to 18 to 19 in 1981/1982. Annual survival rate for radio-
collared wolves declined during each year of the study, from 100% in 1976/1977 to 44% in
1980/1981. Harvest was responsible for the majority of mortality. Reported harvest averaged
30% annually, and annual mortality of radio-collared wolves was 32%. Dispersal was found to
play a key role in wolf population dynamics during this study. Dispersing wolves were highly
vulnerable to harvest (the annual survival rate for dispersing adults was only 38%, compared to
73% for resident adults), but those dispersers who survived to reproduce were critical to the
maintenance of population densities in spite of increased mortality rates (Peterson et al., 1984).
In LCNPP 17 wolves were radio collared in a three-year period (2009 to 2011). Of the 17 wolves
collared, eight died during that time. Three were harvested, two died in intra-specific fights, one
drowned, and two died from unknown causes. Annual survival for all age classes for each year of
the study was 75%, 63%, and 75%. Of five dispersing wolves in LCNPP, survival was estimated
at 60%, but this is likely a high estimate (Mangipane, personal communication).
Population Estimates
There are between 7,000 and 11,000 gray wolves in Alaska. The highest densities occur on the
islands associated with the southeastern panhandle where deer (are the principle prey and the
lowest densities occur in mountainous areas where prey consists mostly of Dall sheep or
mountain goats (Fuller et al., 2003; Person et al., 1996). Two genetically distinct wolf
populations are recognized within Alaska, those occupying the coastal zone of southeastern
Alaska and those inhabiting the rest of the state (Weckworth et al., 2005). During 2000 to 2009,
1,200 to 1,600 wolves, about 14 to 16% of the estimated population, were reported harvested
annually in Alaska. It is estimated that wolf populations can usually sustain 30 to 40% total
annual mortality (Fuller et al., 2003).
Wolf population numbers have not been well studied in the Nushagak and Kvichak watersheds.
Better regional wolf population estimates, gathered using scientifically rigorous methods, are
needed to improve understanding of wolf populations in the study area. Wolf populations are
highly dynamic, so population estimates must be conducted on a relatively frequent basis.
Additionally, wolf population numbers in this area are difficult to obtain due to vegetation and
inconsistent snow conditions. Dense vegetation and sparse or inconsistent snow make sighting of
wolves from the air challenging.
No population estimation surveys for wolves have been conducted in the Nushagak watershed
(GMUs 17B and 17C; Figure 3) (Woolington, 2009b). ADF&G impressions of wolf population
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status in GMU 17 (the Nushagak and Togiak watersheds, west to Cape Newenham) are based on
observations of wolves and tracks, reports from the public, bounty records from 1962 through
1971, mandatory sealing records from 1972 to present, and an annual trapper questionnaire
program initiated in 1988 (Woolington, 2009b). Based on these data, ADF&G biologists
conclude that wolf density in GMU 17 peaked from 1974 to 1977 and then declined sharply by
1980. Wolf densities seemed to increase again until 1989, when a rabies outbreak affected canid
populations in GMU 17. Wolf populations began to increase again in 1992, and wolves are now
thought to be "abundant" throughout GMU 17 (Woolington, 2009b). Woolington (2009)
provided current wolf population estimates for GMU 17, but these were considered too
speculative to rely on for this assessment and are not repeated here.
The Kvichak watershed, the other large drainage in the study area, is located in GMU 9, which
extends from LCNPP to False Pass. Wolf population estimates for the region are available, but
they should be used with caution for several reasons. ADF&G has grouped GMUs 9 and 10 (the
Aleutian Islands) for statistical purposes, so those wolf population estimates include lands
outside the study area. Also, wolf population dynamics have been studied only lightly in the
region, and only limited descriptions of methods and results are available (Butler, 2009b).
Methods consisted of monitoring ten wolf packs using radio-collar tracking, monitoring trends
through observations during other fieldwork, reviewing reports from hunters and guides, and
collecting responses to annual trapper questionnaires. Using these data, ADF&G estimated a
total population of 350 to 550 wolves in GMUs 9 and 10 (Butler, 2009b). Biologists concluded
that wolf densities in GMU 9 and 10 are low to moderate, but wolf numbers in GMU 9 appear to
have increased since the 1990s, despite a decline in caribou populations. Possible explanations
hypothesized for this increase in wolves included an abundance of alternate prey such as marine
mammal carcasses, salmon or snowshoe hares, a population rebound following a high period of
mortality from a rabies outbreak, or wolf immigration from surrounding areas (Butler, 2009b).
Data are not available to directly evaluate these hypotheses. Estimated wolf densities in GMU 9E
(the Alaska Peninsula south to Port Moller) and the southwestern portion of GMU 9C (the
Naknek watershed outside Katmai NPP) are 6 to 7 wolves per 1000 km (Watts, personal
communication).
Human Use/Interaction/Management
Reporting of wolf harvest in Alaska is mandatory, but reporting compliance is suspected to be
weak in some areas (Ballard et al., 1997). The degree of reporting compliance within the
Nushagak and Kvichak watersheds is unknown. The reported wolf harvest in GMUs 9, 10 and
17 for the period 2003 to 2008 are summarized in Table 7.
Table ?. Total In GMUs 9,10 17 to of
Game.
Year
2003-04
2004-05
2005-06
2006-07
2007-08
GMU 9 and 10a
119
64
120
85
110
GMU 17b
141
60
62
79
73
a= Butler 2009; b=Woolington 2009
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Alaska harvests of wolves vary widely due to fur prices, hunter access to wolf habitat, predator
control policies and practices, and population changes in response to prey populations. Hunter
access is influenced by winter travel conditions (Woolington, 2009b), including snow depth and
fuel prices. Wolves in the Bristol Bay area are typically hunted and trapped by local residents,
but are also harvested opportunistically by non-local hunters.
Trappers from southwest Alaska indicated that wolf was the fourth most important species they
targeted (as defined by the trappers themselves), behind otter, beaver and fox (in that order)
(ADF&G, 2010). State trapping regulations do not distinguish between different types of use,
such as "subsistence," "recreational," or "commercial" (ADF&G, 201 Ib). Most rural Alaska
communities are supported by a mixed subsistence-cash economy (Wolfe, 1991). Trapping is
one of many traditional subsistence activities that can provide a modest income for participants.
Some harvested furs are sold to dealers, but others are used locally. Furs are often made into
hand-crafted items, which are more valuable than the raw pelts (Wolfe, 1991). Items commonly
crafted with furs include mitts, coats, boots, fur ruffs, and slippers. In some rural areas,
households use most of their harvested wolf pelts locally for ruffs, hats, and lining for winter
gear, because imported materials are considered inferior (Wolfe, 1991).
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WATERFOWL
Introduction
The purpose of this report is to provide a characterization of waterfowl resources (Anatidae) in
the Nushagak and Kvichak watersheds. The waterfowl family includes swans, geese, and ducks
(dabbling, diving, and sea). This overview is not a comprehensive account of the status, ecology,
and life history of all species of waterfowl that regularly occur in the Bristol Bay region. Instead,
this section briefly summarizes the prominent species, general habitat associations and their
seasonal occurrence by subregion; highlights some primary ecological relationships between
waterfowl and sources of nutrients, particularly salmon, that support habitats and food resources;
and describes human values and uses of waterfowl from Bristol Bay. Waterfowl data and other
biological information are from currently available sources.
Waterfowl information in this report has been organized by geographic subregions (Estuaries and
Inner Bay, Lowlands and Inland Tundra/Taiga) because of substantially different species
composition, seasonal use patterns, ecological settings, and extent of available biological
information. To a large extent, information in this report is constrained to the Nushagak and
Kvichak watersheds, a subset of the greater Bristol Bay region that has been interpreted as
widely as the area from Cape Pierce to the end of the Alaska Peninsula.
Regional Overview-
The Bristol Bay region hosts 34 regularly occurring species of waterfowl (Appendix 2). The
diverse wetlands and other aquatic habitats of the region include boreal forest and taiga lakes and
ponds inland near Lake Clark, river basin wetlands and lakes along the Mulchatna, Nushagak
and Kvichak valleys, tundra ponds and lakes of the lowlands, and coastal tide flats and estuaries.
The diversity and extensiveness of these habitats provide habitat for many species of waterfowl
as breeding birds, migrants during the summer molt, fall and spring migrants, and wintering
birds.
Geographically, the Bristol Bay region is positioned as a major northern spring staging area for
waterfowl destined to breed in western and northern Alaska, Russia, and Arctic Canada. At the
southern extent of Bering Sea ice, the rich estuaries of Bristol Bay provide food and resting areas
for migrants that are advancing north in spring (King, 1982). Spring aggregations include swans,
geese and ducks arriving from Mexico and the western U.S.; sea ducks from the Pacific coast of
North America (Baja Mexico, British Columbia, southeast and Gulf Coast of Alaska); and
emperor geese (Chen canagica) and sea ducks from the Aleutian Islands.
During summer, the estuaries of Bristol Bay and Kuskokwim Bay to the north serve as
traditional molting areas for large numbers of scoters that gather from the Bering Sea and
western Arctic regions. These shallows provide food-rich and secure habitats at a time when
these birds are nutritionally stressed and flightless. Molting occurs from July through September,
varying among subadults and adults, males and females.
Bristol Bay is an important fall staging area for waterfowl migrating south from northern and
local breeding areas. Fall migration tends to be faster and more direct to staging areas than spring
migration. From mid-August through early October, ducks, Canada (Branta canademis) and
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greater white-fronted (Anser albifrons) geese move overland through the passes of the Alaska
Range to Cook Inlet coastal marshes (Redoubt Bay, Trading Bay, Susitna Flats, and Palmer Hay
Flats). Large numbers of ducks and geese orient to the rich lagoons and coastal tundra on the
north side of the Alaska Peninsula west to Izembek Lagoon. Canada, cackling (Branta
hutchimii) and white-fronted geese; brant (Branta bernicla); and most dabbling ducks depart by
early November, either eastward along Alaska's south coast or directly across the Gulf of Alaska
to points between British Columbia and Mexico. Most dabbling and diving ducks from western
Alaska orient to wintering areas in the Pacific Flyway west of the Rocky Mountains, but some
greater scaup (Aythya marila) and other ducks cross the continent to the Gulf of Mexico and
Atlantic Coast (King, 1973; King and Lensink, 1971). Some sea ducks, an increasing number of
Pacific brant (Ward et al., 2009), and even a small population of tundra swans (Cygnus
columbianus) winter along the Alaska Peninsula, while others and emperor geese move into the
Aleutian Islands.
History of Waterfowl Surveys-
Much of the quantitative data on waterfowl numbers and distribution in the greater Bristol Bay
region are derived from surveys conducted over different areas. In many respects, the
distributions and habitat use patterns of waterfowl in the Nushagak and Kvichak watersheds
intergrade with habitats west of the Nushagak Peninsula and down the Alaska Peninsula to
Izembek Lagoon. Information summarized in this report focuses, as much as possible, on the
inner bay, lowlands, and inland subregions of the Nushagak and Kvichak watersheds.
Osgood (1904) provides a broad and detailed description of habitats and wildlife of the Bristol
Bay region from his 1902 reconnaissance survey, mostly by canoe, starting from Cook Inlet and
travelling to Iliamna Lake and Lake Clark, then up the Chulitna River and down the Koktuli,
Mulchatna, and Nushagak Rivers to Bristol Bay. His trip continued by schooner to Egegik, over
the Alaska Peninsula at Becharof Lake, and then by rowboat to Cold Bay. Hurley (1931; 1932
records bird observations in the Bristol Bay region. Murie (1959) summarized the environment,
habitats and wildlife of the Aleutian Islands and Alaska Peninsula, with some coverage of
Kvichak and Nushagak Bays, from his 1936-37 boat-based expedition. Both Osgood and Murie
provide species accounts of waterfowl, reviewing records of earlier observers. Gabrielson (1944)
compiled general records of birds on his extensive trip in summer 1940, including travel up the
Kvichak River, across Iliamna Lake and portage to Cook Inlet. Hine (1919) and Cahalane (1944)
described birds of the nearby Katmai region. Gabrielson and Lincoln (1959) provided the most
thorough compilation of bird records of their day in The Birds of Alaska., including information
on migration patterns, ecological zones and detailed species accounts. Gill et al. (1981) describe
the waterfowl and other birds of the north-central Alaska Peninsula.
Quantitative surveys of Bristol Bay waterfowl were established in the 1950s to monitor ducks
and geese on prime lowland nesting habitats as part of the annual North American breeding
population survey (Hodges et al., 1996). Interest in oil and gas exploration and other resource
development stimulated more extensive surveys of waterfowl in coastal areas (Bartonek and
Gibson, 1972; King and McKnight, 1969). The Coastal Zone Management Act of 1972 provided
funding to states to synthesize information and establish cooperative wildlife resource
inventories in coastal areas (Timm, 1977). In 1974, the Outer Continental Shelf Environmental
Assessment Program (OCSEAP) stimulated many surveys and research projects related to
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waterfowl in the Gulf of Alaska, Bering Sea, and Beaufort Sea. This program greatly expanded
the amount of information on waterfowl and other birds in Bristol Bay (Arneson, 1980).
Information on waterfowl and other wildlife resources in the region have been reviewed broadly
by Timm (1977); USFWS (1976); and USFWS (1983).
The longest-term and most consistent waterfowl surveys in the Bristol Bay region are part of the
Alaska-Yukon Waterfowl Breeding Population Survey (AYWBPS), flown annually by USFWS
since 1957 as part of a North American duck survey program (Mallek and Groves, 2010). This
survey was designed to index breeding dabbling ducks during the early egg-laying period (late
May) to provide annual status data and long term trends. Bristol Bay (Stratum 8 in this statewide
survey) is composed of 11 transects in 23 segments, sampling an area of 25,641 km2 (9,900 mi2)
usually flown in late May.
From 1989 through 1997, USFWS conducted a series of experimental expanded surveys of
major tundra waterfowl areas to assess means of providing more reliable annual estimates of
abundance (Conant et al., 2007). The expanded waterfowl surveys flown by Platte and Butler
(1995) over the Bristol Bay region in 1993 and 1994 covered a much broader area (49,890 km2)
than the traditional AYWBPS, extending west to Togiak Bay and Togiak River drainage,
covering more a northerly band from Wood-Tikchik Lakes eastward to Port Alsworth, and
southwest on the Alaska Peninsula to include Port Heiden and the Seal Islands. The data from
this experimental survey reflect a much wider range of habitats and duck densities than the
traditional AYWBPS survey (Conant et al., 2007).
Coastline and estuarine waterfowl surveys of Bristol Bay, focusing on emperor geese, have been
conducted annually in spring (1981-2011) (Dau and Mallek, 2011) and fall (1980-2011) (Mallek
and Dau, 2011). Steller's eider are the focus of Bristol Bay surveys conducted since 1992
(Larned and Bellinger, 2011).
Waterfowl Resources and Seasonal Occurrence
Estuaries and Inner Bristol Bay-
Bristol Bay estuaries and nearshore waters are important to waterfowl year round, during spring
and fall migration, summer molting, and as winter range for some species. King and McKnight
(1969) made a first attempt to estimate the number of birds in Bristol Bay during October, flying
transects from the high tide line offshore to 12 miles (19.3 km). Their survey covered over
20,700 km2 of coast from Cape Constantine south and west to Unimak Island, and included lines
in outer Kvichak and Nushagak Bays.
During the early 1970s, Bristol Bay became a focal area of the OCSEAP studies, including
offshore transect surveys of birds from Kvichak Bay south along the Alaska Peninsula (Bartonek
and Gibson, 1972; USFWS, 1976). Most of the survey coverage was over the outer bay beyond
the BBWA area, but it provides insights to the seasonal use of Bristol Bay nearshore waters.
Additional coastal bird surveys were flown for OCSEAP between October 1975 and August
1978 (Arneson, 1980). These 33 surveys from the Gulf of Alaska through the Aleutian Islands
were designed to assess seasonal bird densities and distributions in littoral/nearshore waters,
describe coastal habitats, and document migration. The North-Bristol Bay region was surveyed
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in spring (May) by fixed-wing aircraft, helicopter, and boats. Sections 1-6 of the study area
covered the coast of Kvichak Bay to Cape Constantine within the BBWA study area.
Habitats- The estuaries and nearshore waters of Bristol Bay provide diverse aquatic habitats for
waterfowl and other waterbirds (Michel et al., 1982; Selkregg, 1976). Arneson (1980) classified
over 30 types of coastal habitats, ranging from intertidal flats and salt marshes to open waters, to
document seasonal usage by birds. He found about 80% of waterbirds on protected delta habitats
and exposed inshore waters. Dabbling ducks and geese preferred mudflats and delta habitats;
diving ducks and sea ducks were found mostly in exposed inshore and bay waters. A large
proportion of Nushagak and Kvichak Bays has water depths of < 10 m (Schamber et al., 2010),
which provides very accessible benthic habitats for diving and sea ducks that feed largely on
invertebrates.
Geese- Most of the Pacific (black) brant population and the world population of emperor geese
migrate through the greater Bristol Bay and Alaska Peninsula regions during spring and fall.
Pacific brant breed mostly on the Y-K Delta and also along the Arctic Coasts of Alaska,
northeast Russia, and Canada (Pacific Flyway Council, 2002; Reed et al., 1998). During April,
Bristol Bay and the Alaska Peninsula serve as staging areas for birds assessing snowmelt and
Bering Sea ice conditions. Chagvan and Nanvak Bays near Cape Newenham hold about 50,000
brant during spring, but only small numbers of brant use Nushagak or Kvichak Bays. From late
August through mid-September, most brant leave breeding areas and move south along western
Alaska directly to the Alaska Peninsula and eventually to Izembek Lagoon where the entire
population stages until early November. Most brant depart the Peninsula en masse across the
Pacific to wintering grounds from British Columbia to California, but most settle in the large
bays of Baja Mexico (Dau, 1992; Pacific Flyway Council, 2002). Since the mid-1970s, milder
conditions have allowed up to one-third of Pacific brant to winter along the western end of the
Alaska Peninsula (Ward et al., 2009).
Emperor geese breed almost entirely on the coastal zone of the Y-K Delta, with a few in Russia
(Pacific Flyway Council, 2006; Petersen et al., 1994). They winter from the outer Alaska
Peninsula westward into the Aleutian Islands. Like brant, emperor geese migrate in spring and
fall through the greater Bristol Bay area and eastern Bering Sea coast. They stage and migrate
mainly through bays to the west of the Nushagak Peninsula, but a few emperor geese occur in
inner Bristol Bay. During fall large numbers of emperor geese cross Bristol Bay to the large
lagoons on the north Alaska Peninsula, particularly the Seal Islands, Nelson Lagoon, and
Izembek Lagoon (Petersen et al., 1994).
Ducks- The nearshore waters of Bristol Bay host a large variety of ducks, including most of the
common dabbling and diving duck species that use an array of habitats from intertidal marshes to
offshore waters. During spring surveys in 1976-77, diving duck densities were 10-100/km2 along
the north side of Kvichak Bay and around Nushagak Bay (Arneson, 1980). Data from Kvichak
Bay indicated 259 birds/km2, mostly shorebirds, dabbling ducks (8 I/km2), and diving ducks on
tide flats. In south and east Nushagak Bay, high densities (17I/km2) of scaup (Aythya spp.) were
concentrated along Flounder Flats, mixed with flocks of black scoters (Melanitta americand).
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Deeper more open waters are used by diving ducks and sea ducks. Relatively high densities of
sea ducks (26/km2) were recorded during surveys on the east side of the Nushagak Peninsula
(Arneson, 1980), composed of 10% long-tailed ducks (Clangula hyemalis) and 12% harlequin
ducks (Histrionicus histrionicus). The majority of ducks in exposed waters and areas out to > 10
m deep were greater scaup, scoters, and eiders (see below). During spring and fall, smaller
numbers of long-tailed ducks, harlequin ducks, and goldeneyes (Bucephala spp.) occur in the
inner Bay. Up to 48,000 long-tailed ducks have been recorded in greater Bristol Bay during April
(Larned and Bellinger, 2011) on their way inland or northward for breeding; few long-tails occur
during fall (Mallek and Dau, 2011). Harlequin ducks use the bay before and after breeding inland
and on their way west to winter in the Aleutian Islands. Mallek and Dau (2011) counted 3,300
harlequins along the Bristol Bay and Alaska Peninsula coast during spring 2010 (peak count
6,114 in 1992) (USFWS, 1976).
Scoters- Bristol Bay, especially Kvichak Bay, is an important staging and molting area for black,
surf (Melanitta perspicillatd), and white-winged (M. fused) scoters spring through fall. Early
surveys often did not accurately record scoters by species, but over the past ten years, focused
surveys have produced species estimates, especially for black scoters.
Spring OCSEAP surveys along more pelagic transects tallied over 253,000 scoters in May 1972
and 216,000 scoters in April 1973 (USFWS, 1976). Arneson (1980) estimated that black scoters
comprised 97% of all scoters counted during spring surveys in 1976 and 1977. More recent
spring estimates also have documented large number of scoters in the inner bay, including up to
45,000 black scoters (Larned, 2008).
Scoters gather in Bristol Bay during the wing molt from July through September. OCSEAP
surveys counted 180,000 in July 1973 (Dau, personal communication; USFWS, 1976). A high
proportion of 77 satellite-marked black scoters from several wintering areas gathered in northeast
Bristol Bay where they spent an average of 15-20 days from June through September (Schamber
etal., 2010).
Though King and McKnight (1969) did not provide bird distribution data, they documented
about 181,000 scoters during October staging, including approximately 140,000 black scoters.
On the more pelagic OCSEAP surveys, over 285,000 scoters were estimated in the outer bay
during October 1974 (USFWS, 1976). Larned and Tiplady (1998) found about 20,000 black
scoters in the bay during late September. Few scoters are thought to winter in upper Bristol Bay
(Bellrose, 1980; Schamber et al., 2010); most probably disperse westward along the Alaska
Peninsula.
Satellite telemetry indicates that black scoters from widely separate wintering areas (British
Columbia, Kodiak, and Dutch Harbor) all used Bristol Bay from spring through fall (Bowman et
al., 2007). Analysis of cumulative satellite locations throughout the year indicates that black
scoters mostly use specific areas of shallow (< 3 m) waters along the north side of Kvichak Bay,
western Nushagak Bay, and Egegik Bay to the south (Schamber et al., 2010).
King Eiders- King eiders (Somateria spectabilis) in North America breed across the Alaska
North Slope and Arctic Canada, but inner Bristol Bay waters are important to king eiders as both
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a wintering area and as a major spring staging area (Suydam, 2000). Spring, OCSEAP surveys
estimated that about 280,000 king eiders were in outer Bristol Bay in May 1972, and over 1.8
million were found in April 1973 (USFWS, 1976). Arneson (1980) estimated eider composition
was about 45% king, 36% common (S. mollissima), and 19% Steller's (Polysticta stelleri) during
spring coastal surveys in 1976 and 1977.
In the late 1990s, king eiders marked with satellite tags guided aerial surveys to document
staging and molting areas (Larned and Tiplady, 1998). During April surveys for Steller's eiders,
Larned (2008) estimated over 570,000 king eiders in Kvichak Bay in 2008; an average of
194,000 king eiders were estimated from these surveys from 2000 to 2009 (Larned and
Bellinger, 2011).
Bristol Bay has been documented as one of a few important molt areas for king eiders; other
areas include the northeast Russian coast and St. Lawrence Island (Phillips et al., 2006). Based
on satellite tracking, Bristol Bay molters arrive from Alaska's North Slope (Phillips et al., 2006)
and Arctic Canada (Dickson et al., 2001). Molt periods vary by sex and age of birds, but extend
from August to October.
During fall, smaller numbers of king eiders have been recorded in the bay; as many as 20,000
were estimated in late September (Larned and Tiplady, 1998). Southwestern Alaska, especially
inner Bristol Bay, is considered one of three main wintering areas of king eiders breeding in
western North America, though these birds may move considerably within the region between
October and April (Oppel et al., 2008).
Schamber et al. (2010) assessed the distribution of king eiders in Bristol Bay from year-round
locations of satellite-marked birds. Across seasons, king eiders used most of the inner bay
between the Nushagak Peninsula and Egegik Bay (averaging 10.6 km offshore), including areas
with water depths of > 20 m, but they particularly frequented defined areas off Etolin Point, Half
Moon Bay, and Egegik Bay where water depths were < 10 m.
Steller 's Eiders- Bristol Bay coastal waters host Steller's eiders mostly in spring and fall, but
they are not thought to breed in the region. The historical breeding range of the species in Alaska
extends from the Y-K Delta into northwest Alaska and the western North Slope (Fredrickson,
2001). Along with a large number of Steller's eiders that breed west to the central Siberian coast,
the Alaska birds are part of a Pacific (Russia-Alaska) population that probably numbers between
130,000 (Hodges and Eldridge, 2001; USFWS, 1999) and 150,000 (Fredrickson, 2001). Though
historical data are not quantified, the number of Steller's eiders breeding in Alaska declined
sometime between the 1940s and 1960s, especially on the Y-K Delta (Kertell, 1991). Based on
estimates that there were perhaps fewer than 3,000 birds breeding over a substantially reduced
range in Alaska during the 1990s, the USFWS listed the Alaska-breeding component of the
population as threatened under the Endangered Species Act in 1997 (USFWS, 2002).
The primary wintering grounds for the Pacific population extends from the central Alaska
Peninsula westward into the Aleutian Islands (Fredrickson, 2001). Thus, most birds transit
Bristol Bay in spring and fall. Surveys have been conducted to assess the Pacific population
during spring migration (April-early May) since 1992, including survey sections between Cape
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Constantine and the Naknek River (Larned and Bellinger, 2011). In general, spring staging
Steller's eiders were concentrated along the Alaska Peninsula south of Egegik and west of the
Nushagak Peninsula; very few birds used inner Bristol Bay. Coastal emperor goose surveys
flown in late April show similar minimal occurrence of Steller's eiders in Nushagak and Kvichak
Bays (Dau, personal communication; Dau and Mallek, 2011).
After breeding on more northern nesting areas, a large proportion of Pacific Steller's eiders begin
to return to southwest Alaska in late June, in advance of the wing molt (Petersen, 1981). Few of
these birds use inner Bristol Bay as they concentrate in the lagoons along the Alaska Peninsula
(Dau, personal communication; Mallek and Dau, 2011). Molt migration progresses by sex and
age classes, and the actual molt period extends from late July for subadults to October for adult
females (Petersen, 1981). Bird numbers increase during fall staging, mainly in Nelson Lagoon
and Izembek Lagoon, before moving westward to winter in the Aleutian Islands.
Bristol Bay Lowlands-
Habitats- The Bristol Bay lowlands are characterized by old glacial deposits with moraine lakes
and ponds, glacial outwash and riverine deposits along floodplains, and mixed marine deposits
near the mouths of the Nushagak and Kvichak Rivers. Landcover is mostly moist and wet tundra
between Nushagak and Kvichak Bays, and in a broad region of the upper Mulchatna drainage.
Tundra merges into lowland spruce-hardwood forest between the lower Nushagak River and the
Wood-Tikchik Lakes, and in the Kvichak Valley to Iliamna Lake (Selkregg, 1976). This
subregion has a wide diversity of freshwater lakes and ponds, as well as numerous floodplain
wetlands.
Stratum 8 of the AYWBPS generally defines the "lowlands," including the area southwest of
Iliamna Lake to the Nushagak Peninsula and extending southwest from Naknek River to Cinder
River. Table 8 indicates the average indices and densities of 30 groups (32 species) of waterfowl
recorded on aerial surveys flown annually in late May.
Swans- Most swans in the Bristol Bay region are tundra swans that comprise 10-15% of the
Western Population which breeds from Kotzebue Sound to the outer Alaska Peninsula and
winters from British Columbia to central California (Ely et al., 1997; Pacific Flyway Council,
2001). The most recent ten-year average index of swans from Bristol Bay (AYWBPS Stratum 8)
is 15,400 (0.6/km2). Tundra swans arrive as early as mid-March and numbers peak in late April
(Wilk, 1988). The majority of swans move north to the Y-K Delta region, but those that breed in
Bristol Bay initiate nests in early May and young hatch in early- to mid-June. Because Bristol
Bay has an earlier spring thaw, the phenology of local breeding swans is 2-4 weeks earlier than
those nesting on the Y-K Delta and northern Alaska. Wilk (1988) provides indications that
earlier nesting in Bristol Bay supports larger average brood sizes and higher productivity.
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8,
of In May on the
Survey, 8)
Long-term Average
Species/Group 1957-2011
Mallard
Gadwall
American Wigeon
Green-winged Teal
Blue-winged Teal
Northern Shoveler
Northern Pintail
Redhead
Canvasback
Scaup (Lesser, Greater)
Ring-necked Duck
Goldeneye (Common, Barrow's)
Bufflehead
Long-tailed Duck
Unidentified Eider
Common Eider
Spectacled Eider
Steller's Eider
King Eider
Unidentified Scoter
Surf Scoter
White-winged Scoter
Black Scoter
Merganser (Common, Red-breasted)
TOTAL DUCKS
White-fronted Goose
Canada/Cackling Goose
Emperor Goose
(Pacific ) Brant
TOTAL GEESE
Swan (Tundra, Trumpeter)
Sandhill Crane
33,100
1,400
25,200
30,600
0
13,300
57,300
0
200
79,800
400
4,200
500
13,700
900
0
0
0
0
79,400
n/aa
n/aa
n/aa
2,700
346,500
5,100
2,400
0
0
7,600
12,100
3,300
Average Index
2002-2011
68,100
2,000
55,300
71,800
200
33,500
82,100
0
200
94,000
0
1,600
300
5,200
500
100
0
0
0
36,800
400
2,300
37,600
5,300
497,000
5,300
2,300
0
100
7,700
15,400
5,300
10-Yr Average
Birds/km2
2.68
0.08
2.16
2.80
0.01
1.30
3.20
0.00
0.01
3.67
0.00
0.06
0.01
0.20
0.02
0.00
0.00
0.00
0.00
1.43
0.01
0.09
1.47
0.21
19.38
0.21
0.09
0.00
0.00
0.30
0.60
0.20
Scoters have been recorded by species only since 1993.
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Geese- Long-term average indices of geese in the region have been relatively stable over the past
30 years (Table 8), averaging 7,700 geese (0.3/km2) during 2002-11, with greater white-fronted
geese over twice as abundant as Canada geese (AYWBPS). Platte and Butler (1995) tallied 4,255
geese (0.09/km ) over a survey area farther inland and estimated composition as 55% white-front
and 45% Canada geese. Bristol Bay white-fronts currently comprise only a small portion of the
Pacific Flyway population which breeds mostly on the Yukon-Kuskokwim (Y-K) Delta and
numbers over 600,000. During the late 1970s and early 1980s, when Pacific white-fronts were
overharvested and declined more than 80% (Pacific Flyway Council, 2003; Pamplin, 1986),
Bristol Bay white-fronts made up about 15% of the population.
White-fronted geese from Bristol Bay have been shown to be slightly larger than most Pacific
Flyway white-fronts from the Y-K Delta (Ely et al., 2005; Orthmeyer et al., 1995) though they
are not considered taxonomically separate. Bristol Bay white-fronts also migrate south earlier
than others in fall, passing through the Klamath Basin of Oregon and California in September,
and overflying the Sacramento Valley where most Pacific white-fronts winter. Bristol Bay birds
press further south to winter in the northern highlands of Mexico (Ely and Takekawa, 1996).
The Canada/cackling geese that breed in the Bristol Bay region include Taverner's cackling
geese (Branta hutchinsii Taverneri) and lesser Canada geese (Branta canadensis parvipes), the
former found closer to the coast. Taverner's geese breed extensively along the western and
northern coastal regions of Alaska, and lesser Canada geese are found throughout Interior Alaska
and Yukon Territory, but the breeding ranges of these two populations have not been delineated
and there are no reliable population indices (Pacific Flyway Council, 1994). In fall, most lesser
Canada geese migrate through Cook Inlet and along the Alaska coast to winter from British
Columbia into Washington and Oregon while Taverner's geese staging on the western Alaska
Peninsula make a direct migration across the Gulf of Alaska to wintering areas. During winter,
most Taverner's and Lessers aggregate with over 250,000 other white-cheeked ("Canada") geese
in southwest Washington and western Oregon. They also are found in the upper Columbia River
Basin and east of the Cascade Mountains. Intermingling of populations precludes accurate winter
inventories.
The smallest subspecies of cackling Geese (B. h. minima) migrates through the region en route to
and from the Y-K Delta coast where they breed. Cackling geese are assessed annually on their
breeding grounds, with survey results indicating substantial increases from a low of <30,000 in
1984 to 150,000-200,000 since 1997. During fall, nearly all of these cacklers historically staged
along the Alaska Peninsula near Pilot Point and Cinder River (Sedinger and Bellinger, 1987).
Since recovery from a major population decline from overharvest through the early 1980s
(Pamplin, 1986), fall staging of cacklers has been more dispersed westward along the Alaska
Peninsula (Gill et al., 1997) from which they migrate across the Gulf of Alaska. Prior to the
1980s, cacklers wintered in Central California, but now the majority of cacklers winter in the
Willamette Valley of western Oregon and near the Lower Columbia River in southwest
Washington (Pacific Flyway Council, 1999).
Brant and emperor geese move through Bristol Bay coastal habitats in spring and fall (see
Estuaries and Inner Bristol Bay), but the lowlands are not considered a breeding area. Lesser
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snow geese (Chen caerulescens) are occasionally seen in Bristol Bay during migration to and
from Wrangel Island in Russia.
Ducks- Indices of duck abundance generally have been higher than long-term averages since
1995, as measured by the AYWBPS; the most recent ten-year average is about 497,000 ducks
(19.4 ducks/km2) (Table 8). The most prevalent duck species include greater and lesser scaup
(Aythya marila and A. affinis'., 3.7/km2), northern pintail (Anas acuta; 3.2/km2), green-winged
teal (A. crecca; 2.8/km ), mallard (A. platyrhynchos; 2.7/km2), and American wigeon (A.
americana; 2.2/km2).
In their expanded survey area, Platte and Butler (1995) estimated averages of 355,200 ducks
(7.12/km ). Duck species composition was similar to the AYWBPS, with the highest average
densities ranked as scaup (1.9/km2), northern pintail (1.1/km2), green-winged teal (1.0/km2), and
mallard (0.9/km2). Gadwall (A. streperd), wigeon and shoveler (A. clypeata), at lower densities,
were more prevalent south along the Alaska Peninsula. High to medium densities of scaup were
recorded west of Iliamna Lake near the upper Kvichak and Alagnak Rivers.
Scaup- Greater and lesser scaup cannot be differentiated on aerial surveys, but most of the scaup
breeding in tundra regions are assumed to be greater scaup (Hodges et al., 1996). The relatively
high densities of scaup in the Bristol Bay lowlands recorded on the AYWBPS and expanded
surveys suggest that this region hosts a substantial portion of the breeding greater scaup in North
America. About 80% of greater scaup migrate across the continent in fall, stopping in the Great
Lakes, and wintering along the northeast Atlantic Coast (Kessel et al., 2002). Others winter from
south-central Alaska down the Pacific Coast.
Black scoters- Bristol Bay is recognized as one of the most important breeding areas for the
western (Pacific) population of black scoters that occupies Alaska and western Canada. The
Pacific population may number 200,000-400,000 birds (Bordage and Savard, 1995). The
AYWBPS does not provide reliable indices for breeding scoters because it is flown before nest
initiation, which is later (June) than other ducks, and because scoters also are found in taiga and
boreal habitats outside traditional survey areas. Also, in the past, scoters were not identified to
species level during AYWBPS surveys.
Through the Sea Duck Joint Venture, USFWS has been conducting additional aerial surveys
designed for scoters in Alaska since 2007 to improve population estimates (Stehn et al., 2010;
Stehn et al., 2006), particularly for black scoters that have shown historic declines (Bordage and
Savard, 1995). The new surveys have produced recent estimates of 173,000 black scoters on all
western Alaska tundra breeding areas (Stehn et al., 2010). Bristol Bay is an important breeding
area, containing 46,100 black scoters (0.92/km2), about 15% of surveyed ducks, in their
expanded survey area (Platte and Butler, 1995). Densities were highest in a band from western
Kvichak Bay to Lake Iliamna, and also along the western Alaska Peninsula between Egegik and
Ugashik Bays.
Inland Tundra/Taisa-
Habitats- The inland subregion of Bristol Bay is underlain by glacial deposits that are
interspersed with bedrock formations in the upper Kvichak and Mulchatna drainages. Bedrock
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dominates in the montane areas south and east of Lake Clark. Vegetation communities represent
a transition from moist tundra to the west into tall shrub habitats and upland spruce hardwood
forest; alpine tundra rises into the Aleutian Range (Selkregg, 1976). The subregion has abundant
aquatic habitats from alpine lakes and glacial lakes to wet tundra wetlands and floodplain basins
along rivers.
Williamson and Peyton (1962) reviewed the characteristics and historical ecological
classifications of the Iliamna Lake region, concurring with previous observers that the region's
dominant feature is transitional communities where the interior/arctic, southcentral coast forest,
and western tundra ecotypes meet. They indicate that neither the dissected Aleutian Range to the
east nor the open plateaus to the west serve as barriers for avifauna. These authors describe 12
ecological formations (habitat types) and their associated birds species. Generally, Williamson
and Peyton (1962) recorded strong associations of 15 waterfowl species to open lakes and ponds,
secondary preferences for streams and rivers by most species, and use of freshwater marshes by
dabbling ducks.
Aside from less detailed historical accounts, information on waterfowl distribution and
abundance is found in Williamson and Peyton (1962), some aerial survey coverage by Platte and
Butler (1995), and a few observations in montane habitats by Ruthrauff et al. (2007).
The most current waterfowl surveys for the inland/Iliamna region are from contract projects for
Pebble Limited Partnership (PLP). They include aerial surveys for waterbirds from 2004 to 2006
north of Iliamna Lake during spring staging, breeding/brood surveys, and fall. Some surveys
focused on harlequin ducks and tundra swans. Overall, 22 species of breeding waterfowl were
recorded (including scaup, goldeneye, mallard, green-wing, pintail), averaging 10 ducks/km2.
Only very general summary information has been made available on the PLP website and in
public presentations (Pebble Partnership, 2011).
Swans- Trumpeter swans (Cygnus buccinator} are found in the eastern portion of the Bristol Bay
region, associated with the forested and taiga habitats typical of the boreal zone. This area abuts
temperate coastal forest and coastal marshes occupied by trumpeter swans in Cook Inlet. As their
name suggests, tundra swans are primarily birds of open tundra. They have an extensive breeding
distribution from the Alaska Peninsula and Bristol Bay coast inland to the Iliamna Lake and
Lake Clark regions. The Pacific Coast Population of trumpeter swans has been increasing,
including breeders on the Kenai Peninsula, Cook Inlet lowlands and upper Kuskokwim valley.
There is increasing evidence that trumpeters are expanding their range westward into tundra
swan habitats (Pacific Flyway Council, 2008).
Geese- Lesser Canada geese (B.c. parvipes) are found throughout southcentral Alaska and
interior portions of southwest Alaska. Though survey data and descriptions of geese are not
readily available, it is possible that some Taverner's cackling geese (B.h. tavernerf) may breed in
the inland subregion. The ranges of these similar medium-sized geese have not been delineated.
Canada/cackling geese are listed as common in Lake Clark National Park (National Park Service,
2011) and in the Kvichak River valley. Greater white-fronted and snow geese occur during
spring and fall migrations to and from Cook Inlet, and brant have been recorded rarely.
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Ducks- The densities of dabbling ducks are only moderate in the inland subregion, with mallards,
northern pintail and green-winged teal as the most common. Aerial surveys suggest pockets of
spring duck habitat near the head of the Kvichak River and the upper portions of the Chulitna
and Mulchatna drainages (Platte and Butler, 1995). American wigeon and northern shovelers
were less abundant. Scaup and black scoters associated with boreal habitats also occurred in the
same general areas, with densities in some up to >8 birds/km2. Among sea duck species, long-
tailed duck, surf scoter, white-winged scoter, common goldeneye (Bucephala clangula\
Barrow's goldeneye (Bucephala islandica), red-breasted merganser (Mergus serrator) and
common merganser (Mergus merganser} were distributed more sparsely and at low densities.
Harlequin ducks are found in low densities throughout the Bristol Bay region, using clear high-
gradient streams during the breeding season. Harlequins are difficult to detect on aerial surveys
due to the habitat they use and are often underestimated.
Nutrients, Trophic Relations and Foods
For waterfowl, body condition, reproductive success, and survival are dependent on the quantity
and quality of foods available throughout their annual cycle. Waterfowl typically experience the
greatest energetic demands before and during migrations; in the pre-breeding period when
resources are needed for egg-laying, incubation, and territory defense; and during summer molt
when feather replacement taxes reserves. Though stored reserves (body fat and protein) may be
used during migration and incubation, waterfowl need to select habitats that have abundant food
resources, be able to efficiently exploit specific foods, and be mobile to optimize seasonal
foraging strategies.
Nutrients and Habitat Productivity-
In Alaska, the most productive waterfowl habitats are those that have dynamic nutrient systems
that produce seasonally rich plant and animal foods for birds. For example, the mixing of marine
nutrients from upwellings and terrestrial nutrients from rivers and streams enrich the productivity
of coastal estuaries and lagoons. Large river deltas (Stikine, Copper, Yukon-Kuskokwim,
Colville, etc.) are the best examples of complex interfaces of marine and terrestrial nutrients in a
matrix of low-lying depositional wetlands, and they support high densities of breeding
waterfowl. The estuaries and nearshore waters of Nushagak and Kvichak Bays are enriched by
such nutrient mixing (Straty, 1977) in coastal marshes used by swans, geese, and dabbling ducks,
and abundant benthic invertebrates for diving and sea ducks.
The productivity of waterfowl habitats in inland-interior regions are also based on dynamic
nutrient systems mostly associated with river basin wetlands and floodplains. Within the
extensive watersheds of large rivers, nutrient flux (largely nitrogen and phosphorus) is driven by
upstream erosion, in-stream transport and seasonal flooding. In forested areas, mosaic patterns of
wild fires can also be sources of nutrients for wetlands (Bayley et al., 1992). In the Bristol Bay
region, Selkregg (1976) notes a long history of wild fires in the Mulchatna Valley; upland
spruce-hardwood forests are found in the upper drainages of the Mulchatna and Nushagak
Rivers. Seasonal nutrient inputs refresh and subsidize primary productivity and development of
aquatic invertebrates in floodplains and large wetland basins associated with valleys (Heglund,
1992). In the Nushagak and Kvichak River systems, the large volume of sockeye and other
salmon species is a significant source of imported nutrients throughout these watersheds (see
below).
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FoodHabits-
The waterfowl of Bristol Bay exhibit a wide diversity in foraging strategies and food habits;
some species tend to be specialists and some are generalists, but nearly all adapt seasonally to
different foods. Among swans, geese and ducks during the breeding season, some species defend
territories that are selected for sufficient food supplies for nesting adults and growing young;
others adapt more social and mobile strategies to take advantage of temporarily abundant foods.
During non-breeding periods (migration, molt and winter) waterfowl often aggregate and exhibit
fidelity to habitats and sites with reliable food resources.
In general, swans and geese are primarily vegetarians, although adults and young feed
opportunistically on insects, aquatic invertebrates and other animal foods especially during
breeding and brood-rearing season. Both trumpeter and tundra swans feed on submergent and
emergent vegetation year round. The development of cygnets to fledging often extends into fall
when their freshwater habitats begin to freeze and they may move to aquatic beds in flowing
waters and coastal shallows prior to migration (Limpert and Earnst, 1994.; Mitchell and
Eichholz, 2010).
Geese are primarily vegetation grazers, although adults and goslings opportunistically feed on
insects and invertebrates. Canada, cackling, and white-fronted geese consume shoots and stems
of graminoid plants in typical moist and wet tundra breeding areas. In coastal areas, broods are
often brought to wetland basins, tide flats and salt marshes where foods are abundant. During fall
staging, Canada and white-fronted geese often frequent uplands to feed on berries. Canada geese,
emperor geese and brant are associated with coastal habitats where they rely on salt marsh
vegetation, eel grass beds (Zoster a marina) in estuaries, and some marine algae (Petersen et al.,
1994; Reed etal., 1998).
The three tribes of ducks that use Bristol Bay have very diverse foods habits (Table 9). Dabbling
ducks (Anatini) are generally omnivorous, feeding on seeds and aquatic invertebrates, and
focusing on high protein foods prior to breeding. Diving ducks (Aythyini) are also omnivorous,
but they feed in more open waters for benthic invertebrates and focus on animal foods during
staging and wintering on coastal waters. Sea ducks (Mergini) consume mostly animal foods year
round, feeding on freshwater benthic invertebrates during breeding and a wide variety of marine
invertebrates during staging and wintering on saltwater.
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9. by of
Common Name
Dabbling Ducks
Gadwall
American Wigeon
Mallard
Northern Shoveler
Northern Pintail
Green-winged Teal
Diving Ducks
Canvasback
Redhead
Ring-necked Duck
Greater Scaup
Lesser Scaup
Sea Ducks
Steller's Eider
King Eider
General Food Habits
Primarily plant foods with some
invertebrates during pre-breeding
Strongly vegetarian with some
invertebrates during pre-breeding
Omnivorous, mostly plant seeds and
invertebrates, increasing animal foods
during pre-breeding
Small invertebrates and seeds strained
at the surface
Omnivorous, mostly grain and plant
seeds with more invertebrates during
pre-breeding
Omnivorous, mostly plant seeds and
invertebrates, increasing animal foods
during pre-breeding
Omnivorous, mostly plant buds,
tubers, root stock and invertebrates,
increasing animal foods in pre-
breeding
Omnivorous, mostly plant leaves and
stems, invertebrates; increasing animal
foods during pre-breeding
Omnivorous, mostly plant seeds and
invertebrates; increasing animal foods
during pre-breeding
Omnivorous with varied local and
seasonal focus on mollusks,
crustaceans, insects
Omnivorous with tendency toward
animal foods, insects, mollusks,
crustaceans
Mostly animal foods; insect larvae on
freshwater; crustaceans, mollusks,
other invertebrates on saltwater
Mostly insects and crustaceans with
some plant foods on freshwater;
Consumption of Fish
Unknown
Spring herring eggs Oregon coast
(Bayer, 1980)
Incidental fish; fall and winter use of
salmon eggs and flesh in spawning
lakes and coastal waters (Gleason,
2007; Munro, 1943)
Trace offish in winter
Unknown
Use of salmon eggs and salmon flesh in
spawning streams (Gabrielson and
Lincoln, 1959)
Incidental small fish (Cottam, 1939);
alewife fingerlings on fall migration in
New York. Spring herring eggs Oregon
coast (Bayer, 1980). Salmon flesh
Washington coast (Dawson and
Bowles, 1909)
Spring herring eggs Oregon coast
(Bayer, 1980). Largemouth bass eggs
in freshwater (Jarvis and Noyes, 1986).
Incidental small fish (Cottam, 1939)
Incidental small fish (Cottam, 1939)
Incidental small fish (Cottam, 1939);
fall and winter use of salmon eggs and
flesh in coastal streams and lakes
(Munro, 1941). Spring herring eggs
(Bayer, 1980; Munro, 1941)
Light use of 8 species of fish and
salmon eggs (Cottam, 1939); 3 species
of fish during fall in Minnesota (Afton
et al 1991); winter scavenging flesh of
shad and sunfish (Christopher and Hill,
1988); Spring herring eggs Oregon
coast (Bayer, 1980)
Low occurrence of small fish, probably
incidental to benthic feeding (Cottam,
1939).
Spring use of lumpfish eggs in Norway;
small amounts of sculpins and fish eggs
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Common Name
General Food Habits
Consumption of Fish
mollusks, crustaceans and diverse
marine invertebrates on saltwater.
in Alaska during winter (Cottam, 1939)
Common Eider (Pacific)
Mostly animal foods, mollusks,
crustaceans and benthic marine
invertebrates
Spring use of herring eggs (Cantin et
al., 1974); scarce use of sculpins and
sculpins roe in winter (Cottam, 1939)
Harlequin Duck
Foods almost entirely animal;
freshwater invertebrates during
nesting; diverse mollusks, crustaceans
and other marine invertebrates during
most of the year
Herring eggs in spring (Munro and
Clemens, 1931; Vermeer, 1983);
occasional freshwater fish and eggs; fry
of char (Kistchinski, 1968); high use of
salmon roe in streams in late summer
(Dzinbal and Jarvis, 1984); salmon
carcasses in streams and estuaries
(Vermeer and Levings, 1977)
Surf Scoter
Mostly animal foods, insects and
clams on freshwater; mollusks,
crustaceans and other invertebrates on
saltwater
Minor use offish, but more than white-
winged or black scoters. Herring eggs
in spring (Bayer, 1980; Vermeer, 1981)
White-winged Scoter
Mostly animal foods; insects, clams
and some plant material on
freshwater; mollusks, crustaceans and
other invertebrates on saltwater
Minor use of fish on fresh and
saltwater; coastal herring eggs in spring
(Bayer, 1980; Cottam, 1939; Munro
and Clemens, 1931)
Black Scoter
Mostly animal foods; insects,,
crustaceans and some plant foods on
freshwater; mollusks and crustaceans
on saltwater
Some use of freshwater fish eggs
(Bengtson, 1971). Spring herring eggs
(Bayer, 1980; Munro and Clemens,
1931)
Long-tailed Duck
Mostly animal foods; insects,
crustaceans and some plant foods on
freshwater; mollusks and crustaceans
on saltwater
Some use of freshwater fish (Peterson
and Ellarson, 1977). Herring eggs in
spring (Munro and Clemens, 1931)
and some bottom fish on saltwater in
winter (Cottam, 1939; Sangerand
Jones, 1984)
Bufflehead
Mostly animal foods with some plant
material seasonally; insects and
crustaceans on freshwater; crustaceans
and mollusks in saltwater
Minor use offish on freshwater;
salmon eggs in coastal streams; fall and
winter use of small fish; herring eggs in
spring (Bayer, 1980; Munro, 1942;
Vermeer, 1982)
Common Goldeneye
Mostly animal foods; insects,
mollusks and crustaceans on
freshwater; crustaceans and mollusks
in saltwater
Diverse small fish and roe in
freshwaters and saltwater (Cottam,
1939; Jones and Drobney, 1986);
salmon eggs and flesh on coastal lakes
and rivers (Taverner, 1934); herring
eggs in spring (Munro and Clemens,
1931; Vermeer, 1982)
Barrow's Goldeneye
Mostly animal foods; insects,
mollusks and crustaceans on
freshwater; crustaceans and mollusks
in saltwater
Herring eggs in spring (Munro and
Clemens, 1931; Vermeer, 1982); small
numbers of sculpins taken; salmon eggs
and parr on freshwater (Fitzner and
Gray, 1994; Munro, 1923); salmon
flesh (Cottam, 1939)
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Common Merganser
Primarily small fish, also insects,
mollusks, crustaceans, and small
vertebrates
Very diverse fish prey. Salmon are
most important in some regions (Munro
and Clemens, 1932; Munro and
Clemens, 1937; Salyer and Lagler,
1940)
Red-breasted Merganser
Primarily small fish, also insects,
mollusks, crustaceans, and small
vertebrates
Diverse fish prey. Salmon are
important in some regions (Munro and
Clemens, 1939; White, 1957). Also
take herring and roe
Information obtained primarily from species accounts in the Birds of North America (A. Poole, Ed.).
Cornell Lab of Ornithology, Ithaca, NY. Retrieved from the Birds of North America Online:
http://bnii.birds.cornell.edu/bna/
Importance of Marine-Derived Nutrients (Salmon and Herring) to Waterfowl-
Waterfowl benefit from salmon as both direct sources of prey and carrion and indirect nutrient
drivers of aquatic systems (i.e., supporting invertebrate prey species and riverine plant
communities). Roughly 30-40 million salmon spawn annually in the Kvichak and Nushagak
systems (Hilborn et al., 2003; Ruggerone et al., 2010), importing perhaps 20 million kg (44
million pounds) of nutrients throughout the watersheds. The fate of these nutrients (primarily
nitrogen and phosphorus) is divided among the breakdown of carcasses (Cederholm et al., 1989;
Cederholm et al., 1999) and deposition in drainages, the outmigration of smolts (Crawford, 2001;
Moore and Schindler, 2004), and discharge into estuaries. This large influx of nutrients and net
gain to riverine and terrestrial systems strongly affects a wide variety of plants and animals,
including waterfowl (Gende et al., 2002; Holtgrieve, 2009; Willson et al., 1998; Willson and
Halupka, 1995).
Of the 24 duck species that regularly occur in Bristol Bay, at least eleven species are known to
prey on salmon eggs, parr, smolts, and scavenge on flesh of spent carcasses (Table 9). Of these,
greater and lesser scaup, harlequin duck, buffiehead, common and Barrow's goldeneyes, and
common and red-breasted mergansers exhibit directed foraging on salmon. Among dabbling
ducks, mallards feed most on salmon because they are distributed across a diversity of summer
habitats in spawning areas, and they are the principal wintering dabbling duck on the North
Pacific coast where fall-winter salmon runs occur.
From early May through June, salmon smolts emigrate from Bristol Bay rivers, providing
abundant prey (325 million smolt in the Kvichak) (Crawford, 2001) for fish predators like
mergansers (Munro and Clemens, 1932; Munro and Clemens, 1937; Munro and Clemens, 1939;
Salyer and Lagler, 1940; White, 1957; Wood, 1987a; Wood, 1987b). Other duck species may
prey on smolt incidentally.
From late June through early September, salmon eggs are readily available on and downstream
of spawning beds. These eggs are a seasonally rich food source for harlequin ducks, goldeneyes
and scaup that frequent rivers and streams (Cottam, 1939; Dzinbal and Jarvis, 1984; Munro,
1923) and probably for other opportunistic ducks.
From mid-July through September, salmon carcasses are abundant in streams, rivers and
spawning lakes. Though the scientific literature is scarce on this subject, species ranging from
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dabbling ducks (mallard, green-winged teal) and diving ducks to sea ducks that inhabit spawning
waters opportunistically scavenge easy protein-rich meals (Table 9).
Harlequin ducks offer an example of a waterfowl species that utilizes salmon in all life stages.
Harlequins breed in high gradient streams in the upper reaches of drainages (Robertson and
Goudie, 1999), and they forage in portions of rivers and streams occupied by salmon. In the
Kolyma Highlands in Russia, harlequins focused on fry of Dolly Varden (Salvelinus malmd) and
white-spotted char (S. leucomaenis) (Kistchinski, 1968). Gudmundsson (1971) noted a
relationship between harlequin duck nesting areas and Atlantic salmon (Salmo salaf) spawning
areas in Iceland, but thought that ducks and salmon fry preyed on the same insect larvae. Dzinbal
and Jarvis (1984) demonstrated that harlequins breeding in the short streams of northern Prince
William Sound depended heavily on the dislodged eggs of pink (Oncorhynchus gorbuschd) and
chum (O. keta) salmon in the lower stream reaches from the first week of July to early August. In
their study, the drifting biomass of salmon eggs exceeded the biomass of invertebrates in late
July and early August. Harlequin ducks also scavenge from salmon carcasses from August
through September (Vermeer and Levings, 1977). Winter diets did not include notable
occurrence offish (Fischer and Griffin, 2000; Vermeer, 1983).
Spring spawning of Pacific herring (Clupea pallasi) along Alaska coasts provides abundant food
for waterfowl and many other waterbirds in need of energy during migration, and sea ducks
follow the progressive spawning northward (Lok et al., 2008). Though the waters of inner Bristol
Bay are turbid and not conducive to herring spawning, small concentrations may be found. The
region's center of herring activity is the Togiak district west of Cape Constantine from Kulukak
Bay to Cape Pierce (ADF&G, 201 Ic). Sixteen of Bristol Bay's duck species feed on herring and
herring roe during spring (Table 9; (Bayer, 1980; Lok et al., 2011; Lok et al., 2008; Munro and
Clemens, 1931; Vermeer, 1983)).
Populations, Subpopulations, and Genetics
Among waterfowl, the designation of subspecies, populations, and subpopulations has been
applied through increasing research on genetic diversity and relatedness; a long history of
morphological measurements; and evaluation of cohesiveness, philopatry, and annual
distributions of birds from traditional banding and marking studies. Overall, few population units
below the species level have been established for swans, geese, and ducks, probably because
their extensive migrations and mobility across broad ranges provide genetic homogeneity. The
American Ornithologist's Union (AOU) no longer taxonomically designates subspecies
(American Ornithologists' Union, 2011) because of the difficulty in differentiating valid subunits
and the dynamic nature of evolving groups.
In some cases, subspecies and populations have been defined when genetic, morphological and
observational data support designations that are practical for population management, yet are
provisional in terms of taxonomy. Population units also have been designated for purposes of
monitoring biodiversity by programs such as the Alaska Natural Heritage program (Alaska
Natural Heritage Program, 2011), but under varying scientific standards that may be less
rigorous than taxonomic determinations. In addition, populations may be defined and designated
for protection under the Endangered Species Act without strict evidence of discreteness or
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genetic distinction. Following is a summary of current understanding of population structuring in
the principal waterfowl species of Bristol Bay.
Swans-
Few distinctions have been made among trumpeter or tundra swans across their ranges.
Trumpeter swans in Alaska are part of the Pacific Coast Population (PCP) that constitutes over
80% of the species in North America. Recent genetic studies (Oyler-McCance et al., 2007)
indicate that PCP birds are distinguishable from Rocky Mountain Population (RMP) trumpeters
that range from Yukon Territory south to Wyoming and Utah, but within Alaska trumpeters are
fairly homogenous through the Interior and Cook Inlet. Copper River Delta breeders were
somewhat unique because of their geographic isolation. Samples were taken from Susitna Basin
and the Kenai Peninsula, but not from the eastern Bristol Bay region. Based on banding and
marking studies, tundra swans in southwest Alaska are affiliated with the Western Population
(WP) found wintering west of the Rocky Mountains (Pacific Flyway Council, 2001). The small
non-migratory group on the end of the Alaska Peninsula is considered part of the WP, not a
separate entity.
Geese-
The taxonomy of Canada geese has long been debated, but the recent species separation of
cackling geese (B. hutchimii) from Canada geese (B. canadensis) is based on extensive genetic
studies (Paxinos et al., 2002; Scribner et al., 2003) that support divergence of three small
subspecies (hutchinsif) from three larger subspecies (canadensis) during the last glacial period.
The coastal Taverner's (cackling) goose and inland lesser (Canada) goose breeding in the Bristol
Bay region, as well as the migrant Cackling (cackling) goose that passes through, have been
managed somewhat separately for over 60 years. The extensive historical banding information
and genetic evidence, warrants recognition of these populations among six white-cheeked goose
populations in Alaska.
As mentioned previously, white fronted geese breeding in Bristol Bay are slightly different in
morphology and migration patterns from other Pacific white-fronts nesting on the Y-K Delta, but
the differences do not rise to the level of taxonomic significance, nor can they practically be
managed separately (Ely et al., 2005; Orthmeyer et al., 1995; Pacific Flyway Council, 2003).
Dabbling and Diving Ducks-
Historically, there has been little to support identification of distinct populations among dabbling
ducks (tribe Anatini) and diving ducks (tribe Aythyinf) largely because of their extensive mobility
and exchanges across flyways. No population structure has been suggested among these ducks
for the Pacific Flyway or Bristol Bay, though extensive genetics studies have not been
conducted.
Sea Ducks-
Over the past 10 years, declining trends in most sea duck species (tribe Mergini) and the listing
of spectacled and Steller's eiders under ESA have prompted research into the structure and
diversity of sea duck populations. In general, sea ducks are known to be quite philopatric to
breeding, molting and wintering areas, suggesting the potential for discrete population units. In
addition, sea ducks from broad breeding ranges aggregate in winter, making winter a critical
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period of social and genetic interchange. These unique characteristics are important for
understanding the biology of sea ducks and adopting effective management regimes.
The biology and population dynamics of harlequin ducks are not well understood, but they breed
in the upper high-gradient drainages of Bristol Bay and occur in the upper bay, Alaska Peninsula,
and Cook Inlet during the non-breeding season. Though these birds are generally segregated by
river drainage during breeding and they demonstrate fidelity to wintering areas (Esler et al.,
2000), dispersal occurs across regions (Cooke et al., 2000). No evidence has been found to
genetically distinguish wintering harlequins from Prince William Sound, Kodiak and Katmai
areas (Lanctot et al., 1999 ). This suggests that gene flow likely has occurred across broad areas
and regionally discrete populations have not developed.
In a study of genetic structure among king eiders, samples were analyzed from Holarctic
breeding areas from northeast Russia east to Greenland, and wintering areas in the Pacific and
Atlantic sides of North America (Pearce et al., 2004). Results showed little genetic structuring
across the range, indicating that historical or current mobility among regions has not produced
discrete populations, particularly in western North America including the aggregations of
molting and wintering birds of Bristol Bay.
The population structure of Steller's eiders has been of great interest to assess the status and
prospects of the threatened Alaska-breeding component, particularly the small group breeding
near Barrow. In a study that genetically compared samples from breeding and wintering areas of
the Atlantic population (Russia-Norway) and the Pacific population (Russia-Alaska), some
differentiation was found between the two greater populations, but no sign of subpopulation
structure (Pearce et al., 2005). Similar to the situation with king eiders, genetic differentiation
could develop in the future if natural or anthropogenic factors provide isolation of breeding
groups.
Human Use
Bristol Bay waterfowl provide viewing, educational and research values, and harvest
opportunities to people in Alaska, Russia, Canada, the western U.S., and Mexico. As described
above, Bristol Bay is uniquely positioned to host a great diversity of waterfowl that breed,
winter, or pass through the region. This includes birds associated with Arctic breeding grounds,
the Aleutian Islands, the exceptionally productive tundra habitats of western Alaska, and birds
that winter as far south as Mexico and east to Chesapeake Bay. Though most of the common
waterfowl species of North America occur in the region, species such as swans, emperor geese,
and eiders are especially appreciated for their aesthetic, scientific and cultural values. The
richness and abundance of waterfowl from the region supports significant subsistence and
recreational harvests that are important economically and traditionally throughout western North
America.
Nonconsumptive Uses-
Birds that breed in or pass through Bristol Bay are subjects of wildlife viewing opportunities
throughout their annual cycles, from Alaska to Mexico. During spring and fall migration, and
during winter, Bristol Bay birds stop at many local, state, and federal parks and wildlife areas
featuring viewing and interpretive facilities. For some species, special community events revolve
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around the occurrence of migrating birds. For example, there are brant festivals in British
Columbia, Puget Sound, and northern California, and brant are an attraction with whale watchers
in Baja California. Such events and numerous local viewing programs are also common for
swans, aggregations of geese, and sandhill cranes that breed in the Bristol Bay lowlands.
Recreational Harvest-
Because Bristol Bay is an important breeding and staging area for ducks and geese, birds
produced in or supported by the region comprise a notable contribution to fall and winter
harvests in the Pacific Flyway from Alaska to Mexico. Waterfowl harvest data have been
collected by USFWS in Alaska since the early 1960s through a national mail questionnaire
survey (MQS) of federal duck stamp buyers. Species composition of the harvest was estimated
from duck wings and goose tails from a Parts Collection Survey (PCS). From 1971 through 1997
ADFG also conducted a mail questionnaire survey of hunters. Since 1998, waterfowl harvests
have been estimated by USFWS through the national Harvest Information Program (HIP), based
on a sample of all registered migratory bird hunters. The objective of all these surveys was to
produce reliable estimates of duck and goose harvests at the statewide level.
There are no reliable estimates of fall duck and goose harvests in the Bristol Bay region.
Although the ADFG survey and federal MQS surveys collected harvest data by regions, the
hunter sampling rates were not sufficient to provide more than a general sense of harvest across
the state. Bristol Bay was only part of a large sampling region named "Alaska Peninsula" that
extended west to Unimak Pass where ADFG data indicate that 4,000-5,000 ducks and 2,000
geese were harvested annually. Through the 1990s, this amounted to roughly 5% of Alaska's fall
duck harvest and 20-25% of the goose harvest. There are no recent regional harvest data.
Beyond southwest Alaska, birds that use Bristol Bay for some part of their life cycle contribute
an unknown portion of fall harvests in south-central and southeast Alaska, which typically
represent 60-70% of the statewide duck total (-70,000) and 30-40% of the statewide goose
harvest (-6,100) (USFWS, 2010a). There is no practical way to estimate the proportion of ducks
and geese from Bristol Bay harvested in the western states or other jurisdictions. Waterfowl
banding has not been sufficient in the Bristol Bay region to undertake analysis of harvest
derivation for the Pacific Flyway.
Since 2001, a fall tundra swan season has been open in Game Management Unit 17, with
registration permits required. The unit includes the Togiak, Wood, and Nushagak drainages
inland to Lake Clark. On average, fewer than 60 permits have been issued annually, and reported
harvest has been less than ten swans per year (ADFG, unpubl. data). A small number of tundra
swans harvested in Montana, Utah, and Nevada are derived from Bristol Bay.
Subsistence Harvest-
Harvest of waterfowl and other birds has been an important component of Bristol Bay's
traditional subsistence culture and economy. Although spring and summer hunting occurred
historically and continues, it was largely illegal after passage of the Migratory Bird Treaty Act of
1918. Fall and winter hunting, beginning September 1, has been allowed under federal and state
regulations. In 1997 the United States Senate ratified Protocols that amended the migratory bird
treaties with Canada and Mexico. This action authorized the USFWS to open a legal, regulated
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spring and summer subsistence season for migratory birds in Alaska during 2003, the first in
over 80 years. The Alaska Migratory Bird Co-Management Council (AMBCC) was formed,
composed of USFWS, ADF&G, and 12 regional representatives to establish a body of
subsistence hunting regulations and undertake the vital task of assessing spring and summer
subsistence harvests to meet intentions of the amended treaties.
During the 1980s, ADF&G worked with federal agencies, Alaska Native regional organizations,
and village governments to conduct subsistence harvest surveys of 151 rural communities in
Alaska and they characterized the levels and nature of migratory bird subsistence harvest in
Alaska, including Bristol Bay (Wolfe et al., 1990). In response to an intensive goose
conservation program on the Y-K Delta, cooperative village harvest surveys were initiated across
the region and eventually expanded to Bristol Bay villages that harvested the same goose
populations (Wong and Wentworth, 1999). These surveys, begun in 1995, gathered harvest data
by subregions, including Togiak, Dillingham, Iliamna, and villages in the Nushagak drainage and
on the Alaska Peninsula.
In a review of historical and recent harvest information, the AMBCC found the available data to
be insufficient to address management needs, so it designed a comprehensive statewide survey
protocol (Alaska Migratory Bird Co-Management Council, 2003), including Bristol Bay
communities on a rotating basis. Statewide subsistence harvest surveys were implemented
annually from 2004 to 2009, although they were not fully funded and implemented to full
performance standards.
In general, the seasonal harvest of migratory birds in the Bristol Bay region is most prevalent in
spring, with a few species taken in summer, and increased hunting during fall migration and
winter (Wolfe et al., 1990). The importance of the spring harvest reflects the abundance of birds
during spring migration and traditional need for fresh meat after winter supplies have been
depleted. For the Bristol Bay/Diamna region (not including Alaska Peninsula), Wolfe et al.
(1990) estimated annual harvests in the late 1980s of about 8,800 ducks, nearly 2,000 geese, 100
swans, and 1,100 waterfowl eggs. About 70% of households used birds; waterfowl provided 3.4
Ibs of meat per capita.
Conservation concerns in the late 1980s prompted a statewide assessment of harvests for Pacific
brant, emperor geese, and eider species (Wolfe and Paige, 1995). This study characterized
harvest of these species circa the early 1990s and presented harvest estimates by regions
(including North Bristol Bay, South Bristol Bay, and Lake Iliamna-Nushagak) and some
communities from ADFG and USFWS surveys during 1983-94. Harvest estimates for Bristol
Bay subregions provide totals for relative comparisons to other parts of the state, but also reflect
the seasonal availability of species within the region.
Brant harvest was highest (-300) in the North Bristol Bay area where spring staging is
concurrent with seal hunting (Schichnes and Chythlook, 1988), but was low in South Bristol Bay
because most brant bypass that part of the Alaska Peninsula. Brant were rare inland in the
Iliamna-Nushagak region. Emperor geese, which are closely tied to the coast in transit to and
from the Aleutian Islands, provided harvest for North (-300) and South (-200) Bristol Bay, but
were rare inland. Because of diminished numbers, all hunting of emperor geese was closed in
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1986. Eiders were harvested most in North Bristol Bay (-850) where king eiders are abundant
during spring and molt in Nushagak and Kvichak Bay, and common eiders stage during
migrations. Eider harvest was low in South Bristol Bay where king and common eiders are more
transient. Only common eiders (-250) were recorded for the Diamna-Nushagak region, probably
representing either inland migrant birds from lower Cook Inlet or inland households hunting in
coastal areas.
Harvest data from 1995-2005 were collected by subregion, including Dillingham, Nushagak
River and Hiamna subregions (Wentworth, 2007; Wong and Wentworth, 1999). Estimated
annual harvests included -10,000 ducks (mostly dabbling ducks, with -600 scoters, 100-200
eiders, and up to 190 harlequin ducks); 2,500-2,900 geese (up to 1,000 white-front, 800
Canada/cackling, 180-230 brant); up to 300 tundra swans; and fewer than 500 waterfowl eggs.
As part of the statewide AMBCC harvest survey program, Bristol Bay communities were
sampled during 2004 to 2008, including a Southwest Bristol Bay Subregion from Togiak south
to Port Heiden covering King Salmon and 20 villages in the Nushagak and Kvichak drainages,
and a Dillingham Subregion sampled every other year. Recent harvest estimates and seasonality
of species harvests are found in Naves (2010a, b). Approximate five-year average harvests for
the villages are: ducks - 10,200 (includes 600 scoters, 1,600 eiders); geese - 5,200 (includes
2,700 Canada, 1,300 white-fronts, 1,100 brant); swans - 270; and waterfowl eggs - 800.
Dillingham was surveyed on a rotational schedule (2005, 2007, 2008); results indicated
estimated harvests of 1,000-5,000 ducks, 500-800 geese, up to 50 swans, and less than 100
waterfowl eggs (Naves, 2010a; Naves, 201 Ob).
It is difficult to precisely characterize subsistence harvest of waterfowl in the Nushagak and
Kvichak watersheds and the greater Bristol Bay region because of subregional differences in the
geographic diversity and seasonal availability of waterfowl species and other wildlife resources,
as well as differences in cultural preferences and practices (Wright et al., 1985). In addition,
local and regional harvests of migratory birds vary considerably year-to-year because of
variations in bird abundance; timing, rates, and patterns of migrations; and seasonal hunting
conditions (Wolfe et al., 1990). Assessments of changes or potential changes to subsistence uses
of migratory birds will rely on updated status information for migratory bird populations, review
of data from the AMBCC community harvest survey program (Naves, 201 Ob), and compilation
of historic and current harvest data for subregions and communities (Behnke, 1982; Fall et al.,
2006; Fall et al., 1986; Schichnes and Chythlook, 1988; Schichnes and Chythlook, 1991).
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BALD EAGLES
Introduction
Bald eagles (Haliaeetus leucocephalus) range across North America, but are most abundant in
Alaska, where approximately half of the world population occurs. During the first half of the 20*
century in Alaska, the abundance of bald eagles and their attraction to human food sources made
them easy prey for bounty hunters (Hodges and Robards, 1982). Since even before the end of the
bounty days, however, they have been valued by Americans for their stately appearance, and
especially for their intrinsic association with wilderness (King, 2010). Bald eagles and their nests
receive extra protection above and beyond that afforded to other migratory birds by the
Migratory Bird Treaty Act (16 U.S.C. §§ 703-712). The Bald and Golden Eagle Protection Act
(16 U.S.C. 668-668c), enacted in 1940 and amended several times since then, establishes federal
responsibility for the protection of bald and golden eagles and requires consultation with the
USFWS to ensure activities do not adversely affect bald eagle populations. The Act prohibits
anyone, without a permit issued by the Secretary of the Interior, from "taking" bald eagles,
including their parts, nests, or eggs. It provides criminal penalties for persons who "take, possess,
sell, purchase, barter, offer to sell, purchase or barter, transport, export or import, at any time or
any manner, any bald eagle ... [or any golden eagle], alive or dead, or any part, nest, or egg
thereof."
A large apex predator, as well as an opportunistic scavenger, the bald eagles is a key species of
most of the regional food webs across coastal Alaska, from the Aleutians to Southeast. Bald
eagles were proposed as a management indicator species (MIS) for all National Forest lands in
Alaska (Sidle and Suring, 1986) and selected as a MIS for the Tongass National Forest (USFS,
2008). They are also included as a "vital sign"3 for long-term monitoring in the southwest Alaska
national parks (Bennett et al., 2006). The purpose of this paper is to provide a characterization of
bald eagles in the Nushagak and Kvichak watersheds, with particular emphasis on their
ecological relationships with MDNs.
Habitat
Bald eagles are well-known for their association with water. In the Pacific Northwest most nests
are within 1.6 kilometers of large waterbodies (Anthony et al., 1982) and in Alaska they are
almost always found within 200 meters of a stream, lake, or ocean shoreline (Hodges and
Robards, 1982; Stalmaster, 1987; Swaim, personal communication). In the Bristol Bay
watershed they inhabit the spruce and mixed spruce/broadleaf forests along major rivers,
streams, and lakes of the Bristol Bay Lowlands, upper Alaska Peninsula, and Lime Hills eco-
regions (ADF&G, 2006), as well as coastal areas of Bristol Bay.
Throughout most of their range, bald eagles nest in forested habitat associated with riparian and
beach areas (Buehler, 2000; Stalmaster, 1987). The significance of shoreline nest sites appears to
be strongly related to foraging opportunities these areas provide (Armstrong, 2010). Commonly
3 Defined as "a subset of physical, chemical, and biological elements and processes of park ecosystems that are
selected to represent the overall health or condition of park resources, known or hypothesized effects of stressors, or
elements that have important human values."
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used foraging areas are open sites where prey or carrion can be seen and accessed by these large
birds. These are often areas that provide prey aggregations, accessible to many bald eagles at
once. In Alaska, these areas include lakes, rivers, oceans, and their shorelines, beaches, and bars
(Stalmaster, 1987). On the Kenai NWR, nests are located near clear, relatively shallow streams
with spring and fall fish runs (Bangs et al., 1982a). While bald eagle nest sites are positively
correlated with water, they tend to be negatively correlated with lands impacted by timber
harvest and other human uses (Anthony and Isaacs, 1989; Livingston et al., 1990).
In Alaska, where suitable trees are present, bald eagles build their enormous nests near the tops
of live trees, typically in one of the tallest of the stand (Bangs et al., 1982a; Ritchie, 1982;
Stalmaster, 1987). Where large trees are absent, they nest on ridges, hillsides, small islets, or sea
stacks (Savage, personal communication; Suring, 2010). Around Lake Clark and its drainages,
most nests are reported to be in balsam poplar and spruce trees (Wright, 2010). Monitoring in
LCNPP since 1992 identified black cottonwood as the predominant substrate for coastal nests;
interior nests were more equally divided between cottonwood and spruce (from Witter and
Mangipane 2011, in preparation). Of the 165 known bald eagle nest sites on TogiakNWR, most
(-86%) are in balsam poplar trees; 8% are in spruce and 6% are located on the ground (Swaim,
personal communication).
On Kodiak NWR, one-third of bald eagle nest sites are not located in trees (Zwiefelhofer, 2007).
In the Bristol Bay area, at least one ground nest has been noted on an islet in Tikchik Lake
(Wright, 2010) and one on an islet in Katmai NPP (Savage, personal communication). At least
two ground nests have been documented on Flat Island, in interior LCNPP, although each was
occupied for only one year (one was successful and the other failed). Also, there are multiple
years of data on two coastal ground nests near Tuxedni Bay in LCNPP. A third coastal ground
nest was found near Difficult Creek in 2011 and was successful that year (Witter, personal
communication). Besides nest platforms, bald eagles need perches and, often, communal roosts.
In Alaska, bald eagles commonly perch in large spruce or cottonwood trees, often with a good
view of foraging waters. Bald eagles often perch for 90% of the daylight hours (Stalmaster and
Gessaman, 1984), with these locations serving as sites for resting/loafing, foraging/hunting,
feeding, look-outs/sentry posts, displaying (territoriality), and thermal regulation (heating or
cooling) (Stalmaster, 1987).
The extent of communal roosting is not well-known for the Nushagak and Kvichak watersheds.
In general, though, breeding bald eagles roost at the nest tree or on other large trees within their
territory. Non-breeding bald eagles often retire to communal roosts at night, usually in large
spruce or cottonwoods trees. They may be near a foraging area, but are not as closely associated
with shorelines as perches and nest trees (Stalmaster, 1987). Non-breeding bald eagles also
regularly feed, rest, and roost along gravel bars and gravel shorelines (Hansen et al., 1984). On
the Naknek River in March and April, communal roosts are located on hillsides with shrubs and
some balsam poplars (Savage, personal communication). These are often on south-facing slopes
that overlook the river.
An abundant, readily available food supply in conjunction with one or more suitable night roost
sites is the primary characteristic of winter habitat. The majority of wintering bald eagles are
found near open water, where they feed on fish, marine invertebrates, and waterfowl and
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seabirds, often taking dead, crippled, or otherwise vulnerable animals (Buehler, 2000). Over-
wintering on the breeding grounds may provide a competitive edge in territory selection and with
early initiation of nesting (Ritchie and Ambrose, 1987).
The proportion of bald eagles that over-winter in the Nushagak and Kvichak watersheds is not
well-understood, and potential links between over-wintering and open water or winter prey
accessibility remain to be studied. Ritchie and Ambrose (1987) reported that records of bald
eagles over-wintering in northern boreal forests are rare and that one reason for this fact is that
water bodies are frozen. They observed bald eagles in winter along the Tanana River in Interior
Alaska, where open water probably provides access to spawning salmon and waterfowl. Bald
eagles often congregate along the ice/open water interface on the Naknek River, where wintering
common mergansers (Mergus merganser) and common goldeneyes (Bucephala clanguld) are
often found. The 1986 to 2010 King Salmon-Naknek Christmas bird counts reported from zero
to up to 48 adult bald eagles (average 18) (Savage, personal communication). Some over-
wintering occurs in Dillingham and the surrounding area, but in much lower densities than found
in summer. Some of those over-wintering eagles obtain human garbage at the city dump (Swaim,
personal communication).
Food Habits
Diet- Bald eagles are primarily fish eaters (Armstrong, 2010). They do, however, have a variable
diet that can include birds, mammals, and crustaceans, and, as noted above, even human garbage
(Anthony et al., 1999; Knight and Knight, 1983) (also see Stalmaster 1987 and Armstrong 2010
for summaries).
Food habits vary spatially according to specific prey availability and abundance at the site. Bald
eagles nesting near and foraging at seabird colonies during the summer may take primarily bird
prey (DeGange and Nelson, 1982). In the Pacific Northwest, diet varied among sites, with bird
prey items found under nests generally out-numbering fish items (Knight et al., 1990).4 Birds
likely also out-number fish as prey at some sites on Togiak NWR, including Cape Peirce and
Cape Newenham, which support high densities of breeding seabirds (Swaim, personal
communication).
The diet of bald eagles tends to vary temporally, as well, depending on prey availability and
abundance. Nesting bald eagles rely primarily on the availability of salmon resources (Hansen,
1987). Inland bald eagles whose nests are close to spawning streams have higher nesting success
than those with more distant nests (Gerrard et al., 1975). When salmon resources are scarce
during late winter and early spring, coastal populations of bald eagles often shift their diet to
birds (Isleib, 2010; Wright and Schempf, 2010). In one study, birds averaged nearly 20% of
stomach contents by volume over the course of a year, but could range up to 86% and be
especially high during the colder months (Imler and Kalmbach, 1955). In other areas,
mammalian prey may be utilized in winter, because it is as available, or more available, than
birds. For example, on the Kenai NWR, bald eagles may seasonally shift from a diet of primarily
fish to snowshoe hare (Lepus americanus) or mammalian carrion (Bangs et al., 1982a).
4 Note that fish and other soft-boned or bodied prey may be commonly under-counted in both stomach-content and
under-nest methodologies (Ritchie 1982, Knight et al. 1990).
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Specific information regarding bald eagle diet variability in the Nushagak and Kvichak
watersheds is generally not available. Eagles in the Bristol Bay watershed area eat all five
species of Pacific salmon (Savage, personal communication). Bald eagles in the winter along the
Naknek River have been observed to take small fish, which may include eulachon (Thaleichthys
pacificus) (Savage, personal communication). Pacific herring (Clupea pallasi) may also be an
important resource for bald eagles in the Togiak area. In early spring Savage has also observed
bald eagles catching large rainbow trout (Oncorhynchus mykiss). Bald eagles in Bristol Bay may
also scavenge dead marine mammals (Savage, personal communication).
Independent of prey availability, energy requirements may influence prey selection at some level
as well. During the breeding season many bald eagles choose large fish over small fish (Jenkins
and Jackman, 1994). Diets of nesting bald eagles are much more variable than those of non-
breeders (Hansen et al., 1984; Hansen et al., 1986). Non-breeders are able to range farther for
preferred food items (e.g., in late fall birds may leave the Chilkat Valley in southeast Alaska to
go to British Columbia and Washington, where salmon may still be available). Feeding of young
is, as Stalmaster (1987) says, "an enormous chore" and breeders may exploit a variety of food
resources within their home range.
Significance ofMDNs- Fresh salmon and salmon carcasses provide an ideal food resource for
bald eagles, because they are large fish that become available in great numbers when they enter
shallow water to spawn. Shallow water increases the likelihood that living fish will be available
to bald eagles because the limited depth of water brings fish closer to the surface (Livingston et
al., 1990). Returning salmon die after spawning in natal streams, providing a significant seasonal
pulse of MDNs, including nitrogen and phosphorous, to the generally oligotrophic streams and
lakes of northern Pacific watersheds (Hilderbrand et al., 2004; Naiman et al., 2002b; Willson et
al., 1998). Spawned-out salmon carcasses accumulate on stream banks, river bars, lake and ocean
shores, and tidal flats (Armstrong, 2010). Although spawned-out salmon are low in fat and
considered a relatively low-energy food source (Christie and Reimchen, 2005), their large size,
availability, sheer numbers, and other factors (even cold air temperatures, which can increase the
efficiency of digestion of some prey (Stalmaster and Gessaman, 1982)), contribute to their value.
Salmon are approximately 79% edible flesh, compared to 71% for hares and 68% for ducks
(Stalmaster, 1981). Although metabolizable energy is lower for salmon than for hares or ducks,
their greater size means that a bald eagle would require only 57 salmon in one year, compared to
87 hares, or 135 ducks (Stalmaster, 1981; Stalmaster and Gessaman, 1982).
Armstrong (2010) reported that several studies have correlated bald eagle abundance with the
abundance of spawned-out salmon. Simply put, bald eagles in southeast Alaska, the Kenai NWR,
and many other parts of their range likely depend on salmon (Armstrong, 2010; Bangs et al.,
1982b). The nature of the relationship that bald eagles have with salmon, however, is complex
(Hansen et al., 1986). The variable nature of salmon (and other bald eagle food sources)
apparently causes bald eagles to be limited by food availability (Stalmaster and Gessaman,
1984). The abundance of salmon affects not only bald eagle population and distribution, but also
breeding and behavior. Bald eagles, in turn, affect the riparian ecosystem and other areas where
they may range by distributing the MDNs in their excretions (Gende et al., 2002).
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Bald eagles in Alaska also congregate to feed on other species of anadromous and shallow-water
spawning fish, particularly Dolly Varden (Salvelinus malma malma\ Pacific herring, and
eulachon (Armstrong, 2010). Armstrong (2010) also summarized the importance of Pacific sand
lance (Ammodytes hexapterus) to bald eagles and other marine-associated birds and mammals.
Forasins Methods- Bald eagles are opportunistic foragers that exhibit rather complex social
feeding behaviors. They use a variety of methods to obtain food, including active hunting and
killing, scavenging of carcasses, and theft (pirating or kleptoparasitizing) from other eagles or
species (Stalmaster, 1987). They are visual predators that locate their prey by sight. Foraging
methods chosen by bald eagles vary according to both complex relationships among other eagles
and with other predators or competitors, as well as seasonal variability of food sources. Bald
eagles may search for prey themselves or follow other birds, or even mammals, to a concentrated
food source (Harmata, 1984; Knight and Knight, 1983; McClelland et al., 1982). Also, as
summarized in Armstrong (2010), bald eagles will not only steal fish or force other predators
away from fish, but will also exploit fish injured or driven to the surface by others, and scavenge
crippled or dead fish or fish parts left by other predators such as humans or bears. Bald eagles in
the Chilkat Valley of southeast Alaska typically competed amongst themselves for salmon
(Hansen et al., 1984). Dominance in bald eagles may be based on several conditions, but often
includes age and size (Garcelon, 1990; Stalmaster, 1987).
Salmon returning to spawning streams are a relatively easy food source for bald eagles, because
the fish are found either in shallow waters, swimming or floating near the surface, or washed up
or stranded on streambanks (McClelland and McClelland, 1986). Besides their size, abundance,
and availability in shallow water, other unique aspects of salmon life history may contribute to
their importance to bald eagles. For example, large numbers may be frozen into river ice in the
winter, becoming available as food sources again in spring (Hansen et al., 1984). Brown bear
pull salmon from holes too deep for eagles to access, often transporting and discarding portions
of carcasses to other locations, where eagles then scavenge them (Armstrong, 2010). Armstrong
(2010) stressed what he believes to be a particularly important relationship with bears (Ursus
spp.), which scoop salmon out of deep pools, where they may be inaccessible to bald eagles, and
then often eat only the brains and eggs, leaving a significant proportion of the salmon flesh for
the birds.
According to Stalmaster (1987), bald eagles, in general, appear to prefer stealing of food over
scavenging, and scavenging over hunting. Hansen et al. (1984) observed higher frequencies of
stealing over scavenging of salmon carcasses in the Chilkat Valley in southeast Alaska, even
though the cost and benefits of both may be equal. However, others have found that, when food
is scarce, bald eagles will choose scavenging over stealing, if both methods are available (Knight
and Skagen, 1988). Wright and Schempf (2010) state that during seasons of food scarcity,
feeding strategy may switch to more active hunting, particularly of large gulls and waterfowl,
and some bald eagles may steal ducks from hunters or scavenge in garbage dumps.
Behavior
Territoriality- During the nesting period, breeding bald eagles occupy and defend territories
(Mahaffy and Frenzel, 1987). A territory includes the active nest and may include one or more
inactive nests, which the eagles may maintain, even when not in use for nesting in a given year
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(Hansen et al., 1984). They maintain the same territory year after year, using the same nest or an
alternate nest within the same territory (Steidl et al., 1997).
The defended territory contains not only the nest trees, but also favored perches and roost(s).
Territories have been reported to range from 0.2-4.2 km2 (Garrett et al., 1993), but size varies
according to site and other parameters (Stalmaster et al., 1985). The territory is within a larger
home range. Bald eagles, unlike many other birds, do not necessarily use a territory to
monopolize food, but commonly range out of their territory to obtain food communally at a site
where it is abundant (Stalmaster, 1987).
In any given year, not all territories will be occupied, and not all occupants will attempt to
reproduce (Stalmaster, 1987). During the non-breeding season, or if not breeding, bald eagles
generally do not defend territories (Armstrong, 2010), although a pair may remain close to their
nest or return to their territory regularly over the winter (Gende, 2010). Information is not
currently available on characteristics of bald eagle territories (e.g., size, use patterns, average
number of nests, variability according to habitat type, etc.) in the Nushagak or Kvichak
watersheds.
Flocking- When a food resource is concentrated, bald eagles will often forage in large flocks
(Stalmaster, 1987). This is true not only for scavenging and stealing, as can occur when carrion
is present, but also for hunting, when there are large aggregations of forage fish like eulachon or
sandlance (Stalmaster, 1987; Stalmaster and Gessaman, 1984; Willson and Armstrong, 1998).
In the winter, when food availability is limited (e.g., by iced-over rivers or limited daylight), bald
eagles aggregate in large flocks and become very aggressive, often pirating food from other
birds. An available food source will initially draw bald eagles to a site, and the presence of large
numbers of bald eagles will attract additional birds.
At night, non-breeding and wintering bald eagles may congregate in communal roost areas
(Hansen et al., 1980). The same roost areas are used for several years. Roosts are often in
locations that are protected from the wind by vegetation or terrain, providing a favorable thermal
environment. The use of these protected sites helps minimize the energy stress encountered by
wintering birds. Communal roosting may also assist bald eagles in finding food. The use of
communal roosts is poorly documented in Alaska, however (USFWS, 2009a).
Migration and Local Movements- The extent to which bald eagles are migratory or non-
migratory varies with breeding site and the severity of its climate (particularly in winter),
whether the individual is adult or sub-adult, and year-round food availability (Buehler, 2000).
Bald eagles breeding in coastal Alaska typically remain in the vicinity of their nest sites
throughout the year. For example, the southeast Alaska adult population is mostly non-
migratory, remaining in its rainforest habitat year-round (Sidle et al., 1986). Adults in Aleutian
Island populations are generally resident as well (Sherrod et al., 1976). Wintering grounds for
migratory Alaska bald eagles are not well understood, but it is suspected that Interior bald eagles
winter in the Intermountain West and Pacific Northwest (Ritchie and Ambrose, 1996).
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Diurnal and tidal cycles affect the daily activity patterns of fish, as well as enhancing or
inhibiting hunting conditions for bald eagles (Hansen et al., 1986).Variations in these daily
patterns lead to local movements of bald eagles.
Even though the population of bald eagles in southeast Alaska is non-migratory, individuals will
leave their territories to visit foraging areas for several days at a time (Kralovec, 1994).
Southeast pairs also return to their breeding territory periodically over the course of the winter.
Local movement patterns, the extent of over-wintering and migration, and how each may vary
with age, food availability, or other factors are poorly understood for the Bristol Bay watershed.
It is known that at least some adults and sub-adults over-winter in Bristol Bay (Wright, 2010).
Interspecies Interactions
Prey availability has a strong influence on bald eagle reproduction, habitat use and territorial
behavior in Alaska. The studies of Hansen et al. (1984) suggest that salmon availability in spring
is tightly correlated with if and when adult bald eagles will lay eggs in a given year, although this
has not been studied in the Bristol Bay watershed. Bald eagles preferentially select nest sites near
stable food supplies (e.g., salmon in the Chilkat Valley). These studies also indicate that food
(salmon) availability during the nesting period regulates the survival rate of offspring. Hansen et
al. (1984) further determined that, while breeding adults commonly defended feeding territories,
they did not do so when salmon became overabundant. Fall and winter habitat use is often
correlated with salmon availability, too. Hansen et al. (1984) clearly demonstrated this in the
Chilkat Valley and is likely true in Bristol Bay as well.
Bald eagles defend vulnerable young against predators (Stinson et al., 2001). Otherwise they do
not tend to be as aggressive with other species as they are with other eagles, with which
antagonistic interactions regularly occur during feeding and territory defense. One exception is
osprey (Pandion haliaetus), which bald eagles commonly keep from nesting nearby (Stalmaster,
1987), although this behavior has not been investigated in the Bristol Bay area. Great horned
owls (Bubo virgimanus\ which nest earlier than bald eagles, and osprey, which nest later, each
may occupy bald eagle nests in southwest Alaska (Savage, personal communication). Bald
eagles steal and scavenge food from a variety of bird and wildlife species, including river otter
(Lutra canadensis) and sea otter and many others (Stalmaster, 1987).
Breeding, Productivity, and Mortality
Breeding- As with other birds, the timing of bald eagle nesting varies by latitude; in Alaska it
begins with courtship and nest building as early as February and ends when the young fledge in
late August to early September. In the Bristol Bay watershed, initiation may not be until mid- to
late March (Savage, personal communication). The young are attended by the adults near the
nest for several weeks after fledging (Buehler, 2000). A pair's territory frequently contains more
than one nest (Haines and Pollock, 1998), although a pair uses only one nest in a given breeding
year and does not necessarily breed every year (Hansen and Hodges, 1985). The territory (and
pair bond) is usually maintained for life (Jenkins and Jackman, 1993).
Whether or not bald eagle pairs breed in a given year and how early they may initiate nesting in a
given year appear to be related to food availability (Hansen, 1987), particularly in spring
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(Hansen et al., 1984). These studies suggest that there may be a natural long-term population
cycle, at least in southeast Alaska's Chilkat Valley, resulting from a saturated breeding habitat
and surplus of non-breeders, who then compete for food and cause productivity to decline. The
decline may result in less recruitment into the non-breeding population, less competition, and
ultimately increased productivity. Annual occupancy rates at known nest sites within Togiak
NWR varied from 45 to 88% between 1986 and 2006 (Swaim, personal communication). The
lowest occupancy rate occurred in 2006, when spring break-up was particularly late. Occupancy
rates, relationships with food availability and seasonal variability, and other details of bald eagle
breeding are not well understood for the Nushagak and Kvichak watersheds.
Even well-established nesting bald eagles are highly sensitive to disturbance, particularly during
the phases of early courtship and territory establishment, incubation, and the first two weeks after
hatching (Buehler, 2000). As with many birds, a constant level of nest attendance is required
during incubation and brooding. Bald eagles in newly established territories are highly sensitive
to disturbance and prone to abandon nest sites during the courtship and nest-building stage
(Gende et al., 1998). Occasionally, a pair will establish and maintain a territory in urban or semi-
urban areas where some, usually predictable level of, disturbance already occurs (Zwiefelhofer,
personal communication).
Females are larger than males (Buehler, 2000). Both sexes incubate eggs, brood young, and
deliver prey to chicks, sharing the duties more than many other raptor species, although females
still undertake these tasks a greater percentage of the time than males (Cain, 1985).
Productivity and Survivorship- Productivity varies among sites according to prey abundance and
availability, habitat quality, weather, breeding-season length, nesting density, and human
disturbance (Gende et al., 1997; Hansen, 1987; Savage, 1997; Steidl et al., 1997). Measures of
annual productivity include number and percentage of occupied nests, successful nests, and
young produced. Some local information about bald eagle productivity in the Bristol Bay area
can be gleaned from National Park Service nest surveys in Lake Clark and Katmai NPPs.
Average nest success (percentage of occupied nests that produced at least one young) for interior
LCNPP was about 55% between 1992 and 2009, falling to 48% in 2010 (Mangipane, 2010).
Nest success for the Naknek Lake and major associated drainage areas of Katmai NPP varied
from 31 to 65% in the years 1992 through 1997, although the sample size was relatively small
(Savage, 1997).
Dates vary, but generally egg-laying begins in mid- to late April in Alaska (Savage, personal
communication; Swaim, personal communication). Clutch sizes range from one to three eggs.
Successful pairs usually raise one or two young, or rarely, three per nest (Table 9). Bald eaglets
make their first unsteady flights about 10 to 12 weeks after hatching, and fledge (leave their
nests) within a few days after that first flight. The time between egg-laying and fledging is
approximately four months. However, young birds usually remain in the vicinity of the nest for
several weeks after fledging and depend on their parents for food until they disperse from the
nesting territory approximately six weeks later. The entire breeding cycle, from initial activity at
a nest through the period of fledgling dependency, is about six months (Buehler, 2000).
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Numbers of young produced overall in the Nushagak and Kvichak watersheds is unknown.
Annually, Alaska bald eagles may produce roughly 4,200 fledglings, although that figure varies
considerably according to site and year (Schempf, 1989). In Alaska, both inter-annual and site-
specific variability in productivity can be significant (Schempf, 1989) and neither has been
comprehensively studied for the Nushagak and Kvichak watersheds. Productivity appears to be
most commonly related to site-specific habitat features and prey (fish) availability in early spring
during egg-laying and incubation (Anthony, 2001; Gende et al., 1997; Steidl et al., 1997).
Availability of fish increases survivorship of bald eagle offspring, and therefore can cause bald
eagle productivity to fluctuate widely (Hansen et al., 1984). Variability in food availability
appears to be the cause of variability in fledging rates in southeast Alaska (Hansen, 1987).
Productivity can be affected by human disturbance, as well (Fraser and Anthony, 2010;
Stalmaster, 1987).
Mortality- Full-grown bald eagles have few natural enemies, and the most frequently reported
causes of premature adult bald eagle mortality are human-related (Franson et al., 1995; Harmata
et al., 1999; Stalmaster, 1987). Shooting, electrocution, trapping, and collisions cause about two-
thirds of reported deaths. Bald eagles also die from ingesting pesticides and contaminated carrion
used for predator control. Historically, bald eagles experience decreased reproduction and
survival from both intentional and unintentional effects of a wide range of pesticides and
environmental contaminants (Buehler, 2000). Poisoning from a wide variety of sources
accounted for 16% of all deaths in bald eagles necropsied between 1963 and 1994 at the National
Wildlife Health Center (Franson et al., 1995).
Top-level predators, such as bald eagles, are believed to be especially vulnerable to many
contaminants and can be used as sentinel species for contaminated areas (Holl and Cairns, 1995;
Welch, 1994). Eggs from bald eagles in the Aleutian Islands, for example, contained elevated
levels of organochlorine pesticides, but concentrations of these contaminants and mercury were
significantly higher in eggs from Kiska Island than in eggs from the other islands (Anthony et al.,
1999). In contrast, polychlorinated biphenyl (PCB) concentrations were higher in eggs from
Adak, Amchitka, and Kiska islands than in those from Tanaga Island. The most likely source of
these contaminants in bald eagles was their diets, which were spatially and temporally variable.
A similar study found that contaminant concentrations in Aleutian bald eagle eggs were
influenced more by point sources of contaminants and geographic location than by the trophic
status of eagles among the different islands (Anthony et al., 2007).
Mean cadmium, chromium, mercury, and selenium concentrations in bald eagle tissues from
Adak Island were consistent with levels observed in other avian studies and were below toxic
thresholds (Stout and Trust, 2002). However, elevated concentrations of chromium and mercury
in some individuals may warrant concern. Furthermore, although mean PCB and p,p'-
dichlorodiphenyldichloroethylene (DDE) concentrations were below acute toxic thresholds, they
were surprisingly high, given Adak Island's remote location.
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10. in
Site and year(s)
Average number of young raised
to neara fledging per successful
nest
Source
Interior LCNPP (1992-2011)
Coastal LCNPP(1992-2011)
Pacific Coast of the Alaska
Peninsula (1989-1995)
Port Moller( 1976)
Togiak NWR (1986-1988)
Togiak NWR (1986-2006)
1.00-1.87 '
1.09-1.82'
1.55-1.71
1.90
0.95-1.90
1.33-2.00 '
Witter and Mangipane (2011, in
preparation)
Witter and Mangipane (2011, in
preparation)
(Dewhurst, 1996)
R. Gill, unpublished data
reported in Wright (2010)
L Hotchkiss and D. Campbell,
unpublished data, as reported in
Wright (2010)
M. Swaim, personal
communication
Togiak NWR (2006)
Togiak NWR (1999-2005)
KatmaiNPP (1976-1979)
Katmai NPP (1992-1993)
Kodiak NWR (1963 and 2002)
Petersburg area (1967-1969)
Gulkana River (1989-1994)
Copper River (1989-1994)
Chilkat Valley (1979-1983)
Prince of Wales Island (1991-
1993)
1.72
1.62
1.2-1.8 b
1.45-1.67
1.66
1.50-1.65
1.29-1.65
1.34-1.64
1.32
1.10
(MacDonald, 2006)
(MacDonald, 2006)
W. Toyer, unpublished data
reported in Wright (2010)
(Savage, 1993)
(Zwiefelhofer, 2007)
(Corr, 1974)
(Steidl et al., 1997)
(Steidl et al., 1997)
(Hansenetal., 1984)
(Anthony, 2001)
a Nests are normally surveyed just before fledging to assess success and it is assumed that nests are successful if
young are observed. This is because once young fledge and leave the nest it is impossible to determine if they
survived. b Successful nests with three young have been reported in Lake Clark and Katmai NPPs and Port Moller
(L. Witter, personal communication, and as reported in Wright [2010]). Also, 3% of nests on Togiak NWR between
1986 and 2006 had three young (M. Swaim, personal communication).
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Non-human causes of mortality include starvation, fights with other bald eagles, and incidental
diseases and infections (Stalmaster, 1987). When food is limited, mortality rates are probably
higher among sub-adult than adult bald eagles (Stalmaster and Gessaman, 1984). Other causes of
mortality include loss of nests (eggs and nestlings) to spring storms, parental desertion or death,
and predation by gulls, black bears (Ursus americamis), and other predators (Stalmaster, 1987).
Although eggs tend to have a higher mortality rate than nestlings, nestlings also kill each other in
fights, die from starvation when more aggressive nest mates receive the majority of feedings
from the parent, and fall prematurely from nest trees.
Population, Distribution, and Abundance
Bald eagles are one of the most abundant raptors in Alaska, with a population estimated at
>58,000 (Hodges, 2011). Most Alaskan bald eagles occur in the vicinity of the southern coast
(from Dixon Entrance to Bristol Bay) and secondarily along interior rivers and lakes (Schempf,
1989). An estimated 2,775 adult bald eagles were present along the Alaska Peninsula Gulf Coast
in 2005 (Savage and Hodges, 2006).
Surveys of nests and calculations of nest densities and occupancy rates are commonly conducted
to contribute to bald eagle population information, although nesting rates have considerable
temporal and spatial variability. Nesting density is considered to be generally correlated with
food availability (Dzus and Gerrard, 1993), although density of breeding bald eagles in
Saskatchewan was found to be correlated with mean April temperatures (Leighton et al., 1979).
Densities of nests in inland river areas of southeast Alaska are highly variable among sites and
years. This may be correlated with food abundance and weather conditions (Hodges, 1979).
Nests in the Susitna watershed, though, are thought to be more uniformly distributed (Ritchie and
Ambrose, 1996). For Interior Alaska populations, Ritchie and Ambrose (1996) surmise that
densities are greatest in areas adjacent to abundant coastal populations and where weather is
somewhat milder and prey more seasonally accessible and diverse.
Bald eagle densities have been extensively studied in southeast Alaska. A review of several
nesting density investigations in various southeast Alaska locations revealed densities ranging
from 0.33 to 0.50 (perhaps greater on Admiralty Island) active nests per kilometer of coastline
and 0.25 to 0.38 active nests per river kilometer (Hansen et al., 1984; Hodges, 1979; Robards
and King, 1966). There were 0.01 to 0.08 active nests per river kilometer at an Interior location
(Gulkana River) (Byrne et al., 1983). Population and nesting density is also high on Kodiak
NWR, where almost 1,000 nests were located within an area of about 8,000 square kilometers in
2002 (Zwiefelhofer, 2007).
A comprehensive survey has not been published for bald eagles or their nests in the Bristol Bay
watershed. The data that we do have appears to indicate that nest density may be almost as high
in portions of the region as anywhere outside of the highest known densities in southeast Alaska
and Kodiak. The USFWS Bald Eagle Nest Database has accumulated approximately 230 nest
records for the study area (Table 10). Approximately one-quarter (61) of those records, however,
are from the 1970s and 1980s, and those nests may not persist on the landscape. The remaining
169 records were collected between 2003 and 2006. In 2006, a fixed-wing survey of adult bald
eagles was conducted by the USFWS along main-stem portions of some Alaskan rivers. During
that survey, 50 bald eagle nests were incidentally recorded along portions of the Nushagak,
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Mulchatna, and Kvichak Rivers. Of those 50, approximately one-half (24), were identified as
active. Database records for 2004 and 2005 are from a project contractor survey (not flown for
the USFWS Database) that was conducted along the north side of Lake Iliamna. The 2004 and
2005 surveys recorded 75 total nests in this area (Lewis, personal communication). This appears
to be a relatively high nesting density, although we do not know which or how many of those
nests were active, nor therefore, the density of active territories in this area. Data from 2003 are
for only three nests, one active and one empty nest in the lower Nushagak drainage and one of
unknown status on an islet off the north shore of Iliamna Lake.
11. of surveys for in
Survey Survey dates and results
USFWS bald eagle nest surveys (recorded in USFWS bald
eagle nest database)
61 nest records 169 nest records
Nushagak and Mulchatna Rivers survey by USFWS 2006
50 nest records (24
active)
North side of Lake Iliamna survey by contractor 2004-2005
75 nest records
Regarding numbers of individual bald eagles observed, some site-specific surveys have been
conducted in portions of southwest Alaska. For example, summer activity surveys for Katmai
NPP identified between 50 and 87 individuals in the Naknek Lake drainage between 1991 and
1997 (Savage, 1997). Systematic efforts have not been made to identify fall bald eagle
congregation sites in the Bristol Bay area (Wright, 2010). Such sites are known to exist in
surrounding areas (e.g., Port Moller, Savonoski River), however, and are believed to be related to
late-spawning sockeye salmon, fall runs of coho salmon, and fall-staging waterfowl. While bald
eagle densities are undeniably greatest overall in southeast Alaska, salmon also appear to be a
major driving force for the Bristol Bay watershed population of bald eagles, so some
comparisons may be inferred. In the Chilkat Valley, fall and winter bald eagle densities in
habitats adjacent to foraging areas may be ten times those of the same habitats (e.g., gravel bars,
cottonwood stands) located distant from food sources (Hansen et al., 1984).
Human Use
Historically, bald eagles have been important to Native Americans and continue to be so at
present. Bald eagle parts have been of particular importance for rituals and many other spiritually
related uses (Stalmaster, 1987). The Bald and Golden Eagle Protection Act exempts Native
Americans from the prohibition against purposeful take, although a permit is required.
Other humans have not used bald eagles historically for any significant intrinsic purpose.
Humans, however, are the greatest source of the bird's mortality, both directly and indirectly. At
least 128,000 birds, and probably many more, were taken during the bounty years (1917 to 1952)
in Alaska. Bald (and golden) eagles are now protected by law in the United States, with only a
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minimal number of permits for indirect take (incidental to an otherwise lawful activity) allowed.
Take is authorized only when it is consistent with the goal of maintaining stable or increasing
bald eagle populations.
Despite legal protection, illegal direct take still occurs, most commonly when bald eagles are
shot, trapped, or poisoned based on a belief that the birds prey on human-valued resources.
Unpermitted indirect take is probably the greatest source of bald eagle mortality today. Leading
causes of indirect take include pesticides, entanglement in fishing or trapping gear, collisions
with power lines or buildings, ingestion of poisoned prey, plastics or lead shot, and disturbance
or loss of nesting habitat (Buehler, 2000).
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SHOREBIRDS
Introduction
Shorebirds are a diverse group, with members occurring on every continent and in all habitats
ranging from sea level to the highest mountains. They generally are associated with water,
particularly intertidal and estuarine environments, and thus are fairly visible to humans. Due to
their broad geographic distributions, their seasonal migrations are remarkable, regularly spanning
continents and frequently hemispheres. Several species engage in long, nonstop flight, but most
rely on a series of sites where they stop to "refuel" for subsequent legs of their migrations.
Alaska intertidal areas, particularly Bristol Bay estuaries, serve two functions in this regard.
First, during late summer through autumn, the majority of the shorebird population that nests in
western Alaska moves to the benthic-rich intertidal communities of Bristol Bay, where ample
food supports them while they complete their molt and fatten for autumn migrations. Winter
destinations include sites throughout the Americas, the Central Pacific, and Australasia.
Secondly, during spring, hundreds of thousands of shorebirds that staged on the Copper River
Delta and estuaries of Cook Inlet migrate to their western Alaska breeding grounds, through a
broad lowland corridor (the Lake Iliamna corridor) at the base of the Alaska Peninsula, linking
Kamishak Bay in lower Cook Inlet to upper Bristol Bay. In most years, the migration through
this corridor is direct, but in years with late spring or adverse weather conditions, birds stop in
large numbers at Bristol Bay estuaries until conditions improve farther west (Gibson, 1967; Gill
and Handel, 1981; Gill and Tibbitts, 1999). Two major estuaries in the area, Nushagak and
Kvichak bays, have been recognized as Western Hemisphere Shorebird Reserve Network sites
(Western Hemisphere Shorebird Reserve Network, 2011).
Over 70% of the shorebird species or subspecies (30 of 41) that regularly occur in Alaska each
year (Alaska Shorebird Group, 2008) can be found in the Bristol Bay watershed; 21 of these 30
(70%) regularly nest there (Table 12). Shorebird populations worldwide are showing steady
declines (Stroud et al., 2006), with causes most often attributed to loss or alteration of habitats
and environmental contamination. Fourteen species that regularly occur in the Bristol Bay
watershed have been ranked by the Alaska Shorebird Working Group (2008) as being of high
conservation concern.
The Bristol Bay region has had a long history of studies that reported in part on shorebirds.
Several of these studies date to the late 19 and early 20 centuries, but few shorebird-specific
studies emanated from this region until the initiation of the Outer Continental Shelf
Environmental Assessment Program (OCSEAP) in coastal Alaska in the early 1970s (Arneson,
1978; Gill et al., 1977; Gill et al., 1978). At that time, there was also increased interest in
declining populations of waterfowl throughout Alaska and considerable information on
shorebirds was collected in conjunction with waterfowl studies, such as annual spring and fall
emperor goose aerial surveys (Mallek and Dau, 2011). There have also been multiple studies
detailing shorebird activities during the breeding season and migration in the adjacent Kilbuck
and Ahklun Mountains and coastal areas from Kuskokwim Bay to Togiak (Petersen et al., 1991).
Breeding activities of several shorebird species in the Iliamna Lake area have also been reviewed
(Williamson and Peyton, 1962).
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Table 1.2. In
Species
Black-bellied Plover
American Golden-Plover
Pacific Golden-Plover
Semipalmated Plover
Spotted Sandpiper
Wandering Tattler
Greater Yellowlegs
Lesser Yellowlegs
Whimbrel
Bristle-thighed Curlew
Hudsonian Godwit
Bar-tailed Godwit
Marbled Godwit
Ruddy Turnstone
Black Turnstone
Surfbird
Red Knot
Sanderling
Semipalmated Sandpiper
Western Sandpiper
Least Sandpiper
Baird's Sandpiper
Pectoral Sandpiper
Sharp-tailed Sandpiper
Rock Sandpiper
Dunlin
Short-billed Dowitcher
Long-billed Dowitcher
Wilson's Snipe
Red-necked Phalarope
Scientific name
Pluvialis squatarola
Pluvialis dominica
Pluvialis fulva
Charadrius semipalmatus
Actitis macularius
Tringa incana
Tringa melanoleuca
Tringa flavipes
Numenius phaeopus
Numenius tahitiensis
Limosa haemastica
Limosa lapponica
Limosafedoa
Arenaria interpres
Arenaria melanocephala
Aphriza virgata
Calidris canutus
Calidris alba
Calidris pusilla
Calidris mauri
Calidris minutilla
Calidris bairdii
Calidris melanotos
Calidris acuminata
Calidris ptilocnemis
Calidris alpina
Limnodromus griseus
Limnodromus scolopaceus
Gallinago delicata
Phalaropus lobatus
Breeding1
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Current
Trend2
Declining
Declining
Declining
Stationary
Stationary
Stationary
Stationary
Declining
Declining?
Stationary
Stationary
Declining
Unknown
Unknown
Stationary
Declining
Declining
Declining
Declining
Declining?
Declining
Stationary
Declining
Stationary
Stationary
Declining
Declining
Stationary
Declining
Declining
Conservation
Priority3
3
4
3
2
2
3
3
4
4
5
4
4
4
3
4
4
4
4
3
4
3
2
2
2
4ort3
4
4
3
3
3
1 Breeding status of "Yes" requires a record of breeding evidence (nest, eggs, or recently fledged
young on or within 150 km of the Bristol Bay Watershed.
2 Current trends were reproduced from Morrison et al. 2006, Table 1: Estimates, Current trend.
3 Conservation Status scores were reproduced from Alaska Shorebird Plan 2008, Table 2,
Conservation Category. Species in Categories 4-5 are of high concern and in category 2-3
are of low to moderate concern.
In more recent years avian surveys have been conducted by the National Park Service in Lake
Clark and Katmai NPPs (montane surveys including the upper Nushagak and Kvichak drainages)
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(Ruthrauff et al., 2007); by the USFWS (targeting lowland areas of the northern Alaska
Peninsula) (Savage and Tibbitts, In prep); by the Pacific Shorebird Migration Project (including
satellite tracking of godwits (Limosa spp.), bristle-thighed curlew (Numenius tahitiensis), and
whimbrel (N. phaeopus)) (Gill et al., 2009); using color banding (whimbrels) (Tibbetts, personal
communication); and using radio tracking and attachment of geo-locators (plovers
(Charadriidae)) (Johnson et al., 2004; Johnson et al., 2001; Johnson et al., 2011; Johnson et al.,
2008). These surveys have provided additional understanding of the importance of this region to
various stages of shorebird life history. A deficiency in information for upper Bristol Bay during
the early spring still exists, but information for nearby Egegik (Fernandez et al., 2010) and
Nanvak bays (Fernandez et al., 2010; Petersen et al., 1991) are relevant to this characterization.
The paucity of information for the early spring stems in part from the winter-like conditions that
frequently persist in the Bristol Bay region until early May, affecting, if not the birds' use of the
area, then at least the ability of biologists to access it.
The Bristol Bay/Alaska Peninsula lagoon system, of which the Nushagak and Kvichak River
deltas are part, is one of the most important migratory shorebird stop-over areas in the state. Only
the Copper River Delta and the Yukon-Kuskokwim Delta are likely more important (Gill and
Handel, 1990; Isleib and Kessel, 1973; Senner, 1979). The entire set of lagoons supports
thousands of individuals, representing numerous shorebird species, that undertake post-breeding
migrations to the Pacific coast of North America and across the Pacific Ocean to Australia,
Southeastern Asia, and Oceania. For species that migrate directly across the ocean to Hawaii or
other South Pacific islands (e.g. bar-tailed godwit (Limosa lapponica), ruddy turnstone (Arenaria
interpres)\ these lagoons provide the last stopover before their long overwater flights. Western
sandpiper (Calidris mauri), dunlin (C. alpina), and long-billed dowitcher (Limnodromus
scolopaceus) use the peninsula's lagoons to replenish energy reserves before departing non-stop
for British Columbia and points south. The Bristol Bay lagoons are also used by shorebirds as
they migrate north in spring, providing an essential refueling location that enables species not
only to succeed in reaching their breeding grounds, but also to begin breeding shortly thereafter.
The relative importance of each lagoon/delta, including the deltas of the Bristol Bay region, is
likely to vary annually and by species, and the loss of any one site might have a devastating
effect on a species' ability to successfully migrate and consequently might add another factor to
already declining populations.
Habitat
The geomorphology of upper Bristol Bay is shaped by the interaction between the shallow basin
of the Bay and the twice-daily tidal fluctuation in excess of 10 m. These features interact with the
numerous river deltas, including the Nushagak, Kvichak, and Naknek, to form an expansive
intertidal zone dominated by unvegetated sand and mudflats. The intertidal zone also includes
vegetated substrate at Nushagak Bay. These intertidal areas, the Bristol Bay estuary itself, and
other nearby river mouths with extensive mudflats along the Alaska Peninsula characterize the
estuarine portions of the region. Approximately 530 km2 of intertidal habitat is found at Kvichak
Bay and 400 km2 at Nushagak Bay (Gill et al.; Western Hemisphere Shorebird Reserve Network,
2011). In winter, substantial shore ice forms along the coast and sea ice moves through the area
with the tides. The supralittoral (splash) zone varies from gradually sloping unvegetated or
sparsely vegetated shore, to sand and morainal bluffs up to 20 m in height. Beyond the shore
zone, the region is characterized by a mosaic of wetland and tundra habitats, punctuated with low
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and tall shrub communities, located primarily along drainages. At higher elevations are spruce,
mixed spruce, birch or cottonwood forests that give way to ericaceous dwarf shrub or sparsely
vegetated substrates in the alpine zone.
Shorebirds inhabit the Bristol Bay watershed primarily during two phases of their annual life
cycle: migration and breeding season. During each phase they make use of geographically
distinct parts of the watershed. Shorebirds use the expansive intertidal and adjacent supralittoral
areas during both spring and fall migrations. In spring, the mouths of major rivers including the
Naknek, Kvichak and Nushagak, are often the first areas to become ice-free, and provide critical
feeding habitat in the littoral zone (Gill and Handel, 1981). Shorebirds actively forage in the
intertidal zone, often moving toward the declining water level as the tide drops. During high tide
and at night, birds move into the adjacent supralittoral zone to continue foraging or to roost.
Although information for these specific bays is sparse for the spring period, we can make
inference from observations in nearby areas. Beginning in late April through mid-May, the
predominant species to use the Bristol Bay region (western tip of Unimak Island north to Cape
Newenham, as defined in Gill and Handel, 1981) include: bar-tailed godwit (several thousand),
western sandpiper (a few thousand), rock sandpiper (C. ptilocnemis; a few thousand), dunlin (a
few thousand), black-bellied plover (Pluvialis squatarola; hundreds to thousands), Pacific
golden-plover (P. fulva; several hundred), black turnstone (Arenaria melanocephala; several
hundred), red-necked phalarope (Phalaropus lobatus; several hundred), Hudsonian godwit
(Limosa haemastica; a few hundred), and short-billed dowitcher (Limnodromus griseus; a few
hundred)(Gill and Handel, 1981). Other species are found in lesser numbers.
Beginning in mid- to late June, shorebirds return to Bristol Bay in larger numbers and remain for
protracted periods along the intertidal zone. This general shift between terrestrial habitats and the
littoral zone is observed throughout the region (Gill and Handel, 1981; Gill and Handel, 1990;
Gill et al., 1977; Gill et al., 1978) and Alaska (Connors, 1978; Taylor et al., 2011). Shorebirds
also make use of the supralittoral zone and terrestrial habitats near the coast during high tides and
at night for feeding and for roosting (Gill and Handel, 1981; Gill et al., 1981). Initially, the
primary birds present in the region are those nesting locally, but as the season progresses,
populations swell with birds moving into the region from nesting areas in western and northern
Alaska and as far away as eastern Russia (Gill et al., 1994). By early August it is not uncommon
to find hundreds of thousands of shorebirds on intertidal areas of upper Bristol Bay. Species
composition and abundance in fall is similar to the spring migration except for additional
species and increased numbers: bar-tailed godwit (thousands to ten thousands), dunlin (several
ten thousands), red-necked phalarope (several ten thousands), and red phalarope (several ten
thousands), western sandpiper (a few ten thousands), rock sandpiper (a few ten thousands),
short-billed dowitcher (a few ten thousands), black-bellied plover (a few thousands), Pacific
golden-plover (a few thousands), whimbrel (a few thousands), ruddy turnstones (a few
thousands), black turnstones (a few thousands), sanderlings (a few thousands), long-billed
dowitcher (a few thousands), greater yellowlegs (hundreds to thousands), semipalmated plover
(several hundreds), Hudsonian godwit (a few hundreds) (Gill and Handel, 1981; MacDonald,
2000; MacDonald and Wachtel, 1999). Populations awaiting storms to help carry them to
Australasia and the west coast of North America may extend their period of use into October
(e.g., dunlin) (Gill et al., 1978) or early November (e.g., bar-tailed godwit) (Gill et al., 2009).
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Shorebirds breeding in the Bristol Bay watershed use different habitats in the terrestrial areas of
the watershed, from the supralittoral zone (Gill and Handel, 1981; Gill et al., 1981) to elevations
of 1,300 m (Ruthrauff et al., 2007), based on their species' preferences. Many species (e.g.,
greater yellowlegs, dunlin, Wilson's snipe (Gallinago delicata), and short-billed dowitcher)
prefer mesic to wet herbaceous vegetation, while many of the plovers or montane breeders (e.g.,
American golden-plover (Pluvialis dominica), semipalmated plover, surfbird (Aphriza virgata),
rock sandpiper), prefer dwarf shrub/lichen vegetation or even barren areas for nest sites. Several
species are highly dependent on lake or river shores (e.g., spotted sandpiper (Actitis macularius),
wandering tattler (Tringa incand)) (Petersen et al., 1991). A few species (semipalmated plover,
marbled godwit (Limosa fedoa\ black turnstone, dunlin, and short-billed dowitcher) prefer the
coastal fringe (Gill and Handel, 1981; Gill et al., 2004). All of these shorebirds may feed in
marine intertidal zones during breeding, depending on their proximity or their preference for
feeding in these environments.
Food Habits
The shorebird group derives its name from the fact that many species spend migration, and often
winter, associated with shore environments. In many cases, these are marine shores. Food is
likely the most important factor controlling the movements of shorebirds throughout the Bristol
Bay region. Use of Nushagak and Kvichak Bays during shorebird migration is undisputed. In the
spring, shorebirds need to acquire critical food resources, not only to fuel their migration, but
also, for some species, to assure that they arrive on the breeding grounds with sufficient reserves
to initiate nesting and egg production (Klaassen et al., 2006; Yohannes et al., 2010). Beginning
mid-summer and continuing into early autumn, shorebirds in Alaska must again find food-rich
areas to support the process of partial or complete molt (a few species) and to fuel extended
migration (all but a few species). Indeed, some of the longest migrations known to birds involve
shorebird species (bar-tailed godwit) that use Bristol Bay intertidal areas in autumn (Battley et
al., 2011; Gill et al., 2009). Such flights are possible not only due to the extreme abundance of
intertidal invertebrates (polychaetes, crustaceans, gastropods, and bivalves) in the region, but
also because the adjacent uplands are usually rich in fruits of ericaceous plants or tubers that
birds like plovers, whimbrels, and godwits, regularly feed on (Elphick and Klima, 2002; Johnson
and Connors, 1996; Paulson, 1995; Skeel and Mallory, 1996). For species like bar-tailed godwit
and sharp-tailed sandpiper, individuals can gain up to 6% of their lean body mass per day while
feeding prior to migration (Gill et al., 2005; Lindstrom et al., 2011). Other species acquire their
fuel at a different trophic level. Rock sandpiper, for example, often eat the gonads of tide-
stranded jellyfish medusae, while species like whimbrel, Hudsonian godwit, and black and ruddy
turnstones feed on herring roe, carrion and salmon eggs (Elphick and Tibbitts, 1998; Gill et al.,
2002; Handel and Gill, 2001; Nettleship, 2000; Norton et al., 1990). During the breeding season
terrestrial and freshwater environments provide the bulk of the food sources and a wide variety
of animal and vegetable resources are consumed. Most shorebird species make use of terrestrial
invertebrates or their larvae or eggs, and many make use of freshwater invertebrates; small fish
may be consumed by yellowlegs (Elphick and Tibbitts, 1998) and phalaropes (Rubega et al.,
2000). Detailed summaries for each species are found in The Birds of North America (Cornell
Lab of Ornithology, 2011).
Shorebirds play a role in distributing MDNs into the terrestrial system, especially during the
migratory period, but this has not been quantified. They frequently feed in the intertidal zone, but
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roost in the terrestrial zone, where they frequently deposit their waste. In addition, they are prey
items for many larger predators that subsequently cycle the nutrients into the terrestrial system.
Behavior (Movements)
Shorebirds move at multiple temporal and spatial scales that are usually associated with specific
phases of their annual cycle, but within each, there can also be movement driven by more
random events such as weather and opportunistic feeding. The most obvious movements are
associated with migration, a phenomenon that occurs twice a year in most shorebirds. Spring
migration in Alaska is an end to a process that began months prior and often continents away and
is driven by the pending nesting season and the need for birds to establish territories and produce
young. As such, it is characterized as rapid and direct (Gill and Handel, 1981), with little use
made of intertidal areas once birds leave penultimate staging sites such as the Copper River
Delta (Isleib, 1979; Iverson et al., 1996; Senner, 1979).
Spring conditions in Bristol Bay vary greatly from year to year. Shore bound and riverine ice can
vary considerably with regard to magnitude and timing of melt, depending on the severity of the
winter, amount of snow cover, and the onset of spring conditions. Currently, no formal measure
of shore ice is conducted in this region; however remote sensors such as MODIS (Moderate
Resolution Imaging Spectroradiometer) could be used to describe and monitor spring conditions
(Spencer, 2006). Informal observations indicate that breakup may begin as early as late March or
be delayed until early May. The ice conditions may change within a week or may linger for four
to six weeks. Shorebirds make use of the Bristol Bay tidal flats as they become ice-free, typically
beginning in mid- to late April and peaking in early May; the length of their stay depends in part
on conditions in the nearby breeding grounds or further north on their migratory route.
Spring shorebird surveys are limited for this area and most information comes from surveys
targeting other taxa. With that in mind, note that peak numbers for single aerial surveys along the
margins of the two bays range from approximately 7,000 to 10,600 small to medium shorebirds
(Arneson, 1978; King and Dau, 1992). Arneson (1978) conducted several surveys in one spring
and mentions that spring shorebird densities can change dramatically over a short time span.
Although spring migration is abbreviated, variation in migration timing by sex is known for
some species with males generally arriving earlier (Senner et al., 1981).
Once established on the breeding grounds, most shorebirds exhibit territorial behavior.
Movements are driven by the need to defend territories, attract mates, establish and defend nests
and young, feeding, and the need to find shelter from weather and predators. Once young fledge,
or when nesting attempts fail, many species move to coastal habitats; such movement may be
driven by deteriorating food supplies on the nesting grounds or increased availability of food in
the littoral habitats (Gill and Handel, 1981).
Shorebirds return to the coastal zone beginning in mid-June, with some remaining until early
November. Summer and fall food resources in Bristol Bay are diverse and abundant, as
evidenced by the diversity and numbers of shorebirds, waterfowl and seabirds that are attracted
to the area during this time (Gill et al., 1981). A clearly attractive attribute of Bristol Bay is the
short distance birds must move between various components of its large system of
interconnected mudflats and bays, all containing concentrated food resources. In the context of
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post-breeding shorebird use, Nushagak and Kvichak Bays cannot be separated from the context
of the greater Bristol Bay/Alaska Peninsula complex. Rich food resources are in demand,
because adult birds are recovering from the energetic stress of breeding and beginning the
energy-demanding molt. Ground and aerial surveys conducted in Nushagak Bay (MacDonald,
2000; MacDonald and Wachtel, 1999) and other Alaska Peninsula lagoons (Gill et al., 1977; Gill
et al., 1978) provide insight on the seasonality, and duration of use by at least 25 shorebird
species. The magnitude of shorebird use has been captured on single-day aerial surveys in the
later part of the season; for these two bays, high counts range from 20,000 to 67,000 (Gill and
King, 1980; Gill and Sarvis, 1999; Mallek and Dau, 2004). Late summer and fall shorebird use is
likely greater than spring, due to the addition of juveniles in the population, longer residence
times, different pathways of migrants during different times of the year, or different use patterns
of individual species.
The autumn migration is broken into phases based on species, age, sex, and individual breeding
success. Species-specific use patterns have been reported for Nelson Lagoon, on the central
Alaska Peninsula (Gill and Jorgensen, 1979) and patterns for other species common to the
Bristol Bay watershed are reported from studies on the Yukon Delta (Gill and Handel, 1981; Gill
and Handel, 1990). In general, black turnstones, western sandpipers and short-billed dowitcher
move through the area the earliest; black-bellied plovers arrive later and rock sandpipers, dunlin
and sanderlings may arrive at similar or later dates, but remain longer into the fall. Failed
breeders move to the coastal zone sooner than successful breeders (Gill et al., 1983; Handel and
Dau, 1988). On the Y-K Delta, Gill and Handel (1990) observed three age-based patterns of
intertidal use through the late summer. In the most common pattern, demonstrated by western
sandpipers, adults arrived first, followed by a period in which adults and juveniles occurred
together, and finally juveniles appeared alone. In the second pattern (bar-tailed godwits, dunlin,
and rock sandpipers), adults appeared first, followed by a long period of use by both adults and
juveniles. The third pattern was demonstrated by plovers, in which only juveniles used the
intertidal zone in late summer. In addition, some species demonstrate a sex-specific pattern:
female western sandpipers depart before males (Gill and Jorgensen, 1979), but in pectoral
sandpipers (Calidris melanotos\ males depart before females (Pitelka, 1959). The specific
migration patterns demonstrated by individual shorebirds, with regard to micro- and macro-
habitat use and timing, will become clearer as researchers continue to deploy satellite
transmitters and geo-locators.
Interspecies Interactions
Shorebirds act as an intermediate link in the food web between primary producers (berries, seeds,
and tubers of plants) /consumers (invertebrates and small fish) and predatory species. Especially
during migration, when birds are concentrated, their effect on invertebrate populations in feeding
areas can be extensive (Jensen and Kristensen, 1990; Quammen, 1984; Sanchez et al., 2006;
Wilson, 1989). The response of the invertebrate prey varies with time of year, substrate, presence
of other predators, presence of other prey, and age-related factors of the prey items. Any negative
effect on prey abundance is assumedly short-term, however, as these areas are revisited by
shorebirds on the next tide and the next season on a daily and annual schedule. Shorebirds may
compete intra-specifically or inter-specifically for resources during migration as well, as
demonstrated by inter-specific aggressive interactions (Burger et al., 1979).
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Shorebird adults, young, and eggs provide food for a wide variety of predatory birds including
jaegers, gulls, raptors, owls, corvids, and shrikes. Avian predation has long been hypothesized to
be the dominant force in flocking behavior (Lack, 1954); the relationship between the benefits
(predator avoidance) and costs (feeding competition) has been explored (Stinson, 1980).
Nocturnal avian predators alter shorebird use of feeding and roosting areas (Piersma et al., 2006).
The increase in raptor populations following the removal of DDT appears to be altering how
much time shorebirds spend in marine intertidal areas; this threat of danger is potentially forcing
the birds into a trade-off between good food locations and the potential of being eaten (Ydenberg
et al., 2004). In the Bristol Bay ecosystem potential mammalian predators of shorebirds,
particularly eggs and chicks, include canids (especially foxes), lynx, weasels (including otter),
and some rodents. Shorebirds may play a role as prey in multi-species predator-prey cycles
known throughout the Arctic (Underhill et al., 1993).
Direct and indirect interactions between shorebirds and salmon are not well-documented. As
mentioned above, some shorebird species are observed to consume dead salmon and salmon
eggs, but it is unlikely that shorebirds have an impact on salmon populations. No studies have
been conducted to deduce the contribution of salmon to the energetics of shorebird populations;
however, the abundance of invertebrates in the intertidal zone is very likely due in part to
nutrients from salmon that die on the coast and in the rivers feeding Bristol Bay.
Breeding, Productivity, and Survivorship
Using information from studies in and adjacent to the Bristol Bay watershed, approximately 21
shorebird species are known to breed in this area. Most shorebird species form monogamous
pairs, with both sexes defending breeding territories and incubating eggs; however, spotted
sandpipers and red-necked phalaropes will engage in polyandry if conditions are favorable.
Individuals, and especially males, commonly demonstrate site fidelity to breeding territories.
Nesting begins in early to mid-May in the Bristol Bay area (Petersen et al., 1991). Territorial
defense is usually strongest during the early part of the breeding season and lessens as chicks
hatch (Lanctot et al., 2000). All shorebirds nesting in the Bristol Bay watershed, except solitary
sandpiper (Tringa solitarid), nest on the ground. Shorebirds usually produce four eggs per clutch.
After a nest is depredated or lost due to environmental factors, re-nesting may be attempted in
some species, but generally the season is long enough for only one complete nesting cycle
(laying, incubation, and brood-rearing). Incubation may take from 18 to 28 days, depending on
species; chicks can move to forage in habitats outside of the nesting territories within a few days
of hatching. Adults generally brood young for several days or more, until they are thermally
independent, and provide defense against predators for two to three weeks. The time from hatch
to fledging takes from 17 to 45 days depending on species shorebird. Individual breeding
behaviors are discussed at length in the species accounts of the Birds of North America (Cornell
Lab of Ornithology, 2011).
Alaska is known as a nursery ground for shorebirds; however the Bristol Bay watershed has not
been inventoried for breeding shorebird distribution or abundance. Studies from montane areas
in adjacent Katmai and Lake Clark NPPs and the Kilbuck and Ahklun Mountains and from
lowlands of the northern Alaska Peninsula provide some basis for distribution of breeding
species, but cannot be used to estimate breeding densities. For montane areas in Katmai and
Lake Clark, the most common species found during May (breeding season) were semipalmated
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plover, spotted sandpiper, wandering tattler, greater and lesser (Tringa flavipes) yellowlegs,
surfbird, least sandpiper (Calidris minutilla), and Wilson's snipe (Gill and Sarvis, 1999;
Ruthrauff et al., 2007). Ruthrauff et al. (2007) extended the breeding range of several alpine
shorebirds (wandering tattler, surfbird, and Baird's sandpiper (C. bairdiij) and confirmed these
and another three species (black-bellied plover, American golden-plover, Pacific golden-plover),
previously only known as migrants, to be breeders in Katmai. In the area west of the Bristol Bay
watershed, Petersen et al. (1991) found black-bellied plover, semipalmated plover, spotted
sandpiper, greater yellowlegs, western sandpiper, rock sandpiper, dunlin, Wilson's snipe, and
red-necked phalarope to be the most common breeding shorebirds. For lowland areas of the
northern Alaska Peninsula, the most common shorebird species found during May were greater
yellowlegs, least sandpiper, dunlin, short-billed dowitcher, Wilson's snipe, and red-necked
phalarope (Savage and Tibbitts, In prep). Other species that breed in the area include whimbrel
and Hudsonian godwit (Ruthrauff et al., 2007).
Shorebird productivity, survivorship, and mortality) are affected by many factors that may vary
by species, region and annual conditions. Measures of these parameters do not exist specifically
for the Bristol Bay watershed and may not exist at all for many species of shorebirds.
Productivity may be affected by life history (e.g., age at first reproduction, annual participation
in breeding), seasonal abundance of food resources, weather, flooding, predation, and other
forms of disturbance. Productivity in birds is measured in various ways, including proportion of
eggs hatched, proportion of successful nests, and proportion of young fledged. Shorebird pairs
may produce, at most, four chicks per season. Most small and medium-sized birds, including
shorebirds, suffer from high mortality during their first weeks and months of life. It is possible
that some species may experience complete reproductive failure for a region in some years.
Survival may be affected by food availability, weather and climate, predation, and human-caused
mortality (e.g., building strikes, domestic cat predation, and contaminated or degraded habitats);
to date, human disturbance, including habitat degradation, is significantly greater along the
migratory paths and wintering grounds of most shorebirds, than in the breeding grounds. Once
birds reach adult age and have successfully navigated their first migration, survival is generally
higher. The U.S. Geological Survey's Bird Banding Lab maintains longevity records for banded
birds (USGS Bird Banding Lab, 2011). These records indicate that smaller shorebird species
live from 6 to!2 years, while some of the medium to larger species have been recorded to live 21
(Pacific golden-plover) to 23 (bristle-thighed curlew) years. The average age of the majority of
the population is much lower and is not known for most species of shorebirds.
Population, Subpopulations, and Genetics
Shorebird populations throughout North America are experiencing declines (Alaska Shorebird
Group, 2008). Although accurate population data are lacking for most shorebirds, of the 30
regularly occurring shorebird species in the Bristol Bay watershed, 17 are suspected to be
declining and 11 are thought to be stationary; there is not enough information to make a
determination for two species (Morrison et al., 2006). None of the species using the Bristol Bay
watershed are known to be increasing in population.
Two species of shorebirds found in the Bristol Bay watershed may include two or more
subspecies, especially during the migratory period. Of the two subspecies of dunlin that breed in
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Alaska, it is most likely the Calidris alpinapacifica subspecies that migrates through Bristol Bay
and not the C.a. arcticola subspecies. Subspecies status for this species is still under
consideration (Warnock and Gill, 1996). Four to five subspecies of rock sandpiper are found in
Alaska and the most likely subspecies to use the Bristol Bay watershed for breeding would be
Calidris ptilocnemis tschuktschorum; use by C. p. couesi, or C. p. ptilocnemis during migration
may also be possible (Gill et al., 2002).
Human Use and Threats
Shorebirds have been, and continue to be, used as human food. During the latter part of the 19
and early 20th century, shorebirds were harvested commercially, along with waterfowl, for
human consumption. The overhunting of shorebirds and waterfowl and the killing of birds for
the fashion industry in part led to the development of the Migratory Bird Treaty Act of 1918, and
in North America, shorebirds are protected under its provisions. Shorebirds may still be hunted
under regulations formulated for each state. In Alaska, Wilson's snipe may be harvested during
the fall migratory bird season (1 September to 16 December in the Bristol Bay area). Sixteen
shorebird species common to Bristol Bay during some part of their life cycle may be harvested
during the Alaska Subsistence Spring/Summer Migratory Bird season (1 April to 14 June and 16
July to 31 August). A harvest survey is conducted in parts of southwest Alaska, with various
areas assessed on rotating years; however, participation is voluntary and the reports likely
represent minimum harvest levels (Naves, 2011). During the 2009 survey (included the Y-K
Delta Region), 1,688 shorebirds and 1,835 shorebird eggs were reported harvested (Naves,
2011). In general, godwits, whimbrels and curlews are targeted, due to their larger size. The
value of these shorebirds to the diet and thus economy of Native Alaskans, especially in western
Alaska, should not be underestimated.
Non-consumptive uses of shorebirds mostly include shorebird-viewing and tourism associated
with that activity. Other areas of Alaska such as Kachemak Bay, the Copper River Delta, and
Cordova are developing this industry and depend on birds that will pass through the Bristol Bay
watershed. There have been some initial attempts (http://www.visitbristolbay.com/visitor-
guide/wildlife.html) to develop the birding tourist industry in the Bristol Bay area.
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LANDBIRDS
Introduction
Approximately 80 species, representing six orders and 27 families of landbirds breed in the areas
in and adjacent to the Nushagak and Kvichak watersheds (USFWS, 2008; USFWS, 201 Ob).
Published surveys of birds in this area include biological inventories from 1902 through 1959
(Gabrielson, 1944a; Gabrielson, 1944b; Hurley, 193la; Hurley, 193Ib; Hurley, 193Ic; Hurley,
1932; Osgood, 1904; Williamson and Peyton, 1962). More recent work in this and adjacent areas
include: inventories in the Kilbuck and Ahklun Mountains (Petersen et al., 1991), inventories in
the montane regions of Lake Clark and Katmai NPPs (Ruthrauff et al., 2007), breeding bird
surveys in Dillingham, Katmai, and King Salmon, and Christmas Bird Counts in Dillingham and
King Salmon (National Audubon Society: http://audubon2.org/cbchist/count_table.html).
"Landbirds" are species that are generally associated with terrestrial habitats: passerines (or
"songbirds") and other species such as woodpeckers, owls, raptors, and gallinaceous birds
(grouse and ptarmigan). Bald eagles are addressed separately in this document.
Landbirds common in the region during the summer breeding season include, but are not limited
to, Swainson's thrush (Catharus ustulatus), American robin (Turdus migratorius), varied thrush
(Ixoreus naevius), Arctic warbler (Phylloscopus borealis), orange-crowned warbler (Vermivora
celata), and Wilson's warbler (Wilsonia pusilld) (USGS, 2011). Numerous other songbirds
regularly nest here, including several other species of swallows (Hirimdinidae), thrushes
(Turdidae), warblers (Parulidae), sparrows (Emberizidae), and others. Year-round resident
species include northern goshawk (Accipiter gentilis), great-horned owl, common raven (Corvus
cor ax), gray jay (Perisoreus canadensis), black-billed magpie (Pica pica), black-capped and
boreal chickadees (Poecile atricapillus and P. hudsonicus), American dipper (Cinclus
mexicanus), common redpoll (Carduelis flammed), and snow bunting (Plectrophenax nivalis)
(ADNR, 2008; USGS, 2011). Of the relatively common species occurring in the area, two (short-
eared owl (Asio flammeus) and rusty blackbird (Euphagus carolinus)) are on the Continental
Watch List. Twenty-six of the Bristol Bay landbird species are on the Continental Stewardship
list for Partners in Flight, a multi-stakeholder partnership dedicated to the conservation of
landbirds (Rich et al., 2004).
Habitat
Most species of landbirds occupy and defend individual breeding territories during the spring
and summer. Few studies have focused on landbirds in the Nushagak and Kvichak watersheds
and site-specific information is extremely sparse. Migratory species begin arriving in late April
and may remain through late September (Savage, personal communication).
The Nushagak and Kvichak watersheds are within, albeit near the border of, the Arctic avifaunal
biome, as described by Partners in Flight (Rich et al., 2004). The diversity, population,
distribution, and densities of birds here are not well-understood. While no comprehensive studies
have been published for these watersheds, land bird density and diversity, generally speaking,
may be highest along the numerous riparian corridors of the region (Boreal Partners in Flight
Working Group, 1999; Williamson and Peyton, 1962). Greater landbird densities in the riparian
zone often occur in areas where the surrounding habitats have lower plant species or canopy
layer diversity (Stauffer and Best, 1980; Wiebe and Martin, 1998). Riparian habitats in these
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watersheds include ribbons of tall shrub (willow/cottonwood/alder), as well as spruce, birch and
mixed forests, which wind and branch across vast acreages of moist and wet tundra (ADF&G,
2006; Nushagak-Mulchatna Watershed Council, 2007).
From the sparse site-specific data available, we are able to collect the following information to
begin a characterization of local species diversity. For the Western Alaska Lowlands/Uplands
Bird Conservation Region 2 (BCR 2), which includes the Ahklun Mountains and Bristol Bay-
Nushagak Lowlands, passerine diversity is thought to be greatest in riparian tall shrub habitats
(Boreal Partners in Flight Working Group, 1999). 5 Sixteen species of passerines, one
woodpecker species, and belted kingfisher have been recorded along the Alagnak River
(Gotthardt et al., 2010), although proximity to the river and whether or not the records were
associated with riparian habitats is unknown.
Mixed spruce-birch forests of the area may also have a relatively high species diversity, with 25
to 27 species noted by Williamson and Peyton (1962). Forty species were recorded in plots in
LCNPP, within the upper Mulchatna and upper Kvichak watersheds; the survey area likely
included some non-riparian areas (Ruthrauff et al., 2007).
No site-specific density information is available for the Bristol Bay watershed. Regarding
potential densities, Kessel reported between 11.8 and 45.4 passerine/woodpecker breeding
territories per 10-hectare plot (mostly riparian) in Interior Alaska, (Kessel, 1998). However,
significantly wide ranges of variability in landbird breeding densities reduce any potential
usefulness of these figures for the local area.
Landbird diet requirements vary by species and time of year; foods include: vegetation (seeds,
berries), invertebrates (aquatic and terrestrial, as well as flying insects), and vertebrates (other
birds, fish and mammalian carrion, juvenile fish, fish eggs). During the breeding season, adults
face high demands, associated with producing eggs, feeding young, and molting; young birds
require considerable food resources to grow and both young and adults must gain fat prior to
migration. Even birds that consume a high proportion of seeds and other vegetative matter in the
non-breeding season may switch to food in higher trophic levels during the breeding season. In
general, the timing of hatch and the growth of young landbirds is directly related to the
abundance of invertebrate food sources (Ehrlich et al., 1988). In fact, the abundance of emergent
aquatic insects may be one of the reasons riparian habitats are often associated with greater avian
abundance (Iwata et al., 2003; Murakami and Nakano, 2002). Foraging techniques for avian
predators of invertebrates (e.g., foliage gleaning, ground foraging, aerial predating/flycatching)
vary considerably among species, however, so landbirds can exploit a variety of riparian
invertebrate prey types overall (Murakami and Nakano, 2001).
Interspecies Interactions
Regarding the importance of salmon to landbirds, recent studies have indicated that the
abundance of many species of songbirds is related to the presence of salmon carcasses in
5 BCR 2 includes most of the middle and lower areas of the Nushagak and Kvichak watersheds, but extends beyond
them, from the Kuskokwim River to Unimak Pass.
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freshwater streams (Christie and Reimchen, 2008; Gende and Willson, 2001; Willson et al.,
1998). The relationship is complex and not yet fully understood (Christie and Reimchen, 2008;
Gende et al., 2002). One primary component, though, appears to be the initial positive effect of
the seasonal accumulation of carcasses on invertebrate populations (Helfield and Naiman, 2006).
For example, masses of aquatic invertebrate larvae feed on salmon carcasses (Wipfli et al.,
1999), over-winter in the soil, and emerge in the spring as adults, subsequently becoming aerial
prey for songbirds, an important seasonal subsidy that becomes available during the same period
when terrestrial invertebrate biomass is low (Nakano and Murakami, 2001). Terrestrial
invertebrate (e.g., litter detritivore) populations may also increase as a result of salmon carcass
abundance, providing another important food source for passerines (Gende and Willson, 2001).
Passerines such as Pacific wren {Troglodytes pacificus) and other species also feed directly on
fly larvae within dead salmon in the fall (Christie and Reimchen, 2008).
Other important relationships between salmon and landbirds include the effects of increased
plant productivity, particularly in riparian areas, that appears to result from MDN input from
salmon (Gende et al., 2002; Helfield and Naiman, 2001; Hilderbrand et al., 2004; Naiman et al.,
2002a). This increased productivity, reflected, for example, in an abundance of berries and seeds,
in turn provides an increased vegetative food source for landbirds such as Swainson's and varied
thrushes (Christie and Reimchen, 2008). Another positive effect of spawning salmon on Alaska
landbirds is indicated in the case of American dippers. Dippers feed primarily on aquatic
invertebrates, which appear to increase in abundance with salmon carcasses, and females switch
to salmon eggs, fry, and small bits of carcasses during the egg-laying period (Morrissey et al.,
2010). In one study of dippers, there was an apparent positive correlation between consumption
of salmon fry and higher fledgling mass and less brood mortality (Obermeyer et al., 2006).
As noted above, the trophic relationships among salmon, landbirds, invertebrate prey items, and
other organisms are complex, and not yet well understood, particularly in relatively remote and
undisturbed boreal regions, such as the Nushagak and Kvichak watersheds. However, these
relationships are apparently significant to many aspects of landbird life history. In summary,
abundance, distribution, productivity, habitat use, and foraging habits are some of the ways in
which multiple species of landbirds may be affected by salmon. The temporal nature of the
pulses of the abundant food source salmon provide is of particular interest. Several researchers
have examined such seasonal resource subsidies in the riparian forest and it has been suggested
(Takimoto et al., 2002) that seasonal productivity differences between spatially linked habitats
help foster the stability of food web dynamics (Wiebe and Martin, 1998; Zhang et al., 2003).
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APPENDIX 1: LIST OF AUTHORS AND REVIEWERS
Species
Primary Author(s) (Affiliation)
Expert Reviewer (Affiliation)
Overall Report
Phil Brna (USFWS/AFWFO)
Lori Verbrugge (USFWS/AFWFO)
Land Cover
Phil Brna (USFWS/AFWFO)
David Selkowitz (USGS/ Alaska
Science Center)
Jerry Tande (USFWS/NWI
Program)
Julie Michaelson (USFWS/NWI
Program)
Marcus Geist (The Nature
Conservancy)
Brown Bear
Colleen Matt (C.A. Matt; ADF&G/
Retired)
Sterling Miller (ADF&G/Retired)
Sean Farley (ADF&G)
Grant Fiilderbrand (NPS)
Cara Staab (BLM)
Susan Savage (USFWS/Alaska
Peninsula-Becharof NWRs)
Patrick Walsh (USFWS/Togiak
NWR)
Buck Mangipane (NPS/ Lake Clark
NP)
Page Spencer (NPS/Retired)
Lem Butler (ADF&G/Wildlife
Conservation)
Meghan Riley (ADF&G/Wildlife
Conservation)
Jim Woolington (ADF&G/Wildlife
Conservation)
Caribou
Lori Verbrugge (USFWS/AFWFO)
Ken Whitten (ADF&G/Retired)
Layne Adams (USGS/Alaska
Science Center)
Dominique Watts (USFWS/Alaska
Peninsula-Becharof NWRs)
Bob Tobey (ADF&G/Retired)
Andy Aderman (USFWS/Togiak
NWR)
Buck Mangipane (NFS/Lake Clark
NP)
Cara Staab (BLM)
Jeff Shearer (NFS/Lake Clark NP)
Lem Butler (ADF&G/Wildlife
Conservation)
Meghan Riley (ADF&G/Wildlife
Conservation)
Jim Woolington (ADF&G/Wildlife
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Conservation)
Nick Demma (ADF&G/Wildlife
Conservation)
Bruce Seppi (BLM)
Moose
Lori Verbrugge (USFWS/AFWFO)
Chuck Schwartz (ADF&G and
USGS/Retired)
Wolf
Lori Verbrugge (USFWS/AFWFO)
Layne Adams (USGS/Alaska
Science Center)
Buck Mangipane (NFS/Lake Clark
NP)
Dominique Watts (USFWS/Alaska
Peninsula- Becharof NWRs)
Ashley Stanek (UAA/ENRI)
Ken Whitten (ADF&G/Retired)
Bob Tobey (ADF&G/Retired)
Cara Staab (BLM)
Bruce Seppi (BLM)
Page Spencer (NPS/Retired)
Bald Eagle
Maureen de Zeeuw
(USFWS/AFWFO)
Lowell H. Suring (Northern
Ecologic LLC)
Denny Zwiefelhofer
(USFWS/Retired)
Steve Lewis (USFWS/Migratory
Birds)
Michael Swaim (USFWS/Togiak
NWR)
Landbirds
Maureen de Zeeuw
(USFWS/AFWFO)
Susan Savage (USFWS/Alaska
Peninsula-Becharof NWRs)
Meghan Riley (ADF&G/Wildlife
Conservation)
Shorebirds
Susan Savage (USFWS/Alaska
Peninsula-Becharof NWRs)
Bob Gill (USGS/Alaska Science
Center)
Heather Coletti (NFS/Lake Clark
NP)
Rick Lanctot (USFWS/Migratory
Birds)
Steve Kendall (USFWS)
Waterfowl
Tom Rothe (Halcyon Research;
ADF&G/ Retired)
Christian Dau
(USFWS/Migratory Birds)
Maureen de Zeeuw
(USFWS/AFWFO)
Species List
Maureen de Zeeuw
(USFWS/AFWFO)
Susan Savage (USFWS/Alaska
Peninsula-Becharof NWRs)
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APPENDIX 2: SOUTHWEST ALASKA TERRESTRIAL VERTEBRATE
SPECIES
BIRDS
(TogiakNWR, Bird List 2006; D. Ruthrauff et al. 2007; Alaska Peninsula and Becharof NWR
Bird List, 2010; with edits from S. Savage, M. Swaim, D. Ruthrauff)
The following species are thought to regularly occur in the Nushagak and Kvichak watersheds
based on surveys and observations documenting their presence in adjacent federal land
management areas. The species are marked as breeders if they are known to breed in the adjacent
areas. Other species that may occur as accidentals are not included here.
Common Name
Waterfowl
Greater White-fronted Goose
Emperor Goose
Snow Goose
Brant
Cackling Goose
Canada Goose
Trumpeter Swan
Tundra Swan
Gadwall
Eurasian Wigeon
American Wigeon
Mallard
Northern Shoveler
Northern Pintail
Green-winged Teal
Canvasback
Redhead
Ring-necked Duck
Greater Scaup
Lesser Scaup
Steller's Eider
Spectacled Eider
King Eider
Common Eider
Harlequin Duck
Surf Scoter
White-winged Scoter
Black Scoter
Long-tailed Duck
Scientific Name
Anser albifrons
Chen canagica
Chen caerulescens
Branta bernicla
Branta hutchinsii
Branta canadensis
Cygnus buccinator
Cygnus columbianus
Anas strepera
Anaspenelope
Anas americana
Anas platyrhynchos
Anas clypeata
Anas acuta
Anas crecca
Aythya valisineria
Aythya americana
Aythya collaris
Aythya marila
Aythya affmis
Polysticta stelleri
Somateria fischeri
Somateria spectabilis
Somateria mollissima
Histrionicus histrionicus
Melanitta perspicillata
Melanitta fusca
Melanitta americana
Clangula hyemalis
Breeder
119
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Common Name
Bufflehead
Common Goldeneye
Barrow's Goldeneye
Common Merganser
Red-breasted Merganser
Gallinaceous Birds
Spruce Grouse
Willow Ptarmigan
Rock Ptarmigan
White-tailed Ptarmigan
Loons
Red-throated Loon
Pacific Loon
Common Loon
Scientific Name
Bucephala albeola
Bucephala clangula
Bucephala islandica
Mergus merganser
Mergus serrator
Falcipennis canadensis
Lagopus lagopus
Lagopus muta
Lagopus leucura
Gavia stellata
Gavia pacifica
Gavia immer
Breeder
Grebes
Horned Grebe
Red-necked Grebe
Tubenoses
Northern Fulmar
Sooty Shearwater
Short-tailed Shearwater
Fork-tailed Storm-Petrel
Leach's Storm-Petrel
Podiceps auritus
Podiceps grisegena
Fulmarus glacialis
Puffmus griseus
Puffmus tenuirostris
Oceanodroma furcata
Oceanodroma leucorhoa
Cormorants
Double-crested Cormorant
Red-faced Cormorant
Pelagic Cormorant
Hawks, Eagles, Falcons
Osprey
Bald Eagle
Northern Harrier
Sharp-shinned Hawk
Northern Goshawk
Red-tailed Hawk
Rough-legged Hawk
Golden Eagle
Phalacrocorax auritus
Phalacrocorax urile
Phalacrocorax pelagicus
Pandion haliaetus
Haliaeetus leucocephalus
Circus cyaneus
Accipiter striatus
Accipiter gentilis
Buteo jamaicensis
Buteo lagopus
Aquila chrysaetos
120
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Common Name
American Kestrel
Merlin
Gyrfalcon
Peregrine Falcon
Cranes
Sandhill Crane
Scientific Name
Falco sparverius
Falco columbarius
Falco rusticolus
Falco peregrinus
Grus canadensis
Breeder
Shorebirds
Black-bellied Plover
American Golden-Plover
Pacific Golden-Plover
Semipalmated Plover
Black Oystercatcher
Spotted Sandpiper
Solitary Sandpiper
Wandering Tattler
Greater Yellowlegs
Lesser Yellowlegs
Whimbrel
Bristle-thighed Curlew
Hudsonian Godwit
Bar-tailed Godwit
Marbled Godwit
Ruddy Turnstone
Black Turnstone
Surfbird
Red Knot
Sanderling
Semipalmated Sandpiper
Western Sandpiper
Least Sandpiper
Baird's Sandpiper
Pectoral Sandpiper
Rock Sandpiper
Dunlin
Short-billed Dowitcher
Long-billed Dowitcher
Wilson's Snipe
Red-necked Phalarope
Red Phalarope
Pluvialis squatarola
Pluvialis dominica
Pluvialis fulva
Charadrius semipalmatus
Haematopus bachmani
Actitis macularius
Tringa solitaria
Tringa incana
Tringa melanoleuca
Tringa flavipes
Numenius phaeopus
Numenius tahitiensis
Limosa haemastica
Limosa lapponica
Limosafedoa
Arenaria interpres
Arenaria melanocephala
Aphriza virgata
Calidris canutus
Calidris alba
Calidris pusilla
Calidris mauri
Calidris minutilla
Calidris bairdii
Calidris melanotos
Calidris ptilocnemis
Calidris alpina
Limnodromus griseus
Limnodromus scolopaceus
Gallinago delicata
Phalaropus lobatus
Phalaropus fulicarius
121
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Common Name
Gulls and Terns
Black-legged Kittiwake
Sabine's Gull
Bonaparte's Gull
Mew Gull
Herring Gull
Slaty-backed Gull
Glaucous-winged Gull
Glaucous Gull
Aleutian Tern
Arctic Tern
Jaegers
Pomarine Jaeger
Parasitic Jaeger
Long-tailed Jaeger
Alcids
Common Murre
Thick-billed Murre
Pigeon Guillemot
Marbled Murrelet
Kittlitz's Murrelet
Ancient Murrelet
Parakeet Auklet
Rhinoceros Auklet
Horned Puffin
Tufted Puffin
Scientific Name
Rissa tridactyla
Xema sabini
Chroicocephalus Philadelphia
LOTUS canus
Lams argentatus
Larus schistisagus
LOTUS glaucescens
Larus hyperboreus
Onychoprion aleuticus
Sterna paradisaea
Stercorarius pomarinus
Stercorarius parasiticus
Stercorarius longicaudus
Uria aalge
Uria lomvia
Cepphus columba
Brachyramphus marmoratus
Brachyramphus brevirostris
Synthliboramphus antiquus
Aethia psittacula
Cerorhinca monocerata
Fratercula corniculata
Prater cula cirrhata
Breeder
Owls
Great Horned Owl
Snowy Owl
Northern Hawk Owl
Great Gray Owl
Short-eared Owl
Boreal Owl
Northern Saw-whet Owl
Hummingbirds
Rufous Hummingbird
Bubo virginianus
Bubo scandiacus
Surnia ulula
Strix nebulosa
Asio flammeus
Aegolius funereus
Aegolius acadicus
Selasphorus rufus
122
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Common Name
Kingfishers
Belted Kingfisher
Woodpeckers
Downy Woodpecker
Hairy Woodpecker
American Three-toed
Woodpecker
Black-backed Woodpecker
Northern Flicker
Flycatchers
Olive-sided Flycatcher
Alder Flycatcher
Say's Phoebe
Shrikes
Northern Shrike
Scientific Name
Megaceryle alcyon
Picoides pubescens
Picoides villosus
Picoides dorsalis
Picoides arcticus
Colaptes auratus
Contopus cooperi
Empidonax alnorum
Sayornis soya
Lanius excubitor
Breeder
Crows, Jays, Magpies
Gray Jay
Steller's Jay
Black-billed Magpie
Northwestern Crow
Common Raven
Perisoreus canadensis
Cyanocitta stelleri
Pica hudsonia
Corvus caurinus
Corvus cor ax
Larks
Horned Lark
Swallows
Tree Swallow
Violet-green Swallow
Bank Swallow
Cliff Swallow
Barn Swallow
Eremophila alpestris
Tachycineta bicolor
Tachycineta thalassina
Riparia riparia
Petrochelidon pyrrhonota
Hirundo rustica
Chickadees
Black-capped Chickadee
Boreal Chickadee
Poecile atricapillus
Poecile hudsonicus
Nuthatches
Red-breasted Nuthatch
Sitta canadensis
123
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Common Name
Scientific Name
Breeder
Creepers
Brown Creeper
Wrens
Pacific Wren
Dippers
American Dipper
Kinglets
Golden-crowned Kinglet
Ruby-crowned Kinglet
Old World Warblers
Arctic Warbler
Thrushes
Northern Wheatear
Gray-cheeked Thrush
Swainson's Thrush
Hermit Thrush
American Robin
Varied Thrush
Certhia americana
Troglodytes pacificus
Cinclus mexicanus
Regulus satrapa
Regulus calendula
Phylloscopus borealis
Oenanthe oenanthe
Catharus minimus
Catharus ustulatus
Catharus guttatus
Turdus migratorius
Ixoreus naevius
Watgtails and Pipits
Eastern Yellow Wagtail
American Pipit
Waxwings
Bohemian Waxwing
Longspurs and Buntings
Lapland Longspur
Snow Bunting
McKay's Bunting
Wood Warblers
Northern Waterthrush
Orange-crowned Warbler
Yellow Warbler
Blackpoll Warbler
Motacilla tschutschensis
Anthus rubescens
Bombycilla garrulus
Calcarius lapponicus
Plectrophenax nivalis
Plectrophenax hyperboreus
Parkesia noveboracensis
Oreothlypis celata
Setophaga petechia
Setophaga striata
124
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Common Name
Yellow-rumped Warbler
Wilson's Warbler
Scientific Name
Setophaga coronata
Cardellina pusilla
Breeder
Sparrows
American Tree Sparrow
Savannah Sparrow
Fox Sparrow
Song Sparrow
Lincoln's Sparrow
White-crowned Sparrow
Golden-crowned Sparrow
Dark-eyed Junco
Blackbirds
Rusty Blackbird
Finches
Gray-crowned Rosy-Finch
Pine Grosbeak
Red Crossbill
White-winged Crossbill
Common Redpoll
Hoary Redpoll
Pine Siskin
Spizella arborea
Passerculus sandwichensis
Passerella iliaca
Melospiza melodia
Melospiza lincolnii
Zonotrichia leucophrys
Zonotrichia atricapilla
Junco hyemalis
Euphagus carolinus
Leucosticte tephrocotis
Pinicola enucleator
Loxia curvirostra
Loxia leucoptera
Acanthis flammea
Acanthis hornemanni
Spinuspinus
MAMMALS
(Cook and MacDonald, 2005; USFWS, 2009b)
Common Name
Terrestrial Mammals
Shrews
Masked shrew
Pygmy shrew
Tundra shrew
Alaska tiny shrew
Arctic shrew
Montane shrew
Northern water shrew
Scientific Name
Sorex cinereus
Sorex hoyi
Sorex tundremis
Sorex yukonicus
Sorex arcticus
Sorex monticolus
Sorex palustris
125
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Common Name
Bats
Little brown bat
Canids
Arctic fox
Coyote
Wolf
Red fox
Cats
Lynx
Weasels
River otter
Wolverine
Marten
Ermine
Least weasel
Mink
Scientific Name
Myotis lucifigus
Alopex lagopus
Canis latrans
Canis lupus
Vulpes vulpes
Lynx canadensis
Lutra canadensis
Gulo gulo
Maries americana
Mustela erminea
Mustela nivalis
Mustela vison
Bears
Black bear
Brown bear
Ursus americanus
Ursus arctos
Ungulates
Moose
Caribou
Dall sheep
Rodents
Hoary marmot
Arctic ground squirrel
Red squirrel
Beaver
Meadow jumping mouse
Northern red-backed vole
Northern collared lemming
Brown lemming
Northern bog lemming
Meadow vole
Tundra vole
Singing vole
Muskrat
Porcupine
Collared pika
A Ices alces
Rangifer tarandus
Ovis dalli
Marmota caligata
Spermophilus parryii
Tamiasciurus hudsonicus
Castor canadensis
Zapus hudsonius
Clethrionomys rutilus
Dicrostonyx groenlandicus
Lemmus trimucronatus
Synaptomys borealis
Microtus pennsylvanicus
Microtus oeconomus
Microtus miurus
Ondatra zibethicus
Erethizon dorsatum
Ondatra collaris
126
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Common Name Scientific Name
Hares
Snowshoe hare Lepus americanus
Tundra hare Lepus othus
127
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Appendix D
Traditional Ecological Knowledge and
Cultural Characterization of the
Nushagak and Kvichak Watersheds, Alaska
D-l
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TRADITIONAL ECOLOGICAL
KNOWLEDGE AND CULTURAL
CHARACTERIZATION OF THE NUSHAGAK
AND KVICHAK WATERSHEDS, ALASKA
Submitted to the Bristol Bay Assessment:
Environmental Protection Agency
Nushagak River at Koliganek, September 19,2011
Alan S. Boraas, Ph.D.
Professor of Anthropology
Kenai Peninsula College
ifasb@kpc.alaska.edu
and
Catherine H. Knott, Ph.D.
Assistant Professor of Anthropology
Kenai Peninsula College
ifchk@kpc.alaska.edu
March 22, 2012
Public Review Draft
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Boraas and Knott
Cultural Characterization
EXECUTIVE SUMMARY
1. Voices of the People
...Salmon more or less defines this area. It defines who we are. When you look at our art, you
will see salmon.... It is who we are. When you listen to the stories and take a steam, even in the
middle of winter, people talk about salmon. It is in our stories; it is in our art. It is who we are; it
defines us. M-61,9/16/11
...we are relying on EPA to give us a fair shake out here. If EPA is going to crap all over our
people, then take out the checkbook, federal government, and start writing million dollar checks
for these people to move to Anchorage because you are going to kill us culturally, economically
and every other way. M-60, 9/16/11
But Iwouldn 't trade this place for anything. This is home; this is where I find clean water to
drink. M-51,8/20/11
We love the place; it's home. Moving is not an option to me. M-29, 8/17/11
... basically one of the main purposes of the Blessing of the Water is to make that Holy water....
When the Father blesses that particular river, that particular river becomes Holy. M-61, 9/16/11
/ think with us, during potlatch times, during hard times, or Russian Christmas, or if we gather
together, everybody brings out their dry fish or their jarred fish or their salt fish. Nobody goes
hungry, there's always sharing. F-32, 8/18/11
We share with our families, or if anybody does not have fish, we give them fish also. F-27,
8/17/11
2. The Condition of the Indigenous Cultures of the Bristol Bay Region
This section of the Bristol Bay Assessment is based on 53 interviews in seven villages
and an overview of previous research in the study area. The condition of the ecosystems, both
riverine and lacustrine, on which the Yup'ik and Dena'ina depend for wild fish, mammals, and
plants including the keystone species salmon, is nearly pristine. The cultures have proved to be
sustainable in this region for thousands of years. Alaska Department of Fish and Game statistics
indicate wild subsistence resources including salmon provide the Yup'ik and Dena'ina of the
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Boraas and Knott
Cultural Characterization
study area with the bulk of their food resources. Wild foods provide critical nutritional elements
in both quantity and quality in the diet, but subsistence also forms the core of the culture itself,
including knowledge, attitudes, practices, and beliefs important to the Yup'ik and Dena'ina
people in their daily lives.
The villages of the study area are predominantly Alaska Native and the population
remains stable (United States Census, Alaska). The culture has a very high degree of
homogeneity as represented by interviewees' responses to this set of questions revolving around
the importance of salmon and streams in their lives. Interviews conducted in this project relating
to the importance and significance of salmon and clean water resulted in 97% concurrence
among Elders and culture bearers. The Yup'ik people of the region retain their language, and
more than 40% of the population continues to speak it. The Dena'ina are undergoing a cultural
renaissance through language revitalization programs and the emergence of culture camps. Both
languages have a large number of words related to salmon and stream resources reflecting
nuanced understanding developed over time.
Elders and culture bearers continue to instruct young people particularly at fish camps
where not only fishing and processing techniques are taught, but also cultural values. The social
system which forms the backbone of the culture, nurturing the young, supporting the producers,
and caring for the Elders, is based upon the virtue of sharing the wild foods harvested from the
land and waters. Sharing networks extend to family members living far from home. The first
salmon catch of the year is recognized with a prayer of thanks and shared in a continuation of the
ancient First Salmon Ceremony.
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Boraas and Knott
Cultural Characterization
The Yup'ik and Dena'ina consider the land and waters to be their sacred homeland. They
have traditionally considered the salmon as kin in the sacred web of life. The populations of both
Yup'ik and Dena'ina have shown themselves to be spiritually tenacious, combining elements of
traditional practices with those of Russian Orthodox and other Christian churches to create a rich
syncretic religious heritage for their families. The rivers are blessed by priests annually in the
Great Blessing of the Water at Theophany, celebrating the baptism of Christ. This ceremony, for
Orthodox Yup'ik and Dena'ina, is the pure element of God expressed as sanctified nature. The
holy water of the rivers derived from this ceremony is used to bless the homes, churches, and
people and is believed to have curative powers.
3. The Status of the Resource Relative to other Salmon Culture Ecosystems Internationally
The Human Relations Area Files on-line cultural database (Human Relations Area Files,
World Cultures Data Base, http://www.yale.edu/hraf/collections.htm) identifies 23 world
cultures in which anadromous salmon are, or were, a chief component of subsistence. Only in
Alaska are wild, non-farmed, non-hatchery spawned, non-bioengineered salmon abundant. The
Yup'ik and Dena'ina of the study area are among the few remaining cultures to still rely on wild
salmon as a chief source of nutrients and have an intact relationship with the landscape that
supports them.
4. The Causes of the Unique Status of the Resource and the Vulnerability of the Resource
This area is among the last remaining truly viable cultural and ecologically interdependent
human/salmon ecosystem in the world because it is an intact ecosystem largely due to the fact
that it is remote, roadless, and until recently, not thought to contain natural resources of value
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Boraas and Knott
Cultural Characterization
other than fish and game. In addition the unique Alaska State and United States Federal
subsistence laws protect the indigenous people's right to harvest wild resources.
5. Vulnerabilities
The existing culture of the indigenous people of the study area is vulnerable to anything
that would change the quantity or quality of wild salmon resources or the quantity or quality of
water in the Nushagak or Kvichak watersheds. Negative impacts to salmon would leave the
existing culture susceptible to destabilization and affect its present ability to cope with natural
disasters. If significant negative impacts to salmon or streams occur, the cultural stability will be
vulnerable to change in the following ways:
Since the diet is heavily dependent on wild foods, particularly salmon, the diet would be
significantly changed from a highly nutritious diet to one based on store-bought
processed foods.
Since the social networks are highly dependent on procuring salmon (fish camps) but also
sharing salmon and wild food resources, the current social support system would be
significantly degraded
Since significant, meaningful family-based work takes place in fish camp or similar
subsistence settings, transmission of cultural values and language learning would be
impacted and family cohesion impacted.
Since values and the belief system are represented by interaction with the natural world
through salmon practices and clean water practices and symbolic rituals, core beliefs
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would be challenged potentially resulting in a breakdown of cultural values, mental
health degradation and behavioral disorders.
Since Alaskan state and federal subsistence law currently rests on rural and urban
designations, a significant increase in population potentially would result in loss of
subsistence rights if an area were re-designated "urban."
Since a yearly subsistence round rests on having time to harvest and process wild foods, a
shift from part-time wage employment supporting subsistence to full-time wage
employment would impact subsistence-gathering capabilities by restricting the time
necessary to harvest subsistence resources.
Since the area exhibits a high degree of cultural uniformity tied to shared subsistence
practices, significant change could provoke increased tension and discord both between
villages and among villagers.
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Table of Contents
I. INTRODUCTION 9
A. Overview 9
B. Villages, Population, and Ethnicity 15
II. CULTURAL AND HISTORICAL BACKGROUND 20
A. Pre-Contact Bristol Bay 20
1. Voices of the People 20
2. Introduction 20
3. Pre-Contact Salmon Fishing Cultures 23
B. History and Culture of the Yup'ik Area 31
1. Voices of the People 31
2. Introduction 33
3. Pre-Contact Culture 33
4. Post-Contact History and Culture (A.D. 1791 to 1935) 35
C. History and Culture of the Dena'ina 41
1. Voices of the People 41
2. Pre-Contact Culture 42
3. Post-contact History and Culture 44
D. Traditional Yup'ik and Dena'ina Spirituality and Cosmology 47
1. The Yup'ik People 47
2. The Dena'ina People 52
E. The Yup'ik and Dena'ina Languages: Salmon and Streams 57
1. Voices of the People 57
2. Introduction 57
3. The Central Yup'ik Language 57
4. The Dena'ina Language 68
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III. MODERN CULTURE 80
A. Interview Synopsis 80
B. Subsistence 85
1. Voices of the People 85
2. Introduction 87
3. Subsistence in Alaska 88
4. Scope of Subsistence 91
5. The Seasonal Subsistence Round 96
6. The Interplay of Subsistence and Wage Income 97
7. Subsistence as an Economic Sector 100
8. Subsistence and "Wealth" 103
C. Physical and Mental Well-being: the Role of Subsistence 105
1. Voices of the People 105
2. Introduction 107
3. Nutrition 109
4. Fitness Ill
5. Disease Prevention 112
6. Local Wild Fish 114
D. Traditional Ecological Knowledge 116
1. Voices of the People 116
2. Introduction 117
E. Social Relations 124
1. Voices of the People 124
2. Introduction 127
3. Sharing and Generalized Reciprocity 127
4. Fish Camp 130
5. Steam Baths 131
6. Gender and Age Equity 132
7. Wealth 133
8. Suicide in the Study Area 134
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F. Spirituality and Beliefs Concerning Water and Salmon 137
1. Voices of the People 137
2. Introduction 140
3. Great Blessing of the Water 141
4. Respect and Thanks 144
5. First Salmon Ceremony 145
G. Messages From the People 147
1. Voices of the People 147
IV. CONCLUSIONS 150
V. APPENDIX 1. METHODOLOGY and TRIBAL LETTER OF INTRODUCTION 154
VI. REFERENCES CITED 158
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I. INTRODUCTION
A. Overview
The purpose of the Bristol Bay Cultural Assessment is to provide information to the
Environmental Protection Agency on the status of the indigenous cultures of the Nushagak and
Kvichak River watersheds and their dependence on and relationship to salmon and other stream-
based natural resources of the region. The focus of the Bristol Bay Assessment is salmon and
water and this part of the overall assessment portrays the human dimension of modern
indigenous "salmon-cultures" of the region. The Human Relations Area Files on-line cultural
database (http://www.yale.edu/hraf/collections.htm) identifies 23 cultures in which anadromous
salmon are or were a chief component of subsistence. Wild Atlantic salmon populations have
been decimated by high-seas fishing and dam building (Montgomery 2003:111-118) and
consequently indigenous cultures such as the Sami of Fennoscandia, Micmac and Abnacki of
northeastern North America and other cultures once dependent on Atlantic salmon have been
forced to choose non-traditional options (cf Lethola 2004: 72-84). In the Asian Far East wild
salmon have likewise been decimated in Japan and Russia through overfishing and habitat
destruction and cultures like the Ainu of Hokkaido and Nvkh of Sakhalin Island can no longer
depend on wild salmon and cultural institutions based on salmon have been severely affected (cf.
Iwasaki-Goodman and Nomoto 1998: 27-46). In the Pacific Northwest of North America
hydroelectric dam building, overfishing, and habitat degradation have decimated wild salmon
runs and the Northwest Coast cultures from California to British Columbia can no longer subsist
on wild salmon as they once did (cf. Johnsen 2009). The Yup'ik of the Nushagak and Kvichak
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River watersheds and the Dena'ina of the Lake Iliamna, Newhalen River and Lake Clark (also
the Kvichak River watershed) are among the remaining cultures still relying on wild salmon as a
chief source of nutrients. This reliance on salmon has lasted unbroken for 4000 years and salmon
subsistence has shaped cultural patterning in multiple ways. Today modern technology is used
but many beliefs, social practices and components of spirituality are part of this long history and
form both Yup'ik and Dena'ina essential identity and provide the cultural basis for sustainability.
To say they are the last wild salmon cultures is an overstatement, but they are certainly among
the last. Part of the reason they remain is that Alaska in general, and Bristol Bay in particular,
has become the world's last bastion of wild, non-farmed, non-hatchery raised, wild salmon.
This document contains five parts. First, this introduction contains information about the
project and its methodology. Second, it consists of contextualization of relevant prehistoric,
historic, linguistic, and cultural information obtained from anthropological, historical, and other
publications and data bases. Third, this document includes the product of interviews in villages
of the Nushagak and Kvichak River watersheds conducted in 2011, which constitutes original
research on the peoples of the area. Fourth, this document contains conclusions about the
vulnerability of the culture to loss of clean water and salmon resources in the Bristol Bay area.
Between us (Boraas and Knott) we have 48 years of research, teaching, and collaboration with
Alaskan tribes, and that experience is reflected in this study.
As a foundation for this research, all of the federally recognized tribes in the watersheds
were contacted through the Environmental Protection Agency's Tribal Trust and Assistance Unit
in Anchorage following government to government protocols. Since one of us, Alan Boraas, is
an Honorary Member of the Kenaitze Indian Tribe, a letter of introduction from the Kenaitze
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Tribe to village councils was included in the government to government packet following village
protocols (See Appendix 1 which also includes the initial statement of methodology). We
selected seven villages in which to conduct interviews: New Stuyahok, Koliganek, Curyung
(Dillingham), Nondalton, Pedro Bay, Newhalen, and Diamna. Time and funding prevented us
from conducting interviews in Igiugig, Levelok and Ekwok. Kokanok and Port Alsworth did not
respond to the government to government request to conduct interviews.
Table 1 Number of Interviews per Village.
Village
Curyung (Dillingham)
Iliamna
Koliganek
Newhalen
New Stuyahok
Nondalton
Pedro Bay
Total
Males
7
1
5
5
5
4
2
29
Females
0
3
5
6
2
6
2
24
Total
7
4
10
11
7
10
4
53
We interviewed 53 Elders and culture bearers, people whom the village councils or tribal
governments recognize as authoritative sources of information about subsistence, traditional
ecological knowledge, social relations and spiritual aspects of their culture. The village-selected
interviewees consisted of 24 females and 29 males (see Table 1) and ranged in age from mid-
twenties to a man reportedly in his nineties. Most, however, were in their forties or older due to
the intentional weighting toward village-selected Elders and culture bearers. We were not
consulted in the selection of specific interviewees and were assisted by a tribal employee or a
village council member who arranged the time and place of the interview (see Appendix 1,
Methodology). The interviews took place in public tribal or community centers or private homes
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because from the standpoint of the interviewees they are safe, non-threatening places in which to
discuss important cultural matters. We normally interviewed two to four individuals at any one
time but some sessions included as many as six and one was a single interviewee. The interview
session lasted about two hours with a short break. Interviews followed a standard semi-structured
interview process in which a set of questions guided the interview but interviewees were free to
add additional information or perspective, in some cases delving into topics not covered by the
original question. The questions were specifically designed not to be answered briefly but to
probe the subject and allow interviewees to describe cultural structures which for the most part
were familiar and obvious to local villagers, but not commonly understood to others, particularly
those outside the state. If a response was brief we would respectfully clarify or amplify upon the
question to generate a more complete narrative. Interviewees were told they did not have to
respond to a question if they chose not to, although none did. If an interview session exceeded
two hours we occasionally eliminated some questions. If the topic of a question had already been
covered in a previous discussion we eliminated the question. Consequently, not all interviewees
responded to every question. Regularly one person would respond and others would nod
agreement. Since the questions dealt with a cultural standard, there were few alternative points of
view. Some of the interviewees chose to speak in Yup'ik, in which case an interpreter was
present to translate the question into Yup'ik and the response into English. None chose to speak
in Dena'ina. Many Elders "think" in their Native language which we encouraged because
responding in the traditional language generates more accurate and nuanced responses to
questions about culture.
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Figure 1. Nondalton, August 17, 2011
We digitally recorded the interviews and, in the Kenai Peninsula College Anthropology
Lab, made transcriptions from the recordings including both responses to our questions and
additional perspective provided by the Elders or culture bearers.
The interview questions revolved around the theme of, "How are salmon and other
stream-based resources and water important in your lives?" The questions involved the topics of
nutrition, subsistence, social relations, spirituality and beliefs. In addition a final question was
asked: "is there anything you would like to add, or is there anything you would like the
Environmental Protection Agency to know about the situation in your village." The interview
questions are listed in Section III.A.
The transcribed interviews were lumped into a single Microsoft Word document and the
lumped document was searched for key words related to the sub-headings of this report using the
powerful search feature of Microsoft Word 2010. In this way we were able to capture responses
both to the theme of the question we asked and to that theme that might have been discussed by
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interviewees in the context of a question related to a different topic. In this document responses
of Elders and culture bearers titled "Voices of the People," reflecting both the consensus among
those interviewed and the rare deviations from consensus appear in italics before the
anthropological discussion of each section. By the standards of highly pluralistic modern
America, the Yup'ik and Dena'ina villages of Southwest Alaska are culturally much more
homogenous, consequently the narratives reflect that homogeneity. "Voices of the people"
statements were selected through the search process described above because they were concise,
clear, and reflected the intent of the speaker in the context of their broader narrative. The English
response or translation is transcribed "as is" with no grammatical modification; readers must
understand that for some, English is a second language and imperfect English grammar is not to
be construed as imperfect or naive thinking. Following University of Alaska Institutional Review
Board Standards, to protect individual identity of the interviewees, each Elder or culture bearer
has been designated by a code, using an "M" or "F" for "male" or "female" and a number, along
with the date of the interview.l Only we, the interviewers, know the names of the interviewees.
All deviations from consensus have been included in the qualitative "Voices of the
people" responses. In addition, the entire 500 page typed narrative was assessed from a
favorable/unfavorable or agree/disagree standpoint to give a sense of the degree of conformity to
a response. These results, along with the interview questions, are portrayed in Section III. A. and
1 Funding for this project was administered as a contract through the University of Alaska
Anchorage/Kenai Peninsula College and came under Institutional Review Board (I.R.B.)
auspices since it involved human subjects. The UAA I.R.B. reviewed and approved the
methodology, consent forms and research design of this project. I.R.B. stipulates protection of
the identity of human subjects, consequently the names of the participants of this study and not
revealed. Signed consent forms are held by the researchers.
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referenced throughout this document to give a quasi-numerical sense of the culture standards of
the Nushagak and Kvichak drainages.
B. Villages, Population, and Ethnicity
In the 2010 United States Census, the 13 communities of the study area had a total
population of 4118. Table 2 describes the population characteristics of the 13 villages and towns
located in the Nushagak and Kvichak River drainages.
Table 2. Census of the Towns and Villages of the Nushagak and Kvichak River Drainages,
1980 to 2010. (data from U.S. Census, Alaska; Alaska Community Database)
Watershed
Nushagak
River
Kvichak
River
Community
Dillingham
Ekwok
Koliganek
New Stuyahok
Portage Creek
Igiugig
Iliamna
Kokhanok
Levelock
Newhalen
Nondalton
Pedro Bay
Port Alsworth
1980
Pop.
1563
77
117
331
48
33
94
83
79
87
173
33
22
1990
Pop.
2017
77
181
391
5
33
94
152
105
160
178
42
55
2000
Pop.
2466
130
182
471
36
53
102
174
122
160
221
50
104
2010
Pop.
2378
115
209
510
2
50
109
170
69
190
164
42
159
% Alaska
Native,
2010
55.9
90.4
95.7
93.5
50.0
40.0
54.1
80.0
84.1
80.0
63.4
66.7
21.4
41 18 Total
20 10 Population
Ethnic
Majority
Yup'ik
Yup'ik
Yup'ik
Yup'ik
Yup'ik
Yup'ik, Alutiiq/
Caucasian
Dena'ina
Yup'ik/Dena'ina/
Alutiiq
Yup'ik
Yup'ik
Dena'ina
Dena'ina
Caucasian
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4500
4000
3500
3000
2500
2000
1500
1000
500
1980
1990
2000
2010
Port Alsworth
Pedro Bay
Nondalton
iNewhalen
Levelock
IKokhanok
Iliamna
llgiugig
I Portage Creek
iNewStuyahok
IKoliganek
I Ekwok
Dillingham
Figure 2. Population Change for the Study Area: 1980 to 2010. Data from U.S. Census.
Figure 2 indicates the population of the study area grew substantially from 1980 to 2000
and remained stable between 2000 and 2010. 1980 to 2000 village population growth is
probably due to post-ANCSA changes in land-ownership and is related to a similar phenomenon
throughout Southwest Alaska (Fienup-Riordan 1994:39). The population of individual
communities can vary considerably; in small populations only a few large families moving in or
out can change the overall population considerably. Of the 13 communities, four are anomalous:
Dillingham, Port Alsworth, Igiugig, and Iliamna. Dillingham has, by far, the largest population
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in the area (2,329 in 2010) and is a regional center with an economy based on the Bristol Bay
commercial fishing industry, as well as government services, transportation, and professional and
business services (Alaska Community Database). Dillingham has a small branch of the
University of Alaska, a museum, and Alaska Department of Fish and Game (ADFG) offices, as
well as several stores, churches, hotels, and other institutions typical of mid-sized Alaskan
towns. Dillingham, however, is 55.9% Alaska Nativemainly Yup'ikand the Curyung Tribe
and Bristol Bay Native Corporation and associated agencies are a significant presence (Alaska
Community Database).
Port Alsworth is only 21.4% Alaska Native and thus does not have the majority or near-
majority Alaska Native population that other villages in the study area have. The non-Alaska
Native population is primarily associated with two institutions. The Lake Clark National Park
and Preserve, which surrounds Lake Clark, has its regional headquarters in Port Alsworth.
Because of the park, a number of eco-tourism guides unaffiliated with the park but using its
resources are headquartered at Port Alsworth. The Tanalian Bible Camp and associated
ministries, loosely connected to Samaritan's Purse, a national fundamentalist Christian ministry
directed by Rev. Franklin Graham, is also located at Port Alsworth. Yup'iks who relocated to the
area in 1944 (Gaul, 2007:60-61)) account for most of the town's Alaska Native population and
make up its ANCSA-based village corporation, Tanalian Inc. (Port Alsworth is well within
traditional Dena'ina territory). Igiugig has a substantial number of guided sport fishing and sport
hunting operations that have recently moved into the village or near the village which accounts
for the relatively large non-Alaska Native percentage of the population. Iliamna, a traditional
Dena'ina village located on Iliamna Lake, is also a growing center for guided sport hunting and
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fishing. It has also become a staging area for exploration and other activities associated with
proposed copper/gold porphyry mines in the area. Consequently, Iliamna has a proportionately
larger non-Alaska Native population than most other villages in the area, although the Alaska
Native population (54.1%; Alaska Community Database) outnumbers other ethnic groups, and is
still the dominant ethnic group.
The remaining study area communities are Yup'ik or Dena'ina villages with close
connections to traditional practices. They are relatively small, with populations ranging from 510
(New Stuyahok) to 42 (Pedro Bay) (Portage Creek, population 2, is reportedly seasonally
occupied as of 2011, according to interviewee M-26), and from 93.5% Alaska Native (New
Stuyahok) to 67% Alaska Native (Pedro Bay). Most have a single church (Russian Orthodox), a
public school, a health clinic, an airstrip, a small general merchandise store, a post office, a
tribal center or village corporation center, city or village corporation offices, a landfill, cemetery,
and fuel storage tanks (Alaska Community Database and observations).
There are community health aides in the villages of Koliganek, New Stuyahok, Ekwok,
Igiugig, Levelok, Kokhanok, Nondalton, and Pedro Bay (Bristol Bay Area Health Consortium,
BAHC 2006) and some also have dental aides. The clinics are connected via internet to
consulting physicians and the Alaska Native Hospital in Anchorage. New Stuyahok and
Newhalen have completed the Rural Utility Business Advisor (RUB A) process in order to join
the Alaska Rural Utility Collaborative (ARUC) and have a municipal water system
(http://www.anthc.org/cs/dehe/sustops/). Many of the villages are being connected to high-speed
fiber-optic internet connection. Nearly 100% of the population has access to some improved
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sanitation, and 100% of the population has access to the abundant fresh, clean water of the rivers
and lakes.
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II. CULTURAL AND HISTORICAL BACKGROUND
A. Pre-Contact Bristol Bay
1. Voices of the People
Salmon and fresh water has been the lifeline of the people here for thousands of years. If you
look at the water, that is why fish and game has survived so well here, because we have such
clean water. M-62, 9/16/11
[If the salmon were to be impacted], it would stop 10,000 years 'plus tradition, culturally and
spiritually for my people; not only my people, all the other communities and villages in this
region will go away. We would cease to exist. We can't go anywhere. Where are we going to go?
M-33, 8/18/11
Freeze drying is not a new thing. That's been going on with my people for over 10,000 years,
eating freeze dried food. M-33, 8/18/11
There's 10,000 cache pits [at the Kijik archaeological site on Lake Clark] and they are still
counting; over 200 houses, which are huge. So it was pretty big. M-29, 8/17/11
My father, he usually keeps fresh salmon. He would dig a pit and take the topsoil off; dig it out
lay some grass on the bottom and on the side. Then take the salmon, lay them in the pit until he
filled it up. Then he would put grass on top of it. Then he would lay gravel right on top of it, and
he would mark each corner for winter time. Put poles on each corner so he could find where he
buried his salmon. And in the winter time, if he wanted salmon, he would take his axe and cut out
apiece of the soil and dig from there. That was his freezer. That is how my dadwouldkeep
salmon. M-54, 8/20/11
2. Introduction
The pre-contact history (prehistory) of the Bristol Bay drainage is not as well
documented as in other parts of Alaska. The archaeological work is largely due to five projects.
In the 1960s James Van Stone conducted an archaeological survey of the Nushagak River as part
of ethnohistoric research (VanStone 1967); B.I.A. archaeologists have conducted archaeological
surveys in connection with Native Allotment assessments; Lake Clark National Park has
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conducted various survey projects on the Mulchatna River and areas above tree line; the Pebble
Partnership has contracted for archaeological surveys on the footprint of a proposed Pebble Mine
site; and the Alaska Office of History and Archaeology has conducted or required pre-
development archaeological surveys on proposed airstrips and other improvements and
conducted town-site surveys. Within the study area there are a total of 228 historic and
prehistoric sites listed on the Alaska Heritage Resources Survey (A.H.R.S.), the state's database
for officially designated sites. To better understand the patterns of culture change and establish
the time-depth of salmon use in the Nushagak and Kvichak River drainages one of us (Alan
Boraas) generated a database of the 228 sites and from that developed a prehistoric cultural
chronology of which the last 4000 years are depicted in Figure 3.
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Date 1
1000 BP
(-A.D. 1000)
2000 BP
(-A.D. 0)
3000 BP
(-1000 B.C.)
4000 BP
(-2000 B.C.)
Geographic Area within the Nushagak'Kvichafc Watershed
Nushagak
River
K::7cric Yup'i
Pre-Contacl
Yup'ik
Norton
Tradition
(interior)
Kvtchak
Rjver
HiKork Yiip'i
Pis-Contact
Yup'ik
Norton
Tradition
(interior)
Arctic Small
Tool Tradition
Tliamna
Lake
Hi;[. Yjp'ikDer:.
Pra-Contact
Yiqj'ik&
Dena'ina
Norton
Triiiinon
(ulterior)
Arctic Small
Tool Tradition
Mukhatna
River
Si:!. Y^-p'ik.-Dezi.
Sedeniar>'
Dena'ina
*r
Lake
Clark
r.'.'J. I'SLi'oa
Sedeatar," g
D^La'ma
Norton
Tradition
(interior)
Arctic Small
Tool Tradition
Alpme
Areas
Hi;t. Ds^a'ina
Sedentary
Dena:ina
Arctic Small
Tool Tradition
Salmon Cultures No Da:a or not definitive
Radiocarbon Dares
. Probable
BP dates in
Radiocarbon Years
Figure 3. Cultural Chronology of Nushagak and Kvichak River Drainage Salmon-Based
Cultures. From Alaska Heritage Resource Survey database. By Alan Boraas
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The "BP" (Before Present) of the y-axis of Figure 3 is in uncalibrated radiocarbon years and an
approximate B.C./A.D. date is indicated.2 AHRS site data was assembled for six regions (Figure
3) within the Nushagak and Kvichak River drainages, including:
The Nushagak River from its mouth to headwaters.
The Kvichak River, including nearby archaeological sites in the Alagnak River drainage.
The shoreline of Hiamna Lake and the lower Newhalen River.
The Mulchatna River, upstream to Bonanza Creek.
Lake Clark, Sixmile Lake, and the Upper Newhalen River.
Alpine areas above tree line north of Iliamna Lake and west of Lake Clark.
3. Pre-Contact Salmon Fishing Cultures
The study area was occupied as early as 8,000 BP by core and microblade makers of the
Paleoarctic tradition (with two Putu-like fluted points coexisting with microblades at one site,
XHP-00430 extending the possible time range to 12,000 BP). Subsequently, archaeological
cultures of the Northern Archaic and Ocean Bay traditions occupied the area. None involved
intensive salmon fishing as indicated by AHRS records. The Paleoarctic and Northern Archaic
sites are associated with Athabascans (Boraas 2007: 34-7) and establish a time-depth for the
Dena'ina or proto-Dena'ina in the study area.
2 The deviation between calibrated calendar years and uncalibrated radiocarbon years becomes
significant before 1500 B.C. By 2000 B.C. uncalibrated radiocarbon years are ~ 400 hundred
years old (http://www.radiocarbon.com/calendar-calibration-carbon-dating.htm).
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As described below, archaeological records indicate Yup'ik or proto-Yup'ik people have
been fishing for salmon for at least 4,000 years (Figure 3 and Table 3) and may be genetically
related to earlier Siberian salmon fishers. Salmon fishing first appears with the Arctic Small Tool
tradition (ASTt) (see Figure 3) and Table 3 is a list of ASTt sites in the study area. ASTt
cultures are widespread in western and northern Alaska where the site data indicates the
existence of interior nomadic hunters (primarily caribou) or coastal sea mammal hunters. In the
Bristol Bay drainage, three village sites, evidenced by ASTt-style houses and artifacts, are found
on the Kvichak River. Five alpine sites (artifacts only) indicate hunting above tree line. The
houses are permanent structures, generally measuring four meters on a side, indicative of
sedentary or semi-sedentary people and are located adjacent to salmon spawning streams. The
ASTt site at Igiugig (ILI-00002), where the Kvichak River flows out of Iliamna Lake, is an
example of such a site (Holmes and McMahan, 1996).
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Table 3. Arctic Small Tool tradition sites in the Study Area. Compiled From Alaska
Historic Resources Area Files
ARCTIC SMALL TOOL TRADITION AD 200 to 1800 BC
Area
Nushagak R.
Iliamna Lake
Alpine
Alpine
Alpine
Alpine
Alpine
Kvichak
Kvichak
Kvichak
Kvichak
Kvichak
AHRS Site
NAK-00018
B
ILI=00035
ILI-00201
ILI-00205
ILI-00193
ILI-00219
ILI-00218
DIL-00088
DIL-00170
ILI-00002
ILI-00072
ILI-00206
Characteristics
cores and microblades
Lithic tools
Microblade core
Microblade core
Lithic camp: microblades, side
blades, end scrapers, knives.
Microblade core
Microblade core
Village, sedentary houses; C14
Date, 3580+/-150;
Village; Brooks River Gravel
Phase
Cores, microblades, burins,
notched stones, 4000 artifacts;
Brooks River Gravel phase, ca.
1800 BC to 1100BC
3350+/-60 BP radiocarbon date,
possible Norton component
Microblades and other lithics
Village site
Houses
19
2
1
Anadromous salmon remains, while not common, occur in ASTt sites (Dumond, 1984),
suggesting salmon were a significant subsistence human resource in riverine and lacustrine areas
of southwest Alaska. The lack of abundant salmon bones in ASTt sites may be due to small
populations of salmon, decomposition of the relatively delicate bones, or the practice of
returning salmon bones to the watersimilar to ethnographic Yup'ik and Dena'inathereby
contributing to marine-derived nutrients important in salmon habitats. Further research is
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necessary to clarify this point. The fact that one site (DIL-00088) contains 19 sedentary houses
and is located along a salmon stream indicates salmon were a primary resource.
Analysis of human hair from a 4,000-year old ASTt site in Greenland places the
mitochondrial DNA (mtDNA) in the D2c haplogroup reflecting Siberian origins (Gilbert et al.,
2008). Today, haplogroup D2c is present, but haplogroup A is dominant among Yup'iks;
haplogroup A also has Siberian origins where researchers place its origin as early as 7,000 years
before present (Rubicz et al., 2003). Both haplogroups indicate that the time-depth of Yup'ik
people in southwest Alaska is at least 4,000 years and that they derive from Siberian origins,
where their ancestors were also potentially salmon fishers. As described in the section on
nutrition (III.C.3.), evidence is building that Yup'iks are biologically adapted to salmon and
4000 years is the temporal context in which that evolution took place.
In all but the Mulchatna River and alpine areas where evidence has yet to be found, the
Arctic Small Tool tradition is followed by a well-developed salmon culture, the Norton tradition,
dating from -300 B.C. to A.D. 1000 (see Figure 3; Table 4). Like ethnographic Yup'ik, the
Norton tradition has both a coastal and interior subsistence orientation. The coastal Norton
tradition is found in sites as far north as Cape Denbeigh and relied primarily on marine mammals
(Dumond 1984: 99-101). The interior Norton tradition sites, such as those in the study area on
the Nushagak and Kvichak Rivers and Lakes Iliamna and Clark, had a salmon-oriented
subsistence culture based on the following evidence: archaeological features, mainly houses,
similar to those at ethnographic Yup'ik salmon fishing sites: large sedentary villages, villages
located adjacent to salmon fishing locations, and net fishing artifacts. Riverine Norton tradition
sites are similar to ASTt sites in that they consist of large, permanent houses located on salmon
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streams. One large Norton tradition site on the Kvichak River (DIL-00161) consists of 34 to 45
houses representing a population sustainable only through the availability of abundant resources
such as anadromous salmon. In addition, the artifact inventory for the eight Norton village sites
in the study area (see Table 4) contains notched stones that were used as net weights, similar to
the lead line of a modern net (Dumond, 1987:11). In addition to dwelling houses, Norton sites in
southwest Alaska contain large structures indicating a kasheem or kazigi, (local pronunciations
vary), a men's house also found among pre-contact and early historic Yup'ik villages. These
finds indicate that the Bristol Bay drainage Norton culture were Yup'ik or proto-Yup'ik speakers
and relied on salmon as their primary subsistence food.
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Table 4. Norton tradition sites in the study area. Compiled from Alaska Heritage
Resources Survey data by Alan Boraas.
NORTON TRADITION AD 1000 TO 300 BC
Area
Kvichak
Kvichak
Kvichak
Kvichak
Kvichak
Kvichak
Iliamna Lake
Iliamna Lake
Iliamna Lake
Iliamna Lake
Lake Clark
Lake Clark
AHRS Site
DIL-00161
DIL-00174
DIL-00175
DIL-00229
ILI-00073
DIL-00207
ILI-00056
ILI-00127
ILI-00128
ILI-00098
ILI-00012
XLC-
00086
Characteristics
Prehistoric village (6100 artifacts)
1760+/-40BP
Two large house depressions; Smelt
Creek Phase
1920+/-40
Village site, artifacts, pottery; Norton
Brooks River Weir and Brooks River
Falls phases, 1830+/-40BP
Prehistoric Village
Village site, Pottery,
Village, 43 house depressions; lithics
and ceramics
Village, C14 date 860+/-60
Pottery and stone beads
Weir, Early Norton
Village, cache pits no houses
apparent on surface, fiber pottery
Village
Bifaces, scrapers, sideblades, fiber
pottery.
House
s
34-45
2
8
1
4
43
12-15
12
It is not clear how long the Dena'ina have been salmon fishers, but about A.D. 1000, the
Dena'ina of the Mulchatna River and Lake Clark areas developed a method to catch salmon
using weirs and began storing salmon in underground cold storage pits called ebien tugh (Kenai
dialect) that appear in the archaeological record (Boraas 2007). Salmon storage technology
spread to Iliamna Lake, Cook Inlet, and the Susitna and middle Copper River areas (Boraas,
2007). A proliferation of Dena'ina sites65 have been found to date far more than any other
pre-contact periodoccurs in the study area, dating to just after A.D. 1000 (Table 5 and Lynch,
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1982). Forty-one sites are village sites (not necessarily occupied simultaneously) and the Kijik
Site, XLC-00084 and associated sites, are among the largest in Alaska for the prehistoric period.
We can conclude that weir fishing and the underground cold storage technology described in the
pre-contact culture section (II.C.2.) below was an extremely successful adaptation.
Table 5. Pre-Contact or Early Contact Period Dena'ina Sites in the Study Area. Compiled
from Alaska Heritage Resources Survey data by Alan Boraas.
SEDENTARY DENA'INA AD 1000 TO AD 1800
Area
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Mulchatna
River
Iliamna Lake
Iliamna Lake
AHRS Site
XLC-00072
XLC-00076
XLC-00078
XLC-00074
XLC-00075
TAY-00046
TAY-00026
TAY-00030
TAY-00027
TAY-00031
DIL-00200
DIL-00201
ILI-00029
ILI-00046 B
Characteristics
Village
Village
Cache pits
Village, Dena'ina
Village, Dena'ina
Cache pits
Cache pits
Cache pits
Cache pits
Cache pits
Cache pit
Cache pit
Fish camp
Village Complex
Houses
1
2
1
1
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Iliamna Lake
Iliamna Lake
Iliamna Lake
Iliamna Lake
Iliamna Lake
Iliamna Lake
Iliamna Lake
Iliamna Lake
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
ILI-00019
ILI-00135
ILI-00021
ILI-00020
ILI-00001 A
ILI-00047
ILI-00049
ILI-00018B
XLC-00048
XLC-00057
A
XLC-00102
XLC-00167
XLC-00166
XLC-00094
XLC-00165
XLC-00164
XLC-00155
XLC-00163
XLC-00162
XLC-00101
XLC-00100
XLC-00099
XLC-00084
XLC-00092
XLC-00090
XLC-00091
XLC-00093
XLC-00021
XLC-00020
XLC-00012
XLC-00013
XLC-00159
XLC-00158
XLC-00104
XLC-00157
XLC-00156
Village site
Cache pit
Village
Village, houses undetermined
Village
Cache pits
Village
Village 560+/-60 BP
Cache pits
Prehistoric Village
Village
Village
Village
Village
Village
Village
Village
Village
Village
Village
Village
Village
Village (possibly two sites)
Village
Village; C 14 BP 300+/-60
Village
Village
Cache pits
Village
Village
Trapper cabin
Village
Village
Village
Village
Village
3
nd
nd
5
4
nd
30
10
5
2
19
2
2
5
1
2
11
14
2
95
13
10
4
1
2
2
3
2
1
O
12
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Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Lake Clark
Mulchatna
River
Iliamna Lake
XLC-00105
XLC-00088
XLC-00083
XLC-00097
XLC-00098
XLC-00003
XLC-00004
XLC-00008
XLC-00250
XLC-00133
XLC-00134
ILI-00087
XLC-00096
XLC-00249
XLC-00107
DIL-00150
ILI-00031
Village
Cache pits
Village
Village, 1 house
Village
Cache pits
Cache pits
Village
Cache pit
Village
Village
Cache pits
Village
Cache pits
Village
Cache pits
Village
10
6
5
4
3
1
1
1
5
B. History and Culture of the Yup'ik Area
1. Voices of the People
We want to give to our children the fish, and we want to keep the water clean for them ....It was a
gift to us from our ancestors, which will then be given to our children. F-69, 9/18/11
When I was a little girl they had no Snowgo 's [snowmachines], they had no Hondas [Four-
wheeler all-terrain vehicles]. We live up river and they fished all the time. In wintertime they
fished under the ice. They travel with dog teams. My Dad would take me out ice fishing. I used
to be scared of those pikes. I don't know how old I was. That's the only thing they do is try to
catch fish, summer time nets, and winter time they do ice fishing. That's how they pass it on
down. They subsistence fish, usually they travel with dog teams, that's what they did, and that's
how come those people were healthy. They walked, and walked, they worked from morning until
they go to bed. That's how come they were healthy. They eat their fish, they go get wood with
the dog team, they hunt with their dog teams, and they travel to village with their dog team.
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People walk and they eat that fish. That's what makes them live long and healthy, I noticed that.
F-23, 5/18/11
All we have is use the salmon, salmon all the time. The old people tell us you guys have only one
salmon season you guys got to catch it. If you don't catch it you won't have much in the winter,
long winter. F-41, 8/19/11
When you look at the map and where the old villages were they were there because of the
salmon. You go to Igiugig and ?, andPort(?), Levelock, South Levelock andDillingham... all
those villages. Site selection of those communities was very important and it was because of the
production of subsistence foods at each of those sites processed. Most of those produced salmon
in addition to /other foodsy, for example you go to the village ofManokotuk, and it is rich in
berries. If you go to the upriver villages they are rich in caribou and moose and other resources.
Each village was selected by the folks... because of their subsistence resources. M-61, 9/16/11
My father along with other people was very active in fisheries politics. Bristol Bay used to be
controlled by Brindle which was a big cannery superintendent and what he said was law of the
land. Fish and game used to listen to those big processors. One time my dad was talking to a
group Truman Amberg, Joe McGill, Joe Clark from Clark's Point, saying we got to go on strike
this year. I think it was Joe McGill said we 're not going to get any more money [father's name].
Why are we going on strike? You know we are just going to end up sitting on the beach. Dad
says we got to let the fish pass. What that meant was we needed more fish up the river spawning
so we would have better seasons later. Then a group of locals said okay we 're going to strike but
don't tell the processors we or en't striking for more money. Tell them we want more money we
know they 're not going to give it but we will get more fish up the river because the Japanese
decimated our runs in Bristol Bay in the '60 's and 70 's. We had to build our runs back up, M-60,
9/16/11
Like before, you know a lot of people used to put up a lot offish 3000, 4000, 5000 fish. They used
to have a lot of dogs while they were living that is how they try the tradition they have. They used
to hook up their dogs and go wherever they wanted to go. They used to put up a lot offish to eat.
When they get moldy they just wipe it off and eat them. That is the way it was in my living days.
Nowadays people when it is moldy they throw them away, that is the way of life now. You can't
do that anymore. M-49, 8/20/11
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2. Introduction
Perhaps as a result of the relatively recent occurrence of contact with non-Natives, the
Yup'ik have retained their traditional culture and language, ecological knowledge and practices,
social systems, and spirituality, to as great or a greater degree than any other Alaska Native
populations. Where they have adopted non-Yup'ik traditions, such as Russian Orthodoxy, they
have blended their own practices and beliefs with the introduced practices to create a new belief
system that retains the Yup'ik culture as a whole.
3. Pre-Contact Culture
An Eskimo-speaking people have been living in the region for at least 4,000 years as a
recognizable salmon culture, at least as far back as the Norton tradition and Arctic Small Tool
tradition.
The Yup'ik of the Nushagak, Kvichak and lower Mulchatna Rivers historically were
organized in bilateral extended families of up to about thirty people settled in permanent and
semi-permanent villages. Many of the villages contain a kashgee, or men's house, and are
relatively small, averaging five to six houses per village in the 12 pre-contact villages for which
there is house data (see Table 5). Historic Yup'ik village sites, of which 21 are currently
documented, average between 8- 9 houses per village. Today there are only four or five modern
Yup'ik villages along the Nushagak River (Dillingham, Ekwok, Koliganek, New Stuyahok, and
possibly Portage Creek; see also Table 1) and, except for seasonally occupied Portage Creek,
they are much larger in population than their historic or pre-contact counterparts.
The wetland landscape is not easy to traverse, except by river, or in the depths of winter
when all is frozen. The abundance offish and game in the Bristol Bay region allowed the Yup'ik
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to stay within a relatively fixed range, although they moved throughout their range seasonally
from a base village, to hunt, gather, and participate in summer fish camps. The extended families
practiced food sharing and generalized reciprocity, both within and between families. Most
larger villages functioned as independent and self-sufficient social units, and people married
within the village or nearby villages. Sometimes fluctuations in game or fish availability caused
groups or individuals to travel from one region to another. Large disruptions to the population
did not occur until epidemic diseases arrived with European explorers. These diseases devastated
whole populations, decimated villages, undercut social distinctions, and wiped away some of the
boundaries over which the earlier bow and arrow wars had been fought (Fienup-Riordan, 1994).
These population changes resulted in shifts in salmon harvesting, when population remnants
regrouped by joining other villages.
Historically, including after contact, in the winter villages the men and boys older than
seven or eight lived in the qasgiq, the large communal men's houses, while women and girls
lived in a smaller house called an ena, both built from sod and driftwood. During the winter, the
community came together for dances and storytelling, but otherwise, men and women kept in
their separate groups and worked to do gender-specific chores. Men, for example, repaired the
tools for hunting, while women sewed clothes as well as waterproof raingear to protect everyone
from harsh weather.
In the summer, everyone participated in harvesting salmon, whether net fishing, or
processing the fish in fish camps. Women dominated the work of processing in the fish camps.
Family groups might put up as much as 5,000 fish (personal communication to Catherine Knott,
Lena Andree, Yup'ik Elder, Dillingham; July, 2011), including fish for their dogs.
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The Yup'ik traveled to different subsistence sites either overland, by foot or dogsled, or
on the water, in vessels that ranged from small kayaks to larger wooden boats. Traditional
festivals during the year included the Bladder Festival, nakaciuryaraq, the Messenger Feast,
kevgiryaraq, and the Seal Party, uqiquryaraq. Food exchanges played an important part in these
festivals described below.
4. Post-Contact History and Culture (A.D. 1791 to 1935)
At the turn of the 19th century, the bilateral extended family, stretching over several
generations, still formed the basis of Yup'ik villages (Fienup-Riordan 1994). Winter villages
could be just one family, but ranged up to 150 to 300 people in some places. Families did not all
live together in one house; the winter villages had one or more qasgiq, or communal men's
houses, where men and boys over age 6 or 7 lived and worked together, telling stories, making
tools, and preparing for subsistence activities. In the ena, women, girls, and the youngest boys
lived in groups of up to a dozen, and the women taught the girls how to sew and cook. They
cooked the meals there, either in the entryway, or in a central fireplace. Each winter, for three to
six weeks, boys and girls would switch homes, and the men would teach girls survival and
hunting skills, while the women would teach the boys how to sew and cook (Fienup-Riordan,
1990).
The qasgiq also functioned as the communal sweat bath for the men. They would open
the central smoke hole, feed the fire until the heat was intense (possibly up to 300 degrees), then
bathe. Men sat in the sweat house in the order of their social status. The niikalpiaq, or good
provider, held a high social position and contributed wood for the communal sweat bath, as well
as oil to keep the lamps lit; he also played an important role in midwinter ceremonial
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distributions of food (Fienup-Riordan, 1994). There was competition between families to be the
best providers.
Contact between the Yup'ik of the Bristol Bay area and Russians or Americans was later
and more limited than in most of the rest of Alaska. The region was perceived to have few
resources worth exploiting, and the marshlands were difficult to traverse. While some Russian
explorers, traders, and missionaries persisted and made repeated contact with the Yup'ik
throughout the nineteenth century, they did not settle in the area in any numbers until the
twentieth century (VanStone 1967). As a result, the Yup'ik of this region, perhaps more than any
other indigenous peoples in Alaska, have retained much of their language and cultural traditions
to the present time.
When the Europeans came, they brought diseases, to which the Yup'ik and other Alaska
Native populations had no immunity. The first epidemic known to have occurred in the
Nushagak River region was before 1832, but there are no records of the number of dead. The
1838-1839 smallpox epidemic caused several hundred deaths in the Nushagak region and also
occurred in the Dena'ina territory. Vaccines were introduced in 1838, and some Yup'ik received
them, probably reducing the scope of the epidemic and subsequent outbreaks of smallpox. But
each year, while not necessarily counted as an epidemic period, brought more death and illness
to the region. Survivors were often weakened and succumbed later to other illnesses. VanStone
states that during this period "The specter of ill health and death was continually present among
the Eskimo population of all southwestern Alaska" (VanStone, 1967:100). The loss of population
(especially Elders), the disruption of families, the plethora of orphans, and subsequent
rearrangements of the social order created a social and cultural upheaval that the Yup'ik
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struggled to overcome. The European visitors and settlers may not have understood that what
they observed was not the way the Yup'ik had lived even a few short years before.
It is not certain when the first Russian visit to the Nushagak and Kvichak region
occurred, but in the early 1790s Aleksey Ivanov of the Lebedev-Lastochkin Company made an
overland journey to Iliamna Lake from Cook Inlet and then west into the Mulchatna and
Nushagak drainage. His guide was apparently Dena'ina because the place names, including
Dudna (spelled Tutna) the Dena'ina name for Yup'ik's (Downriver People), are Dena'ina,
(Chernenko 1967:9-10). During this early period the region was not well known to outsiders, but
the Russian-American company sent an expedition in 1818 to explore the territory north of
Bristol Bay. In the same year, the company established a post at the mouth of the Nushagak
River, the Alexandrovski Redoubt. Feodor Kolmakov, of mixed Russian and Native American
ancestry, was in charge; he established trade relations with the Yup'ik and baptized some of
them, spreading the influence of the Russian-American Company in several ways (VanStone,
1967:9).
In the summer of 1829, two minor Russian visits had major consequences for the Yup'ik.
Ivan Filippovich Vasiliev led an overland expedition to ascend the Nushagak River, and the
priest, Ivan Veniaminov, visited the redoubt. Veniaminov took away a permanent interest in the
Bristol Bay region and in the Nushagak station which carried over even into his later position as
Bishop. Vasiliev's exploration, in turn, established travel routes that were used by subsequent fur
traders (VanStone, 1967:11).
Christianity was introduced in 1818, at the time that Alexandrovski Redoubt was built,
but it was not until Veniaminov's arrival in 1829 that extensive missionary activity took off.
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Veniaminov was flexible in his approach to the Yup'ik and their traditional religion and
numerous conversions were registered in church documents. Veniaminov noted that "the
Nushagak River was for them [Yup'ik] the River Jordan" (cited in Barsukov, 1887-1888, vol.
2:37). In 1832 Veniaminov visited again and had a small chapel built. By 1842 there were about
200 converts at Nushagak, and in 1844 Bishop Veniaminov had a new church built. The church,
by 1879, was close to 2,400 members. Its success among the Yup'ik may have had much to do
with the flexibility of Veniaminov's approach toward them. Yup'ik people were not required to
fast and many indigenous customs were tolerated (VanStone, 1967:31).
Fur trading accompanied exploration, and sometimes incited it. By the 1840's contacts
between the Kolmakovski Redoubt, on the Kuskokwim, and Alexandrovski at Bristol Bay were
frequent. The company managers of the fur trade created toyons, designated local community
leaders, and rewarded them with silver "United Russia" medals and incentive gifts. These toyons,
motivated by their new prestige and the material rewards offered, then encouraged the members
of their social networks to trap more furs for the Russians (Van Stone, 1967:56). The process of
using village providers to convert the population into loyal company men and women to recruit
fellow villagers into exploiting and extracting the resources of their own region for external
benefit in a colonialist economic system has not changed in over a hundred years. The
researchers observe the practice has helped to dismantle the traditional ecological knowledge and
practices gained from the long indigenous history of subsistence-based culture.
Trade items included wool blankets, tobacco, beads, tent cloth, cast iron kettles, knives,
iron spears, steel for striking a fire, needles, combs, pipes, etc. (VanStone, 1967:56). While these
items did not immediately alter the deeper structures of the culture, the desire for them acted as a
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change agent among the population. Where before, access to status had been open to all, through
skills and responsible sharing with others, access to the time and materials for trapping, open to
fewer individuals, had the potential to change the social dynamics of the Yup'ik. The companies
allowed the Alaska Natives to purchase some items on credit; as debt mounted, some would be
unable to repay for years. After the Alaska purchase, the powerful Alaska Commercial Company
post at Nushagak maintained a trading post through the remainder of the nineteenth century
engaging in about $10,000 in fur trades annually (VanStone, 1967:56),
In the nineteenth century gold mining occurred but was economically unimportant
compared to other activities. In 1887-1888 the prospectors Percy Walker, Henry Melish, and Al
King placer-mined for gold in the Koktuli and Nushagak Rivers, and there was also placer
mining along the Mulchatna. In 1909 a group organized the Mulchatna mining district and
formed the Mulchatna Development Company in Seattle (VanStone, 1967:83). Their activities
were confined to the upper Mulchatna River in Dena'ina territory, and there was only a very
temporary influence of miners on the local Alaska Native population. One Elder (New Stuyahok
Interviewee in a non-recorded interview situation) told the story of his grandfather, who showed
him gold and told him that if he found rocks with gold in them to throw them away, because they
were bad. The grandson thought it was because it would cause social disruption by bringing
strangers to the area who would disrupt the land and the culture of the people. The Elder said he
had thrown a big chunk of gold away once, but he thinks he still knows where it is. The
experience of the Yup'ik people with larger mining corporations has been minimal. Fish have
been far more important both to subsistence and cash-based economies.
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By the end of the nineteenth century, Bristol Bay had become an important commercial
salmon fishing zone. The first salmon cannery, The Arctic Packing Company, began operation in
1884 at the village of Kanulik at the mouth of the Nushagak River (Troll, 2011:3). The fourth
cannery, built at Clark's Point in 1888, is now the oldest surviving cannery in the region (Troll,
2011:4). The commercial fishermen in Bristol Bay used wooden sailboats for drift gillnet fishing
for sockeye salmon and were mostly Italians, Scandinavians, and Finns, hired at Seattle and San
Francisco (Troll, 2011:10), although some Yup'ik also fished commercially including Lena
Andree, now an Elder from Dillingham who fished on one of the wooden sailboats with her
father in the mid-1930s. When World War II began and kept many of the European fishermen
from coming to Alaska to fish, the canneries "discovered that the Native Aleuts and Eskimos
were marvelous boatmen and seemed to have been born to sail," according to Al Andree (cited in
Troll, 2011:35).
The U.S. Bureau of Fisheries visited the Wood River lakes and Nushagak and Nuyakuk
Rivers, and, in 1935, the U.S. Geological Survey conducted the first survey of the region and
produced what would become, for decades, the standard reference for people not from the
region. For the Yup'ik, the Elders continued to convey their traditional knowledge of their
homeland, as they had for thousands of years (Van Stone, 1967). A crevasse of deepening
proportions opened between two contrasting interpretations of the landscape, that of the
outsiders, who saw the region as a land of resources to be exploited, and that of the indigenous
peoples, who saw the region as the sacred landscape of home, and whose culture and way of life
depended upon it.
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C. History and Culture of the Dena'ina
1. Voices of the People
We harvest [subsistence foods] three times for that one person: day of the burial, forty days
later, and then one year later. It is really significant, just for that one person who passed away;
we harvest from the land three times to honor and to pay our respects to ones who lost their
family member. That has been going on for over 10,000 years. M-33, 8-18-11
...from our ancestors, that is how we get all of our information to have fish. The way we put it;
the way we store it for us to eat. That is where we learned it. It is passed on from generation to
generation to have fresh fish. F-48, 8/20/11
/ always think that we are very, very, very lucky people. I know where I came from. I know who I
am. I know where I belong in this world. I know where my ancestors come from. I know the trips;
the walking, the hiking, I know the history of where they were. Every time I come into this part of
the country or fly over it, when I first see the Lake Clark area or coming from the south and see
Sixmile Lake, I know I'm home! F-32, 8/18/11
So the importance of this resource, specifically salmon, has a major impact on my people here.
That's the reason why we live here. We have sockeye salmon until March, when everyplace else
has no more. That's why my ancestors fought over this region... The reason why they 've been
here for so long is it's a healthy environment, and we have been kind of watching over it all these
years. My ancestors fought over it, and they won every battle. We beat the Russians two times. It
was musket against bow and arrow. So, you see, the importance of it has a really long history of
why it is like it is now. We took care of it. Not only that, we have shared with everybody in the
whole world./in reference to commercially caught salmon] M-33, 8/18/11
My Auntie [name] would say, "Don't forget how to live off the land" and I'd think, "Oh, we
could just go to the store and have microwave stuff. " She said, "One day in this world
something's going to happen where you guys are going to rely on living off the land, trapping off
the land. " Like we take things for granted now; we can go on an airplane and shoot a moose or
trap beaver or trap squirrels up on the mountain. We have to. We can't just forget our ways; how
to live off the land, because one day there's going to be something that happens in the world,
where we are going to have to learn to survive out here. F-32, 8/18/11
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But what the spiritual aspect of what they believed was strong... they had energy. Energy from
what they worshipped; everything living. M-33, 8/18/11
That is spring water [at Kijik]. It does not freeze. That is why you can go over there and get a
sockeye salmon in March; it might have a green head, and it's red, but it's still a sockeye
salmon. You can go over there on New Year's Day and get afresh sockeye salmon. F-33, 8/18/17
2. Pre-Contact Culture
Dena'ina origins are described in the section on Prehistory (II.C.2) and indicate the
Dena'ina have been operating as a culture for whom salmon is the primary resource since A.D.
1000. Much can be inferred about the pre-contact Dena'ina culture because of Cornelius
Osgood's (1976, originally published in 1937) comprehensive Ethnography of the Tanaina [sic].
Like the pre-contact Yup'ik culture, the Dena'ina pre-contact culture was sustainable and
egalitarian in terms of equitable access to resources. The fundamental food source was salmon,
but also included caribou, moose, bear, beaver, and other mammals and birds (Osgood, 1976:26)
and about 150 edible plants (P. Kari, 1987:60-188). For the pre-contact Dena'ina salmon were
caught in a number of ways, but primarily in weirs made of poles sunk into the bottom of a
stream and strung with a lattice-like thatch, allowing water to pass through, but trapping
migrating fish (Osgood, 1976:28). When they weren't fishing they simply opened a gate, and the
fish swam through to spawn upstream. To solve the problem of storing this food resource for
later use, the Dena'ina devised a simple but effective underground cold storage pit (Osgood,
1976:42). Two layers of birch bark, with moss in between, lined the pit, which was filled with
dried fish, layered with grass, during fall freeze-up. The frozen fish were eaten throughout the
winter and spring, until the next summer's salmon run. Like modern fish camps, traditional
Dena'ina fishing was an extended family operation. Everyone worked for, and received the
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benefits of, the clan-based family group.
Because of the stable salmon food resource and a means to preserve it, the Dena'ina lived
in sedentary or semi-sedentary villages of substantial log houses, usually spread out along a ridge
above a lake, a river side channel or a tributary to one of the major rivers (Osgood, 1976:55-62).
The married men of a village were members of the same matrilineal clan and their wives and
children were members of a different clan (Osgood, 1976:128-131). Within this family group,
connected by blood and marriage, and allied for economic purposes, various individuals
performed different assigned tasks. The Dena'ina called this group the nakilaqa (ukilqa in
Osgood) (Osgood, 1976:134) or clan helpers. The clan helpers recognized a chief, called a
qeshqa; in the Iliamna area the position was related to being a family head (Osgood, 1976:131-3;
Fall 1987:6-8). The qeshqa had numerous characteristics, among them wisdom, experience, and
generosity. He or she had three primary duties: first, to arbitrate and resolve disputes; second, to
care for the elderly and orphaned; and third, to assure the survival of the clan helpers through the
equitable distribution of food. Regarding the latter, the qeshqa controlled the foods gathered,
processed, and stored by the clan helpers and had authority to redistribute the food (mainly
salmon) back to people throughout the winter on an as-needed basis.
This system provided a safety net. Each qeshqa had a partner in a distant village, called a
slocin. If one village ran low of food, the qeshqa could request aid from his partner, who would
divert some of his village's food resources to the needy village. The second qeshqa would be
willing to do this because, at some point, his village might be short of food, and the partner he
helped would return the favor.
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3. Post-contact History and Culture
In the study area Dena'ina territory includes the Kvichak drainage of Lake Clark, the
Newhalen River and the west half of Lake Iliamna. Today, the Dena'ina villages in the
Kvichak/Iliamna drainage are Nondalton, Iliamna, and Pedro Bay; Kokhanok is mixed Dena'ina
Alutiiq, and Yup'ik. This brief history is germane to the project because it establishes: 1) the
Dena'ina repelled Russian colonization maintaining population superiority in their homeland to
this day: 2) they adopted Russian Orthodoxy which ritually incorporated traditional viewpoints
of a symbolic relationship of people to the land, and, 3) they began to have economic ties to the
Bristol Bay salmon canning industry. Through it all the people retained a strong subsistence
lifestyle.
During the late eighteenth century, two Russian trading companies, the Shelikhov
Company and the Lebedev Company, occupied Dena'ina territory, focusing primarily on the
Cook Inlet region but extending into Iliamna Lake. The Lebedev established a post at Pedro Bay,
on Iliamna Lake, in the 1790s (Ellana and Balluta, 1992:61). About 200 Russians occupied Cook
Inlet and the Iliamna Lake area during the late eighteenth century; by the turn of the century,
their presence had shrunk to a small handful through a complex series of events involving attacks
and counter-attacks as outlined by Boraas and Leggett (in press, 2012). As a result of hostilities
the Russian Lebedev Company left Alaska in the spring of 1798, and subsequent Russian
presence in Dena'ina territory was minimal.
In 1838 a terrible smallpox epidemic decimated the Dena'ina (and most other Pacific
coastal Alaska Natives). Where there are statistics, such as for the Kenai River drainage, about
half the overall population died in two years (Fedorova 1973:164) and, although there are no
specific statistics for the Lake Clark and Iliamna, it is likely the situation was tragically similar in
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the study area. Traditional shamanic practices were ineffective against smallpox and, after 1840,
many Dena'ina were baptized as Russian Orthodox, (Townsend 1981:634-6), accepting the
church's explanation for the epidemic as "God's will" (Boraas and Leggett in press, 2012). In
1853 the Orthodox Church undertook an inoculation program, vaccinating baptized Dena'ina
against smallpox, and an Orthodox Church was built at Kijik in 1884 (Ellana and Balluta,
1992:63). It is probable that by the early twentieth century, most Dena'ina in the Diamna/Lake
Clark area were baptized as Orthodox.
As summarized by Karen Gaul (2007:48) salmon canning in Bristol Bay emerged as a
major industry in the late 1800s. Unregulated Bristol Bay canneries regularly blocked the mouth
of the Kvichak and Nushagak Rivers to harvest salmon; consequently, there were years when
there was little escapement into the rivers, creating extreme hardship for the upriver Dena'ina
and Yup'ik subsistence communities. Starting in the early 1900s, men from the inland villages
traveled to the coast to work seasonally in the commercial fishery, as many still do today. The
fur trade was a second non-subsistence occupation, providing cash for food, guns and
ammunition, traps, cloth, and other items, but commercial salmon fishing remained the primary
source of money for most indigenous families and supplemented subsistence activities (Gaul
2007:48).
Well into the twentieth century Dena'ina practiced a ritual that involved sending the spirit
of the animal to the "reincarnation place." Land animal bones were burned in the fire and water
animal bones, like salmon, were returned to the water. These practices ritualized ecology and
were said to bring the animal back to be hunted or fished again (Boraas and Peter 1996:188-190).
Archaeological evidence indicates the Dena'ina were burning bones in their fire hearths (Boraas
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and Peter 2008:220-222)
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D. Traditional Yup'ik and Dena'ina Spirituality and Cosmology
Many modern practices of Yup'ik and Dena'ina have their basis in traditional spiritual
and cosmological beliefs, though they are sometimes re-contextualized in Christianity. This
section discusses the traditional spiritual and cosmologic beliefs and practices of both peoples
1. The Yup'ik People
Traditional Yup'ik values revolved around not only their extended families, but also their
relationships with the wild animals and other components of the natural landscape. Within this
belief system, the Ellam yua, or creative force, was a universal cosmic presence who coordinated
existence and established a basic ordering framework; tunghitwere powerful spiritual beings
who controlled the recycling of different animals, fish, and bird forms (Langdon, 2002).
The Yup'ik have traditionally regarded animals as other peoples, or categories of
kinsmen, with whom they have fluid relations that often cross species and interpersonal
boundaries. There are numerous stories of half-animal, half-human beings who live in the
villages or of people turning into seals, birds, fish, or other animals, and then turning back into
humans, as well as stories of people who seem to be human, but turn out to be seals or other
animals in a temporary human form. Several major traditional festivals and ceremonies,
described below, honored this relationship. The spiritual values associated with each of these
festivals emphasized sharing between humans and respect and care for animals. Traditional
stories and advice speak of the animals giving themselves to the humans when the humans need
them for food. The good practices of sharing, care, and respect (e.g., being careful with the
animal's body and soul, and not wasting the food) ensured the animals' continued willingness to
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give themselves to the hunters and fishermen in the future. Sharing of the products of subsistence
with their human kin and other relations also strengthened the bonds of family and community.
A version of The First Salmon celebration in the river communities is still celebrated today,
when those who have caught the first king salmon in the spring share them with Elders and all
those in need, as well as with friends and family, emphasizes these values.
The Yup'ik relations with the wild animals and fish of their landscape were primary, and
in many ways still are. The Yup'ik related to the fish, the bear, the caribou, the moose, the ravens
as relations, others equally inhabiting the landscape with them as interrelated peoples. During
spring, summer, and fall the Yup'ik hunt and fish the animals as food, but when processing the
animals as food they treat them with respect and care, and enable their return through rituals and
ceremonies. In winter, a period of rest and renewal for the human population, in the past the
Yup'ik attended to the renewal of life through the rebirth of the animals they had hunted, and
fished, in, according to Fienup-Riordan five ceremonies, "three of which focused on the creative
reformation of the relationship between the human community and the spirit world on which
they relied." (Fienup-Riordan 1994:267). Today, many of the Russian Orthodox ceremonies
continue to be based on this ancient calendar of propitiation of the world of the spirit, in all
seasons. Ellamyua was a universal cosmic presence who coordinated existence and established a
basic ordering framework; tunghit were powerful spiritual beings who controlled the recycling of
different animals, fish, and bird forms (Langdon 2002). During the winter ceremonial season, the
men beat the circular drumtraditionally made from stretching seal gut on a wooden framefor
songs and dances. The drum beats represented the heartbeat of Ellamyua. Thus, the celebrations
were spiritual in the deepest sense. They were also material, involving the exchange and sharing
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of wild subsistence foods from both animals who had given themselves willingly to the hunters
and plants gathered from the landscape, considered to be spiritually alive.
During the Bladder Festival, at or around the Winter Solstice, the women brought out the
bladders of seals, which they had been saving since their husbands brought the seals to them to
prepare, because the Yup'ik believed that the souls or essence of animals are located or retreat to
their bladders when they are killed. By saving the seal bladders and returning them to the sea, the
Yup'ik enable the seals to be reborn, and present themselves again as food for the Yup'ik when
needed. The women take the seal bladders to the qasgiq, or men's house, where the men inflate
them and keep them for about ten days, while they go through a series of rituals to honor the
seals and share food in the community, before returning the bladders under the ice, to the sea,
enabling the seals to be reborn and to present themselves to the Yup'ik when needed again as
food. The men would compose new songs for the Bladder Festival, including songs about
salmon, and sing continuously in the qasgiq; people believed that light from the lamp and the
songs drew the attention of animal spirits (Fienup-Riordan, 1994:284).
At Qaariitaaq, at the beginning of the Bladder Festival, the young boys were painted to
represent the spirits of the dead, and went visiting, going around to the different houses to collect
special food treats. Every house was brightly lit, and the hostesses wore their best clothes. The
boys held out their hand-carved bowls, and the women handed out the special snacks. On the
fifth night of these celebrations, the boys, and men, came to fully embody the spirits of the dead,
and the fifth night was considered the arrival of the spirits. (Fienup-Riordan 1994:271). At
Aaniq, held directly after Qaariitaaq, two men dressed in gut skin parkas, are referred to as
mothers, the "aanak, " and they are taken around to collect newly made bowls filled with
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akuutaq, traditionally a mixture of fat and berries. Small girls and boys referred to as their
"dogs" would accompany them.
The way that people do things
And the way of helping others
And the way of creating friendship
The Bladder Festival is like an opening for these things to occur
And through those events
The people being scattered
Through that too they are gathered
(Toksook Bay Eders, November 3, 1983 NI57 in Fienup-Riordan, 1994: 267).
Today, starting during the Russian Christmas season the modern ritual of "Starring"
follows this familiar pattern - groups go visiting from house to house, and receive special foods.
Other important ceremonies include the Great Feast for the Dead, Elriq, held every ten
years, as well as the annual feast for the dead, and Kelek, a festival that included both serious and
comic masked dances, when "animal spirits and shamanic spirit helpers made themselves visible
in the human world in dramatic form" (Fienup Riordan, 1994:316). Kelek was performed to
influence the animal spirits and elicit successful hunting and fishing through the return of the
animals the following year.
Two other winter festivals underscored the redistribution of goods, including subsistence
foods. The first, Kevgiq, the Messenger Feast, was a celebration and display of the bounty of the
harvest, in which villages challenged each other to exchanges of wealth, with demands for
specific items that were difficult to provide, such as certain game meat in a year when that game
animal was scarce. Kevgiq served to reduce tensions between villages through sharing and
friendly competition. It also provided food security by strengthening ties between villages and
encouraging exchange relationships that could help people in times of food shortages. Sharing
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was considered to be a behavior that would be rewarded by the return of the animals to those
hunters and fishers the following year. Petugtaq., the Asking Festival, was a challenge to
exchange gifts of value between cross-cousins and others, where the person whose gifts were the
most valuable gained the highest prestige. Cross-cousins were in "joking cousin" relations with
each other, and were able to call each other out on bad behavior, embarrassing each other
without repercussions, since they were not permitted to get angry with each other (Fienup-
Riordan, 1994:330). The behaviors were thus made public and frequently resolved through this
tension-reducing mechanism. Both festivals involved teasing, dancing and singing as part of the
ritual celebration of the exchanges. All of the traditional festivals required subsistence foods, not
only for sustenance, but also for the meaningful symbolic and material exchanges.
During their ceremonies, the Yup'ik wore masks they had carved, often representing
animals or those in transition between the animal world and the human world, the half-animal,
half-human. These masks symbolized both the high regard of the Yup'ik for the animals and the
importance of their roles Yup'ik culture. For the Yup'ik, the masks were agayuliyararput, or
"our way of making prayer" (Fienup-Riordan, 1996:xviii).
Dances, including ingulagthe women's loon courtship danceand other bird dances,
filled the evenings and contributed to the festivities. Each dance told a story and many featured
the animals with whom the Yup'ik partnered in their negotiation for existence in the challenging
landscape. Dances were traditionally an essential part of the culture and celebrations and have
returned in force as part of cultural revitalization along the Nushagak and elsewhere. Fienup-
Riordan (1994:288) quotes Billy Lincoln:
And at night, every night, they have what is called nayangaq. They dance. These
young people who are sitting against the far wall go down in front of them and
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dance, sitting down pretending to be some animal, so thus, the nayangaq. They
imitate a certain animal. When the time came whatever animal he is pretending to
be he imitates its noise. They imitate all kinds of animals - loon, hawk, raven,
arctic fox. They make noise accordingly. They dance pretending to be some
animal (July 10, 1985).
The dancers represented the many ways the stories and lives of the animals were woven into
their own, in the richness of shared existence in the watersheds of southwest Alaska. Lincoln
continues:
These dance motions were more than the mere imitation of the motions of the
animals. When the performers danced during Kelek, they actually performed the
animals' dances. Just as married women danced the loon's mating dance during
Ingulaq, so the performers during Kelek danced the dances of the animals whose
presence they hoped to elicit in the year to come. . .
In 1913 Hawkes quoted a Unalakleet chief in an eloquent estimation of the value of these dances
within Yup'ik culture: "To stop the Eskimo singing and dancing," he said, "was like cutting the
tongue out of a bird" (Hawkes cited in Fienup-Riordan, 1994:320-321).
Fienup-Riordan (1994:355; see also Fienup-Riordan 2010) summarizes how the Yup'ik
traditionally saw themselves in relation to the universe: "Yup'ik cosmology is a perpetual
cycling between birth and rebirth, humans and animals, and the living and the dead. Their
relationship between humans and animals reflects a cycle of reciprocity in which animals give
their bodies in exchange for careful treatment and respect."
2. The Dena'ina People
The traditional Dena'ina spiritual world revolved around a quest for k'ech eltcmi, or "true
belief," as a way to understand and interact with the natural world (Boraas and Peter, 1996:183-
4). The Dena'ina believed that social and ecological harmony was affected by an individual's
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attitudes, actions, and even thoughts toward other Dena'ina and to nature. To maintain harmony,
the Dena'ina sought true belief, a kind of mind-set expressed through hunting practices, cooking
rituals, communication with animals and plants (prayer), and other practices that demonstrated
having a "good attitude" toward the forces of nature. Kalifornsky (1991:13) writes that,
"Whatever is on earth is a person [has a spirit] they used to say. And they said they prayed to
everything. That is the way they lived." Achieving k 'ech eltani involved a spiritually torturous
and mentally rigorous quest for understanding (Boraas and Peter, 1996:187).
Many of the Dena'ina traditional stories (sukdu) describe the dire consequences of having
a bad attitude by not practicing the prescribed rituals such as burning the bones of consumed
animals or distributing fish bones in the water as means to symbolically assure the animals
would come back (Boraas and Peter, 2008:222-223). In these stories, a bad attitude would have
the consequence of the animals, believed to be both sensate and willful, withdrawing and not
offering themselves to be taken for food. The result would be starvation. A bad attitude could
result in social turmoil or mental illness. There was immense pressure to behave and think
respectfully toward the natural world including salmon.
In a forthcoming chapter on Dena'ina world view, Boraas (in press) writes the following
about traditional attitudes toward animals:
Attitudes toward bears typify attitudes toward animals. In "Three People in
Search of Truth," (Kalifornsky 1991:164-167) three brothers hunt a brown bear,
the most feared and respected animal. The first fails because he is poorly skilled;
the second fails because he is impetuous, and the third succeeds because he is
skilled, controlled and speaks the correct words to the bear, which then respects
him and does not resist being killed. In Kenai a successful hunter used the phrase
Chadaka, k'usht'a nhu'izdeyeshdle, which translates as "Great Old Man, I am not
equal to you," to communicate humility toward the bear he was hunting
(Kalifornsky 1991:167). In 1966 Mrs. Mike Delkettie, a Nondalton Dena'ina,
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reported that a similar saying was used in that area; moreover, the eyes of the bear
were buried near the spot where it was killed as an offering showing proper
respect (Rooth 1971:62). Francis Wilson, also from Nondalton, told Rooth
(1971:50) that, after a bear was killed, they had to follow prescribed procedures,
particularly in the treatment of the head, lest they never kill another bear, because
"the bear still knows what is happening, so they have to be very careful with what
they are doing." Hunting rituals and prayers were meant to thank an animal for
allowing itself to be killed and sometimes it also involved giving an offering as a
measure of the importance of proper attitude (Rooth 1971:50).
The First Salmon Ceremony (Osgood, 1976:148; Kari and Fall 2003:184-190) expresses
the intimate relationship of Dena'ina and salmon. The First Salmon Ceremony was based on a
traditional story. As the Osgood's retelling goes, a qeshqa 's (chiefs) daughter was admonished
not to go near the fish weir. The determined girl went anyway to find out what was in the trap,
promising to return later. At the fish trap she saw a king salmon, began talking to him, and
gradually transformed into a salmon and disappeared with him. The desperate qeshqa looked for
his daughter to no avail. Years later, the qeshqa was collecting fish from the weir. He put them
on the grass and took them to be cleaned, but forgot one little one. He returned to find a little boy
sitting there. He walked around the boy three times and realized it was his grandson. The boy
then told his grandfather the things that should be done to ensure the salmon return each year,
and those things became the First Salmon Ceremony, a world renewal ceremony3 which ritually
recognized the salmon's return and the Dena'ina as salmon people whose spirit is merged with
the fish.
In 1862 Hegumen Nikolai, the first missionary priest stationed in Dena'ina territory
3 World renewal ceremonies are important identity-building ceremonies that recognize the
beginning or end of a year's subsistence activity and social cohesion. In American culture
Thanksgiving is a world renewal ceremony.
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wrote in his travel journal, "In the middle of May the king salmon reached our area [writing
from Kenai]. This is the best red fish we have here, and the Kenaitze celebrated the fish run with
some sort of festivities, during which they treated each other with food" (Znamenski 2003: 91).
Fr. Nikolai was clearly referring to the First Salmon Ceremony.
Water was particularly important in Dena'ina spirituality in the act of moving into a
spiritually liminal state. One kneeled beside a river or lake and took three sips of water (Boraas
in press). This was practiced well into historic times and also occurs in mythological stories
(sukdu). For example in "The Woman Who Was Fasting" (Kalifornsky 1991:168-9) a young
woman was ritually fasting and spoke these words "People will learn something from our
beliefs" as she took three sips of water. She was then able to perform a spiritually power act
upon which she said, "When we pray and we fast there is another dimension."
Some places took on special importance. The Giants Rock, Dzelggezh, was along an old
Dena'ina trail that became the Pile Bay Road between Old Iliamna and Kamishak Bay on Cook
Inlet, one of the major trails connecting eastern and western Dena'ina territory. The rock was the
site of a mythological story and was a spiritual place (Johnson, 2004:49-54). The rock was
dynamited in 1955 as part of road building activities by the Territory of Alaska; Dena'ina still
regularly leave votive gifts at the site in homage to the place and the mythological event that
happened there. Other sacred rocks and sacred locations exist in Dena'ina territory, but for most
their locations are privileged cultural information (Boraas 2009:10-20).
Not only are there sacred sites but the Dena'ina believed the landscape retained a sense of
events that happened there: events which could be good or bad. Spiritually powerful people and
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animals could detect information about these events and, thus, to travel was to encounter morally
good and morally bad events encoded into the landscape (Boraas 2009:8-10).
Figure 4. Nushagak River, January 18, 2012
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E. The Yup'ik and Dena'ina Languages: Salmon and Streams
1. Voices of the People
Talk Native, no English.... They talk Native [Yup 'ikj better [than English], [in reference to Elder
interviews in Yup'ik] M-25, 5/18/11
That's why we quit using our Native tongue because we get our... ears pulled. I don't know how
many times I sit in the corner because I use my Native tongue. We couldn 't speak our own
language in school because we get abused. F-46, 8/20/11
When we first went to school they took our dialect away from us and told us to speak English
only. If we spoke our Native tongue we would get hit by the teacher which isn 't right. Now they
call it abuse. Anyways none of us speak our Native tongue [Dena 'ina] because of that. My mom
didn 't speak English.... F-48, 8/20/11
2. Introduction
Language is intimately tied to cultural identity and Yup'ik and Dena'ina have evolved as
languages of place for their respective areas over thousands of years. Landscape, subsistence,
social relations, and spirituality are reflected in both languages. The variety of words a language
has for a given topic generally reflects the importance of that topic to the people who speak it.
Given their cultural importance, it is not surprising that both Dena'ina and Yup'ik have
numerous, highly detailed terms involving salmon, other fish, and fishing. Streams are also
intimately tied to Dena'ina and Yup'ik psyche and their languages reflect that fact.
3. The Central Yup'ik Language
The Yup'ik people of the Nushagak and Kvichak River watersheds are part of the Central
Yup'ik group, of whom there is a population of about 25,000 in an area that also includes coastal
communities and the lower and middle Kuskokwim River drainage (Krauss, 2007:408) (See
Table 6). Ten thousand four hundred of this population, or 42%, speak Central Yup'ik of which
the 7,000 mostly Yup'ik of the Nushagak and Kvichak River drainages are a part. Central Yup'ik
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has one of the highest percentages of speakers among indigenous languages in the U.S and is an
indicator of strong cultural heritage. Yup'ik is the first language for many residents in the study
area and the language in which many feel most comfortable expressing complex or heartfelt
ideas, which is why, for this project, we encouraged interviewees to respond in Yup'ik if they so
choose. Eight of fifty-five interviewees spoke in Yup'ik. One Yup'ik interviewee (M-25; 5-18-
11) spoke about helping set up a 2011 Elders Conference which occurred a few days before our
interviews in New Stuyahok in which the entire discussion was in Yup'ik. He said, "I set up that
meeting [Elders Conference], I try to do it for a long time.. .yes, talk Native [Yup'ik], no
English. Get somebody else to translate.. .they talk Native better [than English]."
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Table 6 Estimated Number of Central Yup'ik and Dena'ina Speakers. Data from Krauss
(2007:408)
Language Family
Eskimo- Aleut
Athabascan-Eyak-
Tlingit
Language
Central Yup'ik
Dena'ina
Population
Estimate
25,000
1,000
Speakers
10,400
50
Percent
Speakers
42%
5%
Table 7 presents Yup'ik terms for salmon, related fish, and fishing activities. In many
cases there are multiple words and/or dialect differences. As indicated the sheer number of words
are indicative of a long history with salmon and fishing activities. Moreover, the nuanced
meaning of some words is indicative of a deep knowledge of salmon and related activities. For
example the word kiarneq' means "unsalted strip or fillet offish flesh without skin, cut from
along the backbone and hung to dry"
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Table 7. Yup'ik Words for Salmon and Other Fish Species and Related Fishing Terms, (x
means literal translation same as English term.) From Jacobson (1984)
English Term
Yup'ik Word
Literal Translation
salmon (generic)
(Oncorhynchus spp.)
neqaraq
any species of salmon
dog salmon, chum salmon
aluyak
iqalluk
kangitneq
mac 'utaq
teggmaarrluk
x
'fish'
'old dog salmon after
spawning'
x
boiled half-dried salmon
humpback salmon, pink salmon
amaqaayak
amaqsus
cuqpeq
terteq
amaqatak
sayalleraam amaqatii
neqnirquq
x
x
X
X
'back offish, hump on
back'
'back of spawning red
salmon is tasty'
silver salmon, coho salmon
caayuryaq
qakiiyaq
qavlunaq
uqurliq
'streak or wake made on
surface by fish'
red salmon, sockeye salmon
cayak
sayak
sayalleq
sayagcurtuq
imarnikaralegmun
'he is fishing for red salmon
at a deep calm place'
spawning salmon
masseq
masruuq una neqa
nalayaq
nalayarrsuun
'old salmon near spawning'
'this fish is a spawning
salmon'
x
'fish spear to catch
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king running under smelt
salmon egg
salmon strip
salted fish or meat
scale (fish)
rolled oats
smelt
stick(w) fish-spreading
stickleback
talayaq
talmag (NUN)
talmagtut
aciirturtet
cilluvak
culunallraq
taryitaq
culunaq
culunanek ajurciuq
sulunaq
sulunanek ingqillruuq
taryitaq, taryiraq
taryirki sulunarkat
kapciq
qelta
akakiik qeltairru suu
pirniaraqa
qeltengalnguut
cemerliq
cimigliq
ayagta
ayagtekartellruunga
cukilek
angun cukilegnek
spawning salmon'
'calico salmon'
'to spawn (of fish)'
'they are spawning'
'the first group of king
salmon running under the
smelt'
'salmon egg, especially
aged salmon egg'
'salted and dried salmon
strip'
'salted fish or meat that is
eaten after it is cut up and
soaked to remove excess
salt'
'she is soaking some salted
fish'
see culunaq
'my wife cut up the salted
fish'
'salted salmon strip'
'put salt on the pieces of
fish to be preserved'
X
'fish scale',
'take the scales off the
whitefish so that I can make
soup with it!'
'things like fish scales'
X
X
'prop, support, especially a
small stick used to keep a
cut fish open as it dries'
'I gathered material to use
as spreaders for drying fish'
'one with quills'
'the man is dipnetting for
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supper
tail, fish
preopercle
fish cheek
trap, fish
whitefish with pointed head
young whitefish
frozen raw whitefish
To fish (v)
Fish
Boiled fish
qaluuq
ilaqcungaq
quarruuk
atakutaq
papsalqitaq
papsalquq
ulluvalqin
ulluvalquq
taluyaq
cingikeggliq
esevsiar(aq)
iituliar(aq)
qassayaaq
akakiigem meluanek
qassallruunga
neqsur
iqalluk
ilaqcuugaq
neqa
neqet amllertut maani
qimugtet neqait
nangyarpiartut
neqtulnguunga
neqa unguvangraan
uklia
neqngurtuq
nereneqaiq, neqiaq
egaaq
sticklebacks'
X
'needlefish'
'supper, evening meal'
'dried fish tail'
'tail or caudal fin offish'
'gill cover of a fish,
preopercle'
'cut from the fish'
'fish tray'
X
X
'whitefish fry'
'frozen whitefish aged
before freezing and served
frozen'
'I ate the whitefish eggs
raw'
?
'dog, chum salmon, fish'
'small fish found in lakes'
'food;fish'
'the fish are plentiful here'
'the dogs' food is almost
gone'
'i'm tired of eating fish'
'even though the fish is still
alive he is cutting it up'
'there was food
everywhere', lit. 'it became
food'
'food-stealing bird'
'any cooked fish or other
food'
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Bundled fish
Canned fish
Cut fish
Fish cut in half
Dried fish
Dried small fish
Dried fish heads
inartaq
paankaraq
qakiiyak paankarak
uksuqu nernalukek
cegesseg-
cegtuq
cegaa, ceggaa
ceg'aq, cegg'aq
seg-
ulligte-
ulligtuq
ulligtaaa
ulligciuq
ulligtaq
ingqii-
inguqin, inguqitaq
neq 'liur-
neq 'liurtuq
qup 'ayagaq(NUN)
neqaluk (NUN)
neqerrluk
palircima
nevkuq
ulligtaruaq
nasqurrluk
X
X
'he is canning two silver
salmon so that he can eat
them in winter'
'to cut fish for drying'
'she is cutting fish'
'she is cutting it'
'a fish cut for drying'
(see ceg-)
'to cut fish for drying, in the
traditional manner, making
cuts so that air can reach all
parts of the flesh; (NUN) to
turn over'
'it is cut for drying'
'she cut it for drying'
'she is cutting it for drying'
'fish cut for drying'
'to make the horizontal cuts
in fish flesh while preparing
it for drying'
'board on which one
prepares meat or fish'
'to work on fish (cleaning
it, etc.)'
'he is working on fish'
'fish cut in half to hang and
dry'
X
X
'to be burnt by the sun (of
dried fish)
X
'split and dried small fish,
such as whitefish, pike or
trout'
'cut and dried fish-head'
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Dried frozen fish
Air dried fish
Fish dried in a basket
Fish partially dried and boiled
Frozen fish
Poke fish
Fish partly smoked and stored
in seal oil
Fish in strips
Dried Fish tails
Fish strung to dry
Fish hung to dry
Raw fish
Raw frozen fish
Cooked piece offish
qamiqurrluk
irniani nerevkaraa
tepnek
yay 'ussaq
tamuaneq
tut 'at (plural)
egamaarrluk
teggmaarrluk
cetegtaq
kumlaneq
nutaqaq
qercuqaq
uqumaarrluk
arumaarluk
kiarneq
palak 'aaq (BB)
parmesqatak
papsalqitaq
piirrarrluk (Y, HBC)
kanartaq
qassaq, qassaulria
qassar-
qassartuq
qassaraa
quaq
ukliaq
(see above)
'she let her child eat some
aged fish heads'
'dried tomcod or whitefish
that has been frozen all
winter'
X
'fish packed down and dried
in a basket'
X
'boiled, half-dried salmon;
dog salmon, chum salmon'
'fish slightly aged and
stored in seal oil'
X
'unsalted strip or fillet of
fish flesh without skin, cut
from along the backbone
and hung to dry'
'strip of dried flesh'
X
9
'small fish, such as tomcod
strung up for drying'
X
'raw fish or meat'
'to eat raw fish or meat'
'he is eating raw fish'
'he is eating it raw'
'fish to be eaten raw and
frozen'
X
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Fish bin
Fish trap
Fish rack
Fish wheel
Fish fence
Fish spear
Fishing line
Fish camp
Fish Village
Fisherman
Fish hook
qikutaq
taluyaq
initaq
her 'aq
qer 'aq
akalria
capon
angutet capcirtut
uqvianek
manignarrnaluteng
taluyakun
kalgun
aggsuun
ag 'ssuun
ipiutaq (NSU)
kiagvik
neqlilleq
neqlercurvik
neqsurta
neqsurtuq
neqsurvik
neqsurtuq
tuniarkaminek
aataka neqsurtenguuq
iqsak
iqsag/manaqutaq
'bin used for temporary
storage offish before they
are cut up for drying'
X
'part of a fish rack on which
the fish is directly hung'
X
'weir, fish fence; wall'
'the men set a weir of
willows to catch loche with
a fishtrap'
'weir, fish fence extending
from the bottom of the river
and leading fish to a place
where one can catch them
with a dipnet'
X
X
X
'summer fish camp'
(see above)
'fish village, site on the
lower Yukon'
X
'he is fishing'
'fishing place'
'he is fishing commercially'
'my father is a fisherman'
X
'to fish with a hook and
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iqsagtuq/manartuq
iqsagaa/manaraa
manaq
manor
manaryartuq
qerrlurcaq
line, to jig for fish'
'he is hooking for fish'
'he hooked it'
'fishing lure with hook'
'to fish with a hook, lure,
and line, usually (though
not necessarily) through a
hole in the ice in winter'
'he went to fish with a hook
and line'
'fishhook which is baited
and set below the ice, held
in place with a stick across
the hole, and left unattended
to be checked periodically'
Fish net
kuvyaq, kuvya, kuvsaq
kuvya
kuvyauq
kuvyaq cangliqellruuq
nutaranek
qemiraa kuvyaq
qilagcuutmek aturluni
kuvyaq civtaa
kuvyaq takuua
kuvyarkaq
qelcaq (Y)
'to fish by drift-netting or
purse-seining'
'he is drift-netting'
'the net caught lots of fresh
fish'
' he is stringing the net
using a net shuttle'
'he set the net'
'he checked the nets'
'twine for making nets'
'net into which fish are
driven by peopoole who
walk in and thrash the
water'
Set net
petugaq
Fine mesh net
caqutaugaq(NUN)
'fine mesh net for dog
salmon, worked by hand by
men standing in the water,
not left unattended'
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Net shuttle
Net setting line
Net sinker
Fishing rod
Roe
aged roe
herring roe
fish rack
trout
imgutaq
qilagcuun
amun
atlirneq
nuvun
qemiq
qemirtuq
qemiraa
kic 'aqutaq
manaq
piqrutaq
tin 'aq
cilluvak
imlauk
meluk
melug
cuak
imlauk (NUN)
qaarsaq
qiaryaq (NUN)
ker'aq(NSU)
qer 'aq
anerrluaq (BB)
anyuk (BB)
X
X
'line used to set and reset a
net under the ice'
'lead line offish net'
'threading device (such as
the line used to set a net
under the ice, or a needle
threader)'
'lead line or float line of a
net'
'he is stringing (a net)'
'he is stringing it'
X
'fishing lure with hook'
'salmon egg, especially
aged salmon egg'
'fish egg, roe'
'fish eggs, roe; fish eggs
prepared by allowing them
to age and become a sticky
mass'
'to suck; to eat roe directly
from the fish'
X
'dried herring egg'
X
'herring eggs, so called
because they crackle when
eaten'
X
X
'type offish, salt-water
trout'
X
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lake trout
steelhead trout
rainbow trout
dolly varden (char)
herring
Arctic cod
Pike
Wolf Fish
Smokehouse
Smoked Fish
Subsistence
cikignaq
irunaq
talaariq
iqallugpik
iqalluarpak, iqallugpak
iqalluaq
uksumi-llu iqsagnaurtut
cuukvagnek
qugautnaq (NI, NUN)
elagyaq
puyurcivik
talicivik
neqnek aruvarqiyartua
talicivigmi
aruvarqi-
aruvir-
puyurqe
puyurte-
angussaag-
yuungnaqe-
X
X
X
X
X
'boreal smelt'
'and in the winter they
would hook for pike'
X
'partially underground
cache; pit for cleaning fish;
smokehouse'
X
'shelter for smoking fish,
smokehouse'
'go smoke the fish in the
smokehouse'
'to smoke fish'
'to be smoky; to smoke
(fish)'
'to be smoked; to feed the
fire when smoking fish'
'to smoke (fish)'
'to hunt, to try to catch
game'
9
4. The Dena'ina Language
There is a dramatic difference in language retention between the Yup'ik of the Nushagak
and Kvichak River watersheds and the Dena'ina of the Iliamna Lake and Lake Clark area. In
contrast to the Yup'ik, the Dena'ina population is much smaller, estimated by Krauss (2007:408)
at 1,000 for the Iliamna/Lake Clark area and Cook Inlet Basin. Krauss estimates that within this
population there are only 50 Dena'ina speakers remaining (see Table 6), most of whom live in
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the vicinity of Nondalton or Lime Village (the latter outside the study area in the Kuskokwim
River drainage). The youngest active Dena'ina speaker is 64 years old. Dena'ina is, thus, one of
the world's most endangered indigenous languages (Boraas 2010:2). The reason for the disparity
between Dena'ina and Yup'ik language usage is complex but the main reason for Dena'ina
language extinction was the Alaska Territorial School's federally mandated policy of punishment
for children speaking their indigenous language in school. This forced assimilation policy
occurred to various degrees throughout Alaska but its application seems to have been particularly
harsh in Dena'ina territory (Boraas 2010:2).
Given the importance of language to cultural identity, the Dena'ina have begun to
revitalize their language and significant efforts are underway to avoid its extinction both in
spoken and written form (cf Boraas and Christian 2010). There is a history of Dena'ina Elders
working with linguists dating back to Anna Brigitta Rooth's (1971) work in 1966 in Nondalton
followed by dozens of bilingual publications by James Kari working in collaboration with
Dena'ina speakers starting in the 1970s and the bilingual publication of Joan Tenenbaum (1984).
More recently a number of speakers from Nondalton and Lime Village have participated in
Dena'ina Language Institutes, sponsored by a consortium of institutions including the Alaska
Native Language Center, Alaska Native Heritage Center, the Sovereign Nation of the Kenaitze,
and Kenai Peninsula College. The one to three-week institutes have been held at various
locations including Nondalton and include workshops on Dena'ina language learning and
teaching. Recently, two speakers from the study area, Andrew Balluta of Nondalton/Newhalen
and Walter Johnson of Pedro Bay, now of Homer, have collaborated with linguist James Kari on
important bilingual publications: Shtutda 'ina Da 'a Shet Qudel: My Forefathers are Still Walking
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with Me (Balluta 2008) and Sukdu Net Nuhtghelnek: I'll Tell You a Story: Stories I Recall from
Growing Up on Iliamna Lake (Johnson 2004). Finally, numerous speakers living and deceased
(through archived recordings) contributed to Dena 'ina Elnena [Dena 'ina Territory]: A
Celebration edited by Karen Evanoff (2010).
The language is indicative of the importance of water and salmon and other fish to the
Dena'ina. Streams are intimately tied to the Dena'ina psyche through language. The Dena'ina
words for directions are not based on the cardinal directions, but on the concept of upstream or
downstream. A Dena'ina description of direction results from combining one of five stems,
indicating upstream, downstream, and related terms; one of six prefixes, indicating proximity;
and a suffix indicating general direction or location (Kari, 2007:336). For example, the word
"yunif combines the stem "«/'" (upstream) with the prefix "_yw" (distant) and the suffix "f (at a
specific place) and means "at a specific place a long way up upstream." If one were using that
phrase at Iliamna, yunit would mean the direction toward Nondalton, which is a specific place far
upstream; in this case, the direction would be north, because from Iliamna the Newhalen River
flows south.
Because of the importance of stream stems reflecting a fundamental cultural construct
affecting a wide range of cultural activities (subsistence, diet, travel, directions, spirituality etc.)
Kari (1996) has proposed migration theory for Dena'ina and other Athabascans (who employ a
similar directional system) based on variants in the stream stem morpheme. Kari suggests a
movement of people from northern British Columbia, to the Yukon River area to the Kuskokwim
piedmont, to Dena'ina territory. Boraas (2007:35) believes this to be the best hypothesis of
Dena'ina origins to date.
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The spirituality of water is also embedded in the language. The Dena'ina have 36 terms
for streams (Kari 2007:123-4), among those the primary word for 'water' is of special note. The
Dena'ina word for "water" vinilni (in the Inland dialect, mibii in the Outer dialect) is unique
among other Athabascan/Dene languages and Dena'ina linguist James Kari considers it to be
esoterogenic meaning a special word reflecting special importance or sacredness (personal
communication, Dr. James Kari, UAF Professor Emeritus, December 6, 2011). Dena'ina Elders
Clare Swan and Alexandra Lindgren (2011) state "the Dena'ina word for water was held sacred"
and by implication the water was sacred. The word vinilni and its sacred connotations is reflected
today in the Orthodox Great Blessing of the Water ceremony described in section III.F.3 in
which river water is annually baptized and made holy.
The Dena'ina named a general category of animal or plant by the name of its most
important representative. For example, the name for animal is ggagga, for brown bear, and the
name for tree is ch 'wala, for white spruce. Not surprisingly, the name for fish is the name for
salmon, liq'a. Table 8 is a compilation of Dena'ina terms for salmon, freshwater fish, and fishing
technology which, like the Yup'ik counterparts, shows an intimate connection with salmon, fish,
and fishing.
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Table 8. Dena'ina Terms Involving Salmon, Freshwater Fish and Fishing Technology.
(x means literal translation same as English term.) Data from Kari (2007)
Dialect notations: I = Inland, U=Upper Inlet, O=Outer Inlet, L=Lime Village,
Il=Iliamna, S=Seldovia, Lk-i=Kuskokwim Deg H'tan, Su=Susitna Station, E=Eklutna,
Ty Tyonek, T=Talkeetna, Kn=Knik
English Term
salmon (generic) (Oncorhynchus
spp.)
Male fish
Female fish
Small fish
Fry, baby fish
Bottom fish
Spring fish run
Spring fish caught under ice
king salmon, Chinook salmon (O.
tschawytscha)
king; salmon sizes: smallest
two-foot king salmon
largest king salmon
middle-sized king salmon
humpback salmon, pink
salmon (O, gorbuscha)
red salmon, sockeye salmon
(O. nerka)
nickname
old fall sockeye
dog salmon, chum salmon
(O. keta), (I) early summer
Dena'ina Word
tiq 'a (IU)
luq 'a (OSl)
Hest 'a, qest 'a (IO)
Tl 'ech 'I (U)
Q 'in 'i
Q 'inch 'eya (IO)
Q 'inch 'ey (U)
Chagela gga (U)
Shagela gguya (I)
Shagela ggwa (O)
Lch 'eli, dghelch 'eli
Tahliq 'a (IU)
Tahluq 'a (O)
Litl'eni(UI)
Ten t'uhdi (U)
tiq 'aha 'a (IU)
luq 'oka 'a (O)
chavicha, tsavija (O)
tiq 'agga (U)
ggas ten 'a (L)
q 'inagheltin (U)
tiq 'aka (U)
vigit'in (L)
tl 'istqeyi (U)
qughuna (OUSl)
liq'a(I)
t'q'uya (LNOSl)
k 'q 'uya ON)
q 'uya (U)
veghutna qilin (I)
bendashtggeya (U)
dghelbek'i (UO)
alima (Oil)
seyi (U)
Literal Meaning
X
X
'roe one'
'shiny one'
'underwater fish'
X
X
"big salmon'
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chum salmon
August run dog salmon
silver salmon, coho salmon
(O. kisutch)
steelhead trout (Salmo
gairdneri)
running salmon
fish laying eggs
spawned-out salmon
dead salmon
fall salmon, esp. sockeye
fingerling, baby salmon, alevin
first fish run
last fish run
old female salmon
red-colored salmon
spring (early) salmon run
summer salmon run, sockeye
season
fall-winter running salmon
dead salmon that drift ashore
salmon captured in weir
Non-salmon fish
nulay (NL)
shighat 'iy (Lk-i)
nusdlaghi (I)
nudlaghi (O)
nudlegha, nudleghi (U)
usdlaghi (O)
telaghi (II)
turn, turn denlkughi (N)
shagela (U)
tuzdlaghi (OI)
tuydlaghi (U)
tag 'innelyaxi (I)
tag 'innelyashi (UO)
nudujuzhi, dujuzhi (I)
dujuyi (U)
itak'i (O)
tilani
hey luq 'a (O)
hey liq 'a (IU)
tuyiga (OI)
liq 'agga (U)
liq 'a gguya
qtsa ghelehi
q 'ech 'en ghelehi (I)
unhtl 'uh ghelehi (UO)
unhtl 'uyeh (I)
q 'in ch 'ezhi (I)
q 'in ch 'eya (U)
nuditq 'azhi (I)
nishtudghiltani (U)
ts 'iluq 'a (O)
litl'eni(UI)
chiluq 'a (O)
hchiliq 'a (UI)
shanlaghi (UI)
tuleha (OU)
tulehi (I)
niqatayilaxi (I)
q 'anughedeli
Shagela (IO)
Chagela (UIl)
'runs again'
icn;i
'one that swims back'
? 'one that swims past'
'one that runs'
'water one'
'fish'
'one swimming in water'
X
X
X
X
X
'winter salmon'
'water spirit'
'little salmon'
X
X
'infested roe'
'one that is red'
'that which floats in midstream'
'straight salmon'
'spring one'
X
'summer run'
'one running in water'
X
'those swimming back'
'fish'
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Alaska blackfish
Freshwater sculpin
Burbot, lingcod
Burbot's chin barbell
Arctic char
Eel, lamprey
Large lamprey
grayling
Grayling's dorsal fin
Freshwater herring, least cisco
Three-spined stickleback
Spawning stickleback
Northern pike
Small pike
sheefish
sucker
Brook trout, Landlocked Dolly
Varden char
Lake trout
Rainbow trout
Chebay (U)
Huzheghi, huzhehi (L,N)
Ch 'qenlt 'emich 'a
Ch 'qenlt 'emch 'a (NL)
Ch 'qeldemich 'a (II)
Ts 'est 'ugh 'I, ts 'est 'uhdi
(U)
Ch 'unya (I)
Ch 'any a (U)
K'ezex (Lk-i)
Veyada k 'ich 'aynanik 'et 'i
Vat (NL)
Suy liq 'a
Liq 'a q 'ints 'a
Lizil (O)
77 'eghesh (I)
Ts 'ilten hutsesa (U)
Ch'dat'an(I)
Ch 'dat 'ana (U)
Vech 'eda
Ghelguts 'I k 'una (N)
Dghezhi, dghezha (O)
Dgheyay (U)
Dghezhay (I)
Vek 'eha qilani (NL)
Tuyiga (II)
Bente qiyuya (U)
Ghelguts 'I (I)
Tl 'egh tuzhizha
Shish (L)
Zdlaghi (L)
Duch 'ehdi (IU)
Dehch 'udya
Lih (O)
Dghilijuna (NL)
Dghili chuna (II)
Dghelay tsebaya (T)
Zhuk 'udghuzha (I)
Bat (Su)
Tuni (I)
'gaping thing pointing up'
?
'the one beneath rocks'
'one that hands out from chin'
'sand fish'
? 'salmon roe female'
'dog windpipe'
'arrow nock'
'one with a blanket'
'It's blanket'
'pike's food'
'thorny one'
'one with quills'
'water spirit'
'one going in lakes'
'swift swimmer'
'grass water beak'
'one that runs'
'open mouth one'
'mountain dark one'
'mountain fish'
'spiny mouth'
'water one'
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Dolly Varden trout
Whitefish (any)
Alaska whitefish
Broad whitefish
Broad whitefish stomach
Round whitefish, pin-nose
whitefish
Fish guts (all)
Fish bones
Fish backbone
Fish belly
Dark fish blood along backbone
Dark salmon meat near skin
Fins (any)
Pectoral fin
Dorsal fin
Pelvic fin
Anal fin and cartilage
Telaghi (U)
Shagela (II)
Qak'elay(I)
Qak 'elvaya (II)
Telch 'ell (O)
Chebay (U)
Liq 'a k 'qen (I)
Lih (UI)
Hulehga (I)
Q 'untuq ' (Lk-i)
Telay (L)
K'jida (I)
K'eghezh (Lk-i)
Hasten (IT)
K'inazdliy, vinazdliy
K'iztin (IO)
K'iytin (U)
K'eyena
K' eve da
K'tl'ech' (I)
K'kuhchashga (I)
K'kukelashch 'a (L)
K'chashga (U)
K'kuhchash 'a (O)
Beyes tut ' tsen (UO)
K'ts'elghuk'a(I)
K'ch 'elna (OU)
K'tay'a(U)
K'ch 'enla (U)
K'ts'elghuk'a(I)
K'iniq ' ts 'elghuk'a
Ghuk'a (I)
Biniq ' ch 'elna (U)
K'inhdegga (O)
K't'egha(U)
nilk 'degga (O)
k 'eveda degga (I)
nich ' k 'eltin 'a (O)
K'tselts 'ena (U)
K'tseldegga (IO)
'one that swims, runs'
'fish'
?
?
'shiny one'
'fish'
'salmon's husband'
'runs up'
'ridge on top'
'swimmer'
'oval'
'pus handle'
'inner objects'
'inner long object'
X
X
X
X
'wings'
'paddle'
'wing'
'back fin'
'back swimmer'
'back wing'
'back collarbone'
'paddle'
'paddles together'
'belly fin'
'one in the middle'
'anal bone'
'anal collarbone'
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Adipose fin
Tail fin
Fresh air sack
Fish collarbone, pectoral girdle
Fish head gristle
Fish meat
Fish tail
Meat next to fish tail
gills
Gut with stringy end (pyloric
caecum)
Fish heart
Hump on salmon's back
Male sperm sac
Sperm, milt
Nose cartilage
Oily strip of meat in front of dorsal
fin of salmon
Roe, fish eggs
; OO
Roe sac
scales
Fish slime
net-making tool, net stringer
net rack
K'tagh 'a (IO)
K'tach 'etvasha (N)
Tak 'elbasha,
k'tach 'ebasha (OU)
K'kalt'adegga(O)
K'kalt 'a ts 'elghuk 'a (I)
K'kuhlet'
K'degga
K'enchigija
K'enut'
Duni (II)
K'kalt 'a
K'kalt' a veghun
K'q'eshch'a
K'delchezha (Oil)
K'delcheya (U)
K'jida
K'ggalggama (I)
K'ggalggamam 'a (IIOL)
K ' ghalggamama (U)
K'qaldema (T)
K'eyenghezha (OI)
Hest 'a vekuhlashga (I)
K'tl'ech'
K'ingija, k'engija (IOU)
K'ingeja (II)
K'ints'isq'a (U)
K'yin tseq'a (I)
K'intsiq'a (OI)
Q'in
K'q'inyes
K'gguts'a(O)
K'ggisga (IU)
K'esM'a (Oil)
K'tl'eshch'a (IU)
tahvil vel k 'etl 'iyi,
tahvil qeyltl 'ixi
tahvil dugula (I)
veq ' k 'etl 'iyi
veq ' nuk 'detggeni
'paddle'
'submerger'
X
X
X
'head cartilage'
X
'food'
X
'body of fish tail'
X
'rattle'
X
X
X
X
X
'back strip'
X
X
X
X
'with it he weaves net'
'on it he weaves something.'
'on it, it is dried'
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net mesh measure
fishing clothes
awl for stabbing salmon
bale offish
cutting board
dipnet, long-handled dipnet
short-handled dipnet
salmon dipnet (longer handle)
trout dipnet
dipnet frame
fish bait (on hook)
rabbit or ptarmigan guts used
for tomcod bait
natural rock hole fish bin
rock fish bin, fish cutting hole
fish box
fish club, seal club
angled fish fence, dipnetting dock
fish fermenting hole
gaff hook, branch hook, leister
fish hook
Note: eleven separate types of
named fish hooks
fishing hole, fish trap location
fish trap location
fish jigging hole in ice
ve» k'ettl'iyi
va liq 'a ch 'el 'ihi
ts 'entsel (U)
vava hal
veq ' huts 'k 'del 'esi
tach 'enil 'iyi (UO)
nch 'equyi (LN)
tach 'enil 'i (I)
shanlaghi tach 'nil 'iy (I)
taztin (I)
taztin duves (I)
k 'enelneha (O)
k 'inlneha (I)
k 'indneha (U)
k 'egh dghichedi
bel ch 'k 'nulneq 'i (O)
k 'entleh, k 'entleq ' (U)
tsaq 'a (I)
k 'usq 'a (NL)
k'esq'a(OIl)
k 't 'usq 'a (U)
shagela yashiga
tsik'nigheli (IO)
tanatl 'ini
chuqilin q 'a (O)
chaqilin q 'a (IU)
qishehi (IU)
k 'isheq 'i (II)
sheh (L)
shehi (O)
ihshak, iqshak (OI)
k 'inaq 'i, k 'eninaq 'i (U)
k'enq'a(OU)
k 'inq 'a, -k 'inq 'a 'a (I)
tach 'k 'el 'unt
tasaq 'a
tatsiq 'a (II)
ges aq 'a (L)
'with it, it is woven'
X
'dry fish pack'
X
X
X
'summer run dipnet'
X
X
X
X
X
'cutting cavity'
X
X
'woven into water'
X
'hooker'
Eskimo origin
X
'where we set object'
'water head hole'
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fishing line
fishing pole
fishing reel
fishnet
net-like fish drag
Russian-era fishnet
drift net
gunny sack net
seine net
sinew net
twisted willow bark fiber net
small hole, net mesh,
net drying rack
lead line
corks, floats
cork line
fish pew, pike
fish sealer, ulu knife
fish spreader stick
hoop fish spreader
small fish spreader
hand-held fish snare with handle
spruce root fish snare
fish stringer
willow fish stringer
fishtrap, woven basket style trap
she hi tl 'ila (O)
k 'inaq 'i tl 'ila (U)
iqshak tl 'ila (I)
iqshak ten (IO)
shehi ten (O)
k 'inaq 'i ten, k 'inaq 'i
nikena, k'niten, k'neten
(U)
shehi tl 'ila telcheshi (UO)
tahvil
nich ' nuk 'tasdun (SITy
setga (O)
saiga (U)
te»edi (I)
chidayiztl'ini tahvi» (I)
vel niqak 'idzehi
nebod (O)
ts 'ah tahvil
ch 'eq ' tahvil (IU)
k 'eniq ' (IO)
k'eneq' (OU)
tahvil denluh
duyeh vetsik 'teh 'i
duyeh vetsittehi (I)
tahvil ts 'esa (IO)
tahbiljija (U)
vetsik 'teh 'i
liq 'a el dalyashi (OU)
liq 'a vel telyayi (I)
vashla
bel k 'elggits 'i (U)
k 'enun 'i
nuk 'ilqeyi
dnalch 'ehi (I)
t 'utseyyi (O)
k 'entsa quggil (I)
qunqelashi quggil (OU)
k'e'eshtl'il(OU)
q 'eyk 'eda (IU)
taz 'in (IO)
toy 'in (U)
'hook line'
X
'underwater snare'
'in back is hole'
Russian origin
'one that floats'
'with it one scrapes in circle'
Russian origin
X
X
X
X
X
X
X
X
'little stone'
X
X
X
X
X
X
'tough willow'
'object that is in water'
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Note: Seventeen types of fishtraps
for different species and conditions
fishtrap funnel
inner basket
angled leads to trap
long stick ribbing on fishtrap
spiral sticks on fishtrap
branch drag material put in weir
inner spruce bark reflectors pinned
to bottom of weir
vertical stakes for weir
fish wheel
lead line
net-making tool
net rack
k 'eshjaya (I)
k'jaya (OU)
taztin (I)
talyagi (IO)
talyashi (U)
k'etnalvesi (L)
k 't 'un dighali (U)
k 't 'un dalghali (I)
tah 'iggeyi (U)
vejink'ehi (I)
dik 'ali
niqak'uquli (I)
niqaghetesi (U)
naqak'ulqu»i taz'in (O)
duyeh vetsik 'teh 'i
duyeh vetsittehi (I)
tahvil vel k 'etl 'iyi
tahvil dugula (IL)
veq ' k 'etl 'iyi
veq ' nuk'detggeni
X
'heart'
'long object that is set'
X
X
X
'under water turns white'
'stg. swims over it'
X
'scoop that turns'
X
X
X
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III. MODERN CULTURE
A. Interview Synopsis
Table 9 is a synopsis of respondents to the semi-structured interviews. The interview
process is described in the Introduction and readers should refer to that section (II. A) and note
the questions were not designed to elicit a simple yes/no-type response (nominal data) but rather
to elicit a narrative of how the interviewee felt about or understood the topic in order to give a
richer and more nuanced understanding of cultural patterns and values. The "Voices of the
People" are a reflection of those deeper understandings. However, Table 9 has been derived from
the interviews in order to give the reader a sense of the overall consensus or variation from
consensus of the respondents. To accurately depict cultural practices, we read the interviews and
characterized the response as Agree, or Disagree/Neutral for each interview question, generating
nominal data. This data includes 47 interviews, the number transcribed at the time of the analysis
(un-transcribed interviews were from Dillingham and Pedro Bay). Sometimes respondents in a
group took up a topic at a later time during the interview in which case we included that response
as it applied to a previous question. As discussed in the Introduction, not everyone responded to
every question. In a small-group setting often one person would respond and others would nod or
otherwise express agreement with the speaker. We only recorded the verbal response, not non-
verbal indications of concurrence in formulating the data in Table 9. A second reason not every
responded to every question concerned the well-being of Elders. If Elders were tiring in the
course of the two-hour sessions, or if the session went long, we often skipped questions to
shorten the interview time.
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The responses represent consensus or near consensus: 694 responses were positive and 18
were negative or neutral. The data indicate Elders and culture bearers reflect indigenous cultural
standards that have a very high degree of homogeneity as represented by this set of questions
revolving around the importance of salmon and streams in their lives. Responses to interview
questions are used in the Modern Culture sections (III) that follow with statements like:
"interviewees universally felt...," "interviewees predominantly stated...," or "interviewees
indicated...."
Table 9. Nominal Evaluation of Interview Responses to Semi-Structured Interview
Questions. Based on 47 Interviews.
Question
1 . Are salmon critically important in your lives?
Note: often asked: "If the salmon were to disappear for whatever
reason, how would it affect your lives?"
Agree means people perceive salmon to be critically important in
their lives. Disagree means salmon are not perceived to be
critically important.
2. How many times in a week or a month do you eat salmon or
other fish? Is it different during different seasons?
Agree means three or more times a week or "all the time. "
Disagree is less than three times a week or "seldom. "
3 . Do people in your village need to eat salmon to be healthy?
How does salmon maintain or improve physical or emotional
health?
Agree means people perceive they need salmon and other wild
foods to be healthy. Disagree means they do not perceive salmon
to be necessary for health and wellbeing.
4. Which foods are important to give to a child so that he or she
will grow up to be smart or strong?
Agree means salmon and other wild foods are perceived to be
necessary for children 's health. Disagree means salmon and wild
foods are not necessary and children can eat commercially
purchased food and be healthy.
Agree
40
35
37
30
Disagree or
Neutral
0
0
0
2
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5. Does it matter to you if the salmon you eat is wild salmon? Does
it matter to you if the salmon comes from the streams and rivers in
your area?
Agree means people perceive that the salmon they harvest and
consume must be wild salmon from local streams. Disagree means
it doesn 't matter where the salmon comes from.
40
6. Does it matter to you that the salmon are connected to the
salmon your ancestors ate?
Agree means salmon genetically connected to fish their ancestor's
ate is perceived to be important. Disagree means there it is not
important that the salmon are genetically connected to ancestral
harvests.
27
7. If the fishing practices and care for the streams and rivers are
good (what the ancestors call, 'without' impurity, Dena'ina
beggesh quistlagh), does it result in salmon coming back?
Agree means proper practices are perceived to result in the
salmon's return. Disagree means practices have no effect on the
salmon's return.
37
8. Have you observed changes in the numbers of salmon that come
back each year? Is there a big difference some years? If there is,
what do you think causes these differences?
Agree means people have observed changes in the number of
returning salmon. Disagree means people have not observed
changes in number of returning salmon.
31
9. Are salmon important for the lives of other animals or birds that
are important to the Yup'ik or Dena'ina ? What would happen to
these animals or birds it they can't eat the salmon?
Agree means salmon are important to other animals. Disagree
means salmon are unimportant to other animals.
35
10. Who do you share food with? Relatives in Anchorage,
Dillingham? Elders? Who decides how to share the salmon, and
who to give salmon to?
Agree means wild food is shared with family and/or friends living
outside of the area. Disagree means wild food is not shared
outside the area.
31
11. Do you share salmon with people who don't do subsistence
and what type of things to you get in return?
Agree means salmon are shared with people who don't do
subsistence. Disagree means salmon are not shared with people
who don't do subsistence.
14
12. What does it mean for families to go fishing together? Do
young people learn a lot at fish camp? How do you teach the
41
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young people to catch salmon? Do you teach young people to
respect the salmon?
Agree means it is important for families to fish together. Disagree
means it is not important for families to fish together.
13. How do you feel when you give salmon? How do you feel
when you are given salmon?
Agree means people feel good when they give or receive salmon.
Disagree means people have no particular emotion when they give
or receive salmon.
33
14. Do you feel an obligation to return the favor when someone
gives you salmon?
Agree means people feel no obligation to return the favor of a
salmon gift. Disagree means people feel an obligation to return the
favor of a salmon gift.
15. Are salmon and other wild foods eaten in community
celebrations? Is this important?
Agree means it is important to include salmon and wild foods in
community celebrations. Disagree means it is not important that
salmon and wild foods are included in community celebrations.
27
16. It has been said that most Yup'ik/Dena'ina believe that a
wealthy person is one with a large family. Do you think that family
is more important that material wealth?
Agree means the person believes family is more important than
material wealth. Disagree means material wealth is more
important than family.
36
17. Do you do anything to make sure the salmon will return?
Agree means people do specific practices or rituals to assure the
salmon return. Disagree means people do not do any specific
practices or rituals to assure the salmon return.
37
18. What would it mean to treat salmon badly? Why is this bad?
Agree means there are specific things that are identified as bad
practices with disagree consequences. Disagree means there are
no specific things identified as bad practices with disagree
consequences.
19. Did the old people tell of a time when there would be a disaster
and the fish would disappear?
Agree means people heard elders tell prophetic stories of the
disappearance of salmon. Disagree means people never heard
Elders tell prophetic stories of the disappearance of salmon.
15
20. Do you ever thank the salmon for offering itself to you? Do
you ever pray when you catch salmon? Do you make an offering
when you catch the first salmon?
37
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Agree means individuals give thanks through a prayer and give an
offering when the first salmon is caught. Disagree means no
prayer, offering or other recognition is given with the first salmon
catch.
21. Do you ever hear the Elders talk about the salmon having a
spirit?
Agree means people perceive salmon to have a willful spirit.
Disagree means people do not perceive salmon to have a willful
spirit.
19
22. Did you ever hear Elders talk about a stream having a spirit or
being like it was alive? Do some people still think that way?
Agree means people perceive of a stream as having a spirit and
being alive. Disagree means people do not perceive of a stream as
having a spirit and being alive.
23. Do rivers or streams have events - or stories - associated with
them that are good or bad? Is it appropriate to tell any of them
now?
Agree means there are stories associated with streams that have a
moral implication. Disagree means there are no stories associated
with streams that have a moral implication.
24. How do people get money to buy boats and motors for
subsistence fishing?
Agree means people commercially fish in Bristol Bay or engage in
other part time employment. Disagree means people do not engage
in Bristol Bay commercial fishery or other part-time employment.
16
25. Do you feel a connection between the way you fish today and
the ancestors' way of fishing?
Agree means people feel an emotional connection between
subsistence fishing today and the subsistence fishing of their
ancestors. Disagree means people feel no such connection.
26. Why do you live in your village?
Agree means people desired to live in their village and felt an
emotional attachment to their lifestyle. Disagree means people
were ambivalent or disliked living in their village or felt they had
no future there.
39
27. Is there anything else you'd like to say? Is there any message
you'd like to convey to Washington/EPA (Environmental
Protection Agency)
Total
N.A.
694
N.A.
18
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B. Subsistence
1. Voices of the People
It may be different, the way we gather it nowadays, but it's the same end product. It's the same.
F-69, 9/18/11
If you get out in these outlying villages, about 80-90% of what they eat is what they gather from
their front yards. I was in Igiugig this spring. A can of SPAM... Do you know how much a can of
SPAM is in Igiugig? Eight dollars for a can of SPAM! ... There are fewer jobs, so subsistence is
one of the main cultures and the driving force of the economy within a community. M-60,
9/16/11
Our fish is more important for them. I tell my kids andgrandkids with fish they are very rich;
without fish you are hungry. This is the important thing all over in Alaska for us. It is very hard
out here in the bush. We have to pay double for every food we get, double to get our heating fuel,
double for gas, and without gas, we cannot travel. It is very hard in a rural area. In a big city it
is easy; you just grab everything from the store, department store. Out here we don't have
grocery stores; our grocery store is very expensive. They give us prices that, if you buy one item,
you pay for four. So it is very hard for us, but we grow our kids, and you ask us if it is important
for us to have fish. We have to have fish every day because the fish is most important. F-48,
8/20/11
For two families we put up in jars 32 cases [of salmon].... that doesn 't include frozen stuff. M-60,
9/16/11
We get them [smelt] until freeze-up here. Then, when the river freezes up, people go up and fish
through the ice for them with hooks. They seine them up in the lake, too, but you have to catch
them at the right time. M-62, 9/16/11
When that first salmon is caught, it is in the news. KDLG [Dillingham radio station]. Everybody
knows about it. M-61, 9/16/11
And he still, to this day, goes to fish camp. He gets all excited about fish camp. He's down there
getting his net ready, and he still, at 89 years old, still go out and sets his own net, picks his own
net, and work on his own fish, because he knows, and he always tells us how important it is to
save our fish and salmon for the winter months. F-32, 8/18/11
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We would starve if we don't have fish or salmon. In this area we have lived with fish all our
lives, from generation to generation. The people that stayed before us and kids that are behind us
will be living on fish. Salmon is very important; all kind of.... Without fish we are very poor; we
have no food to eat. With fish we are very rich; our stomach is full. That's the way I look at it. F-
48, 8/20/11
Salmon is one thing. They make you feel rich because you have something to eat all winter.
Smoked salmon, sun-dried spawned-out fish; all of those make you feel good, because you grew
up with it, it is in your body. Any subsistence food; what you eat, like him and I [gestures]; we ate
it for a long time. M-53, 8/20/11
Salmon is very important to us. I don't think we could live without fish.... I'm seventy-six years
old, and I have never been without fish, since I was small. I don't know how I would feel without
it. I think I used fish more than meat when I was growing up, because my Grandma raised me,
and that's all she could get, was fish, because it's easier to get. She used to help people put up
fish for us to have her share in the wintertime. Then she would put up salt fish for us to have in
the winter, so we use it year round. F-27, 8/17/11
Minority View Subsistence
We couldn 't live like our parents lived, because it doesn 't exist anymore. I mean, we could fish
and catch fish and stuff like that. You know, nowadays, you can't live on fish like you used to.
You can't even get meat like you used to; you can't even go out hunting for moose or caribou.
Nothing is here anymore; everything is disappearing. I know, you know [name] could verify too.
There used to be so much caribou, we would see them all over the road, all over the lake,
everything. F-44, 8/19/11
Like she was saying right now, even with subsistence, we can't live on that. We have to have
money to pay for our bills, telephone, our lights, our heat and trash, our toys, water, and sewer.
You have to pay so much a month for that. I myself will support any kind of entity that comes and
bills for jobs. I don't think subsistence; we love subsistence, but I don't think it is going to last
forever.... We need money to pay our bills. That is why a lot of people are moving to Anchorage.
M-44, 8/19/11
We can't just go out there and get money from nowhere. You know, subsistence is gone in this
village [Iliamna] and in Iliamna. Subsistence, we can't live on subsistence anymore. We have
car payments to pay, we have Honda payments to pay, andwe have our snowmobile payments to
pay. How on subsistence; how are you going to pay all of those bills? Some pay $500 a month
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for car payments. How are you going to pay $500 a month on subsistence? You can't do that
anymore; you have to live to make money nowadays for those young kids. M-49, 8/20/11
2. Introduction
In southwest Alaska subsistence is a fundamental non-monetized economic activity of the
region and forms the basis of cultural life. Though the economy involves both cash and
subsistence sectors, most of the food comes from subsistence activity as indicated in the ADF&G
Division of Subsistence data reproduced below. Moreover, cultural and personal identity largely
revolves around subsistence. This concept is expressed in a 1988 film by Brink and Brink where
Dena'ina leader Fred Bismark highlighted the importance of subsistence when he said, "If they
take subsistence away from us, they're taking our life away from us." Two decades later that
remains true; Fall et al.(2009:2) wrote of the Nushagak and Kvichak drainages, "At the
beginning of the 21st century, subsistence activities and values remain a cornerstone of area
residents' way of life, a link to the traditions of the past, and one of their bases for survival and
prosperity." Bismark's statement and Fall's analysis as well as interview generated "Voices of
the People" at the beginning of this section illustrate the idea that subsistence is "life" and the
foundation of culture for the Nushagak and Kvichak watershed villages. Everyone who
responded to Question 1, Table 9 felt the loss of salmon would impact them negatively and
subsistence based on salmon and other wild foods is the cultural foundation for the region. Four
of the 53 interviewees felt subsistence was no longer tenable.
Subsistence is not a return to practices of earlier centuries but employs modern
technology. Nylon nets have replaced spruce-root or sinew nets; aluminum skiffs and four-stroke
motors have replaced kayaks or canoes; metal pots have replaced birch-bark or willow baskets;
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modern clothing has replaced sewn hides and skins; and freezers have replaced underground cold
storage pits. Moreover, subsistence activities follow management practices formulated by the
ADF&G, dictating bag limits and seasons. The results of these interviews and ADF&G research
cited below confirm that the diet is still largely based on wild foods caught and processed by the
people who live in the area; values, such as respecting the salmon and not taking more than you
need, among others, are still honored; and the identity of the people is shaped by the subsistence
process, just as it was in the past.
As described in the Pre-Contact and History sections (II A & B).), indigenous people in
the study area have been harvesting wild resources for at least 12,000 years and have intensively
caught salmon for at least 4,000 years. This immense time depth has shaped all aspects of the
culture, including social structure, political structure, and religion. Because Dena'ina and Yup'ik
are the dominant populations in the study area, and because healthy wild salmon stocks and
many other components of their traditional way of life still persist such as language, sharing wild
foods and sharing beliefs related to nature, the area has a cultural continuum with the past that is
rare in North America. In few places do the same wild foods as their ancestors ate dominate the
diet and shape the culture as they do today in the Nushagak and Kvichak watersheds
3. Subsistence in Alaska
The importance of salmon and other wild food resources in the study area is tied to
federal and state subsistence legislation. No other state in the United States so broadly grants a
subsistence priority to wild foods to indigenous peoples as does Alaska. Both federal and state
subsistence legislation apply to Alaska but they differ, and have resulted in two sets of legislation
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because of an inherent conflict between federal and state legislation over indigenous rights vs.
inherent rights.
Federal subsistence legislation began with the 1971 Alaska Native Claims Settlement Act
(ANCSA), which extinguished aboriginal hunting and fishing rights and, in return, charged the
Secretary of Interior and State of Alaska to "take any action necessary to protect the subsistence
needs of Natives" (La Vine 2010:30-34). The federal subsistence intent of the 1971 ANCSA
legislation was clarified in Title VIII of the 1980 Alaska National Interest Lands Conservation
Act, (ANILCA). ANILCA recognized the cultural aspect of indigenous subsistence stating: "the
opportunity for subsistence uses by rural residents of Alaska...is essential to Native physical,
economic, traditional, and cultural existence and to non-Native physical, economic, traditional,
and social existence (emphasis added)" (La Vine 2010:32). The language describing the
importance of subsistence to Alaska Native and non-Native rural communities is the same with
the only difference that "cultural" importance is included in Alaska Native subsistence users' list
of essential rights while that term is not included in the non-Native list of essential rights. That
language became the basis for federally recognized indigenous subsistence rights. Federal
ANCSA and ANILCA legislation set up a legal conflict between indigenous rights and state law.
The "Inherent Rights" clause in Article 1, Section 1 of the Alaska Constitution specifies equal
treatment under the law for all Alaskans and makes no provision for indigenous rights.
Consequently, subsistence became an important political issue in the early 1970s and remains so
today (cf AFN Federal Priorities, 2011, pp. 1-9).
The State has developed subsistence legislation within the context of the "Inherent
Rights" clause cited above. As depicted in the 1988 documentary Tubughna: The Beach People
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by Brink and Brink, in 1973 Governor William Eagan made a promise to Alaska Native people.
Speaking at a meeting in Anchorage, Governor Eagan said:
Let me assure you that the state's commitment to preserving subsistence
capability in our fish and game resources is of the first priority and will continue
to be. Continuing attention to the Native for maintaining subsistence capability is
an integral part of the state's overall fish and game management program. It
always has been, is now, and will be so in the future. (Brink and Brink 1988)
That promise was partially realized as law in the 1978 State of Alaska Subsistence Act,
which provided for a Division of Subsistence within the ADF&G and defined subsistence as
"customary and traditional use." The act also specified a subsistence priority in wild resource
allocation over commercial or sport caught resources. The act did not limit subsistence to rural
(largely Alaska Native) residents and did not recognize indigenous rights; to do so would have
been unconstitutional in state law. The act also directed establishment of a Division of
Subsistence within the Alaska Department of Fish and Game to "quantify the amount, nutritional
value, and extent of dependence on food acquired through subsistence hunting and fishing" (AS
16-05.094) and has resulted in three decades of the most detailed subsistence data collected
anywhere in the world, some of which is used in this report.
As a result of over forty years of legislation and adjudication revolving around the
"Inherent Rights" issue among stakeholders, a dual management system has emerged. As
summarized by La Vine (2010:34) the state now manages fish and game for subsistence purposes
on state and private land including regional and village corporation land, while the federal
government, through the U.S. Fish and Wildlife Service or cooperative agencies, manages fish
and game in federally designated subsistence areas as determined by criteria applied and
regularly reviewed by the Federal Subsistence Board. On state lands all citizens are eligible to
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harvest fish and game for subsistence purposes but are bound by the customary and traditional
use criteria. On rural federal lands only rural residents are eligible to practice subsistence. On
non-rural lands subsistence is prohibited. Alaska Natives of the communities of the Kvichak and
Nushagak drainage fit both the "customary and traditional" and "rural' criteria and have engaged
in subsistence fishing and hunting throughout this time period and will continue to do so as long
as they remain rural. Significant non-Alaska Native population increases constituting a shift from
rural to urban would potentially change subsistence access as has happened, for example, on the
Kenai Peninsula where the Dena'ina do not have full subsistence rights because the area is
largely determined to be urban.
4. Scope of Subsistence
Table 10 is an indication of the importance of subsistence activities and salmon to the
people of the Nushagak and Kvichak River systems. Essentially everyone in every village and
town (98% or more of the households) uses wild food subsistence resources, and most (88% to
100% of households) use salmon.
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Table 10. Use and Reciprocity of Subsistence Resources. Data from Fall et al. 2009, Krieg
et al. 2009, Fall et al. 2005
Community
Dillingham
Ekwok
Igiugig
Iliamna
Kokhanok
Koliganek
Levelock
Newhalen
New Stuyahok
Nondalton
Pedro Bay
Port Alsworth
Year
1984
1987
2005
2004
2005
2005
2005
2004
2005
2004
2004
2004
All Wild Resources;
% Households that:
Used
98
100
100
100
100
100
100
100
100
100
100
100
Gave
62.7
86.2
100
53.8
82.9
92.9
85.7
80
73.5
92.1
88.9
72.7
Received
88.2
82.8
100
76.9
94.3
89.3
92.9
96
98
97.4
100
90.9
Salmon'
% Households that:
Used
88.2
89.7
100
100
97.1
100
92.9
100
89.8
92.1
100
100
Gave
34.6
48.3
83.3
30.8
62.9
60.7
35.7
64
55.1
55.3
72.2
45.5
Received
43.8
51.7
83.3
38.5
60
53.6
78.6
32
63.3
63.2
77.8
54.5
(Recent data collected by Steve Braun and Associates funded by Pebble Limited Partnership for
Environmental Impact Statement assessment includes more recent data not available as of this
draft.)
The data of Table 10 also indicates reciprocal sharing of wild foods is a fundamental
aspect of subsistence culture in the study area. In most villages almost 100% use wild food
resources and more than 80% of households receive shared subsistence food resources of some
kind. Sharing of salmon is lower than for all resources probably because, typically, extended
family units work together at subsistence fish camps (Fall et al. 2010) and the fish they
collectively harvest is not considered to be "shared" as much as "earned" among contributing
extended family members. Further research could clarify the matter. Sharing is further discussed
in Social Relations section (III. E.3). Table 11 presents subsistence resource data on a per capita
basis.
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Table 11. Per-Capita Harvest of Subsistence Resources. Data from Data from Fall et al.
2009, Krieg et al. 2009, Fall et al. 2005
Community
Dillingham
Ekwok
Igiugig
Iliamna
Kokhanok
Koliganek
Levelock
Newhalen
New
Stuyahok
Nondalton
Pedro Bay
Port
Alsworth
Year
1984
1987
2005
2004
2005
2005
2005
2004
2005
2004
2004
2004
Total
Harvest
Pounds
494,486
85,260
22,310
34,160
107,645
134,779
17,871
86,607
163,927
58,686
21,026
14,489
Estimated Per-Capita Harvest in Pounds
.Till
Resource
242
797
542
469
680
899
527
692
389
358
306
133
Salmon
141.4
456.2
205.2
370.1
512.8
564.7
151.8
502.2
188.3
219.4
250.3
89.0
A I*
7 ^ £
^H t« MH
17.5
68.6
59.4
34.1
36.3
90.4
39.9
31.8
28.0
33.9
15.3
12.0
Land
Mammals
65.9
249.2
207.8
32.7
95.9
186.2
257.4
104.5
143.4
81.8
30
24.7
Marine
Mammals
2.97
0
29.2
6.5
1.7
0
37.7
4.4
0
0
0
0
Freshwat
er Seals
1.7
0
7.4
6.5
1.7
0
4.5
4.4
0
0
0
0
c3
60
"S
PQ
0
0
21.9
0
0
0
33.2
0
0
0
0
0
(Recent data collected by Steve Braun and Associates funded by Pebble Limited Partnership for
Environmental Impact Statement assessment includes more recent data not available as of this
draft.)
Table 11 presents the range of some of the important subsistence resources used in the
region and their relative importance to each village on a per-capita basis. This data does not
include vegetation foods, birds/eggs, and marine invertebrates which are seasonally important,
nor does it include salmon retained from commercial fishing. While all subsistence foods are
important particularly for the physical and emotional benefits derived from a varied diet
salmon is, by far, the most important subsistence food ranging up to 82% of the subsistence diet.
Land mammals, including moose and caribou among other species, are the second most
important form of subsistence food for most villages. Many villagers but particularly Iliamna,
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Newhalen and Nondalton interviewees indicated that in recent years they are experiencing
reduced subsistence returns of caribou. They feel the Mulchatna herd is declining or moving out,
possibly due to overhunting from guided trips or seismic blasting and helicopter traffic from
mining exploration.
Non-salmon fish (northern pike, Dolly Varden/char, various whitefish, trout, etc.)
constitute a third important type of subsistence resource. Subsistence use of marine mammals
includes beluga whales, which regularly move up the Kvichak River, and freshwater harbor
seals, a unique freshwater population that lives year-round in Iliamna Lake. These are significant
subsistence resources for the Kvichak River villages of Igiugig and Levelock.
The data indicates as much as 899 pounds of dressed meat is harvested per-capita
(Koliganek) and an average of 503 pounds of meat per-capita is harvested per village. According
to the U.S. Department of Agriculture's "Agriculture Factbook," in 2000 Americans consumed
an average of 277 pounds of meat per year per-capita (USDA Factbook). The difference, of
course, is the subsistence data presented here is pounds per-capita harvested, not pounds per-
capita consumed. A substantial amount of subsistence-harvested food is shared which partially
accounts for such high numbers of per-capita harvest. The numbers are high, however, because
the people eat a lot of wild food and subsistence foods are the staple of the culture.
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Table 12 Per-Capita Harvest of Salmon Resources. Data from Data from Fall et al. 2009,
Krieg et al. 2009, Fall et al. 2005
Community
Dillingham
Ekwok
Igiugig
Iliamna
Kokhanok
Koliganek
Levelock
Newhalen
New
Stuyahok
Nondalton
Pedro Bay
Port
Alsworth
Year
1984
1987
2005
2004
2005
2005
2005
2004
2005
2004
2004
2004
Total
Harvest,
Pounds
494,486
85,260
22,310
34,160
107,645
134,779
17,871
86,607
163,927
58,686
21,026
14,489
Per-Capita Subsistence Harvest in Pounds
All Wild
Resources
242.2
796.6
542
469.4
679.6
898.5
526.7
691.5
389.2
357.7
305.5
132.8
o
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the most important subsistence salmon species in the villages of the Kvichak and Newhalen
River drainages and are also taken in significant numbers in the Nushagak River drainage.
5. The Seasonal Subsistence Round
As illustrated in Figure 5, the villages in the Nushagak and Kvichak River drainages have a
seasonal subsistence round that involves harvesting wild resources at an optimal time throughout
the year. Evanoff (2010:66) and Fall et al. (2010) have described the seasonal round for the
Kvichak drainage Dena'ina and it is summarized as follows. In the spring, with the return of
ducks, geese, and other waterfowl, small groups travel to hunting or egg gathering areas. In
addition, villagers also gather early spring plants, such as fiddlehead ferns. In late May and early
June, villagers begin harvesting salmon returning to spawn. Some families net salmon near their
villages while others travel to fish camp. Subsistence salmon activities occur throughout the
summer although many also engage in commercial fishing in Bristol Bay, depleting the fish
camp personnel but providing cash to support subsistence activities. Late summer and fall
subsistence activities involve berry and plant gathering. In late fall or early winter villagers
engage in caribou and/or moose hunting depending on the ADF&G-determined hunting seasons
for the specific area. Winter subsistence activities revolve around ice fishing for whitefish and
other freshwater species, ptarmigan hunting, wood harvesting to supplement home heating and
for steam baths, and trapping of furbearers.
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December
January
November
February
October
September
March
April
August
Subsistence Economy
Cash Economy
Religious Events
Time Spent in Village
Figure 5. Significant Aspects of the Subsistence Seasonal Round. Modified from Evanoff
(2010:66).
6. The Interplay of Subsistence and Wage Income
Interviewees indicate that, for those fully engaged in it, subsistence is a full-time job, but
it is necessary to supplement subsistence with cash from part-time wage labor or commercial
fishing, to defray the costs of subsistence activities. With gasoline costs presently in the $6 per
gallon range (summer 2011), trips to fish camps and other subsistence areas are expensive.
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Guns, ammunition, fishing gear, and modern winter clothing, among other expenses, also add to
the subsistence investment. While conducting village interviews, researchers observed that
besides having a skiff and motor powerful enough to navigate rivers like the Nushagak,
Mulchatna, Newhalen, and Kvichak, most families must also rely on one or more all-terrain
vehicles (ATVs) and snowmachines for subsistence, all of which require considerable initial
investment and maintenance costs. Rather than being recreational vehicles, these means of
transport have become necessary for the longer travel distances required for modern
subsistence. During the nineteenth century, dog teams, canoes, kayaks, and foot power via
snowshoes or hiking were the primary means of transportation, and people, by necessity, lived
in small villages located close to subsistence resources. In contrast, the twentieth-century
establishment of trading posts/stores, schools, churches, and health services led to residents
consolidating in fewer, larger villages. For example, today, there are only three interior villages
on the Nushagak River whereas, in the mid- to late nineteenth century, there were eight
(VanStone, 1967:114-115). The result of the consolidation is that village residents must now
travel farther to obtain subsistence resources, requiring mechanized transportation to do so, and
there is overlap among the range of village subsistence activities.
Interviewees indicate that to deal with these costs, many families report holding
commercial fishing permits and fish the sockeye run in Bristol Bay during late June and into
mid-July or engage in other forms of part time employment. Besides providing needed cash,
these forms of employment, with their short duration and/or seasonal nature, are ideally suited
to provide another ingredient critical to a subsistence lifestyle, time to engage in subsistence
activities. Thomas Lonner indicates that in Bristol Bay villages cash is obtained from wage
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employment such as working in the commercial fishery (also corporate dividends from
membership in Alaska Native Corporations and social welfare payments) and states "wage
employment is intended to underwrite subsistence equipment; the time, energy, and opportunity
cost in wage employment may be seen as an investment in subsistence" (Lonner cited in Lowe
2007:40). Table 13 is the number of 2010 Bristol Bay Fishing permit holders and crew member
licenses for the study area villages reflecting the major source of cash to support subsistence
activity.
Table 13. Commercial Fishing Permit and Crew Member Licenses
Dillingham
Ekwok
Igiugig
Iliamna
Kokhanok
Koliganek
Level ock
Newhalen
New Stuyahok
Nondalton
Pedro Bay
Port Alsworth
Commercial Permit
Holders, 2010
227
3
4
15
9
18
6
11
24
6
O
2
Commercial Crew
Member
Licenses, 2010
272
5
4
26
19
25
10
1
43
6
0
4
Subsistence
Permits, 2007
n.d
n.d
6
54*
29
n.d
1
n.d
n.d
29
19
30
2010 Data from ADF&G Commercial Fisheries Entry
Commission, http://www.adfs.alaska.sov/index. cfm?adfs=fishinscommercial. main
2007 Data from Fall et al. , 2009, page 19
* Combined data for Iliamna and Newhalen
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Figure 6. Subsistence Skiffs, Nushagak River, New Stuyahok. May, 2011
7. Subsistence as an Economic Sector
Labor statistics do not identify subsistence as an employment category because it is not
based on wage-labor or a salary and, hence, people engaged in subsistence are considered
"unemployed." However, those who choose the subsistence lifestyle work long hours, utilizing
considerable skill to provide food for themselves and their families and in interviews described
subsistence as a full-time occupation.
The official unemployment rate in the study area ranges from zero (Igiugig, Iliamna,
Pedro Bay and Port Alsworth) to 31.1% (Koliganek). The weighted average is 10.9%; compares
to 8.0% for Alaska and 9.6% for the U.S.
(http://www.census.gov/compendia/statab/2012/tables/12x0629.pdf). The unemployment rate
includes only people actively seeking wage-based employment and does not include villagers
for whom subsistence is their non-wage employment. The percentage of working-age
population "not in labor force" (http://www.bls.gov/cps/cps_htgm.htm#nilf) may better reflect
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how many people might seek employment should subsistence salmon/other resources no longer
be available.
Based on 2010 U.S. Census Data, 4.0% (Port Alsworth) to 44.5% (Nondalton) of the
residents in the study area communities have wage incomes below the poverty level. The
weighted average for all communities (excluding Pedro Bay) is 17.1%. These rates compare to a
9.1% rate for Alaska and a 15.1% for the U.S. (DeNavas-Walt et al. 2011:14). These numbers
are high but do not reflect the role of wages in a subsistence economy: wage income which for
many is not considered the primary source of sustenance but functions to support non-wage
subsistence activities. Neither do the statistics consider the non-monetized value of subsistence
foods to the economies of the villages.
Subsistence is dictated by the seasons, is time-consuming and must be understood
differently from recreational fishing or hunting. It is not critical if a recreational fisher or hunter
misses a season due to work obligations or other demands, but, for many Bristol Bay village
residents, subsistence is one's work obligation and employment in the cash economy impinges
on the time that is necessary to obtain and process food for a family for a year.
Thornton (1998) writing in the on-line edition of Cultural Survival Quarterly, considered
Alaska subsistence to be the leading employment sector of rural Alaska because of the number
of people engaged in subsistence and the economic benefits derived from harvesting one's own
food Several attempts have been made to measure subsistence economically by monetizing
wild food resources. Fall et al. (2009:3) measured the economic importance of subsistence by
calculating the cost of replacing wild foods obtained from hunting, fishing, and gathering with
similar foods obtained in a market. Their published data indicates the average annual per-capita
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harvest of wild foods in the villages of the Nushagak and Kvichak River drainages is 304
pounds of salmon, 123 pounds of land mammals (mostly moose and caribou), 39 pounds of
other fish, 23 pounds of plants and fungi (mostly berries), 9 pounds of marine mammals
(freshwater seals and beluga whales), 8 pounds of birds and eggs, and one pound of marine
invertebrates (mostly clams). To supplement their subsistence harvest, households in the
Nushagak and Kvichak River drainages spend 15 to 26% of their annual cash income on store-
bought food (Fall et al., 2009:3). In the ten villages for which there is recent data (i.e., excluding
Dillingham and Ekwok), the annual per-capita cost of purchasing food ranged from $1,467 to
$2,622. At 2004 prices (when the initial analysis was done), the annual replacement cost for the
average subsistence harvest described above would be an additional $7,000 per capita, which
would increase the demands on the annual cash income an average of nearly 80% ranging from
23% for Port Alsworth to 157% for Koliganek. As high as they are, the estimate may be an
under-representation of the estimated worth of subsistence resources. With rising food prices,
the replacement value would be significantly higher today. King salmon fillets, for example
were $17/pound on December 30, 2010 at 10 and M Seafoods, Anchorage, Alaska. The
replacement value of 193 pounds of king salmon alone for Koliganek, for example, would be
$3281 per-capita.
While monetizing subsistence gives a measure of its importance to the economy, these
values do not reflect the fact that the people of the region unanimously reject replacing their
traditional subsistence foods with farmed fish or other imported products, should deterioration
of wild salmon runs occur (Interviews). This is based on the belief that such products are of
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inferior quality and that doing so would result in cultural degradation. See Section III.C.6 for a
discussion of the importance of wild salmon from one's home river.
Figure 7. Salmon Drying. Koliganek. September 17, 2011
8. Subsistence and "Wealth"
In Alaska many non-Native people perceive subsistence as an activity for impoverished,
unemployed rural people who live in employment-poor communities and cannot afford to buy
food so they have to hunt and fish for it. Thornton (1998) asserts that this perception relates to
the "minimum food and shelter necessary to support life" dictionary definition of subsistence and
has given rise to the "subsistence-as-welfare" concept and associated negative implications. The
Yup'ik and Dena'ina perceive subsistence quite differently. Interviewees spoke of the cultural
value of subsistence as a chosen lifestyle. As indicated in the 2011 interviews, subsistence is a
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lifestyle chosen by both old and young. Subsistence is a job, in which the wages are healthy wild
foods and the benefits include not only vigorous outdoor activity shared with friends and family,
but also a large measure of self-determination supported by a community of like-minded people.
Subsistence is coterminous with culture, and the entire range of social and spiritual activities that
"culture" implies. Consistently, the Yup'ik and Dena'ina communities of the Nushagak and
Kvichak River drainages define a "wealthy person" as one with food in the freezer and the
freedom to pursue a subsistence way of life in the manner of their ancestors (see Social
Relations).Their ability to continue their reliance on subsistence and their concept of wealth has
contributed to the maintenance of vital and viable cultures for the last 4000 years.
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C. Physical and Mental Well-being: the Role of Subsistence
1. Voices of the People
We crave it [salmon] when we don't have it. We just need it. F-30, 8/17/11
You know, it's got that one oil in it that is a cancer-fighting oil, and it's really good. F-3 8,
8/18/11
/ think it [salmon] is healthier than probably beef or pork or something like that. M-68, 9/18/11
Yes, to be healthy, like I say, if we don't eat fish we won't have anything to eat. That is our
health. F-48, 8/20/11
When you are eating fish...you get a drink of water to flush yourself out. If you don't eat fish, you
will starve. You got to flush yourself out with water every day; that is what your health is about.
God put us on this earth to eat fish every day. That's what it is. Without fish, like I said, we are
hungry; with fish we are full. F-48, 8/20/11
We have... to live healthy to be free from diseases if we eat healthy food. Not breathe air that's
no good or drink water that is no good; it will affect your whole body. So, on the subsistence, I
say let's protect Mother Earth; I demand it. If we don't protect Mother Earth, we are gone. M-
51,8/20/11
We don't buy meat very much. Salmon is our most important dish. F-27, 8/17/11
Salmon is a really an important part of our diet. I think it has things that meat [domestic beef for
example] does not have. You are always hearing things about fish oils and how healthy [they
are], but we already have that, so we must be healthy. F-34, 8/18/11
We can't live without salmon. We 'II be missing something. F-27, 8/17/11
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Well, we grew up with it. We need it. If we don't have it, we miss it. I can't see anybody that lives
around here without it. F-30, 8/17/1 Ic.
I've seen kids teethe on smoked salmon strips. They 're hard. They get all fishy and smelly, but
man, they just chew. It's better than the rubber toy. F-38, 8/18/11
... [salmon] is one of our healthiest foods we can give to our child.... It is really healthy. F-69,
9/18/11
To me, I think eating salmon has sustained our ways of life. I think by eating a lot of salmon, we
are a healthy, healthy Dena 'ina. I always tell children there atpotlaches or wherever; I say that,
"Ifyou eat this piece of fish you 're going to be a smart Dena 'ina woman, you might be able to
be a lawyer or a doctor. " It's surprising that, just by telling them that, they ...eat it, and they will
say, "Oh, taste good. " F-32, 8/18/11
When my kids grew up, I mostly gave them fish and moose meat. F-44, 8/19/11
I definitely limit my child; you know, the fast foods, we eat it once a week, sometimes more...
[They eat] moose meat, the fish...berries, and wild plants as well... We want to give to our
children the fish and we want to keep the water clean for them. It was a gift to us from our
ancestors, which will then be given to our children. F-69, 9/18/11
The school system here does get volunteers who donate fish to the schools. Prior to that they
used to order codfish and other fish from out of the area. The kids didn 't like it. Not from here.
They finally started the donation program, and the fishermen stepped up to the plate and said,
"Yes, definitely. " The crew members didn't balk. There were no qualms whatsoever about
donating fish to the schools. M-61-9/16/11
It is the best hot lunch program we have; the kids just love it when they have salmon day. M-60,
9/16/11
Yes, and that it is healthy [wild salmon] ...and something they [Yup'ik] wouldn 't have without....
But if we ever lose it, then we won't have anything at all. M-68, 9/18/11
/ think it would matter [that the salmon be wild]; that would be our concern. We like to take our
wild natural renew able resource salmon rather than farmed salmon because you never know
what they've [farmed salmon] been eating. M-26, 5/19/11
Wild salmon is more important for us, or wild fish. I don't believe in farmed fish, because wild
fish is better for all our health. It has all natural oil, and we don't paint it with artificial paint
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like the farmed fish you get. You can sell your farmed fish all you want, but wild salmon is more
important to us. F-48, 8/20/11
...people from Kenai or Anchorage, they can go to Kenai and get their salmon, but they always
say there's nothing like the lake salmon. There's nothing like salmon that comes from Sixmile
Lake. We hear that all over.... I always try other people 'sfish, but there's nothing like salmon
from our own stream, salmon from the lake that comes up. Well, I guess we 're spoiled having
our own. F-32, 8/18/11
There is nothing better than wild salmon... I have talked to many people all over the state, and
the best salmon comes from this area, Bristol Bay. M-29, 8/17/11
One year we got a farmed salmon.... What a difference! It came in with the usual run, and it was
salmon that was raised in the University of Washington [salmon farm]. They have a big place out
there in Seattle. We went in there, and they had a lot offish. The meat was soft, and the skin was
not firm and scaly. I remember, my daughter was cleaning salmon that year, and she said,
"Where 'd this fish come from? It looks like a salmon, but it's terrible. " It was soft. It wasn 't
like a wild salmon. F-38, 8/18/11
Matter of fact... I had [salmon] for breakfast this morning before I come over. They stay inside
all day. M-53-8/20/11
In the summertime it is every day [we eat fish], as long as the fish are running. We eat fish every
way we could: boiled, baked, fried. Every way we could, we eat fish. In the wintertime, what we
preserve in the summertime is what we eat in the wintertime, like the dried fish, the canned fish.
The fresh canned is something we eat a lot, because you can do so many different things with it.
F-35, 8/18/2011
2. Introduction
As described in Section H.A.3., archaeological evidence indicates that salmon were an
important component of the diet of the probable genetic ancestors of the Yup'ik and Dena'ina,
who left evidence of their presence in this region up to 4,000 years ago. These genetic ancestors
of the present day Yup'ik had an important component of salmon in their diet as long as 4000
years ago, according to the archaeological record (see Section II.B.3). The Dena'ina track back
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to the Paleo-Arctic tradition, as old as 10,000 years ago, although evidence for intensive salmon
utilization in Dena'ina territory does not occur until A.D. 1000.
Based on studies of other Yup'ik populations in the nearby Kuskokwim River villages,
there is a strong possibility that, within their long history, the Yup'ik may have become
genetically adapted to eating salmon. Several recent studies have shown that physical adaptation
and evolution based on dietary factors (e.g., lactose intolerance) can occur in 3,000 years or less
(Tishkoff, et al., 2007; Bersaglieri et al., 2004: Hollox et al., 2001). Other studies are
demonstrating genetic changes at the population level in humans in a similarly short time frame
based on adaptation to environmental stressors such as living at high altitudes in Tibet (Peng et
al., 2010 :1075-1081; Xin et al., 2010: 75; Simonsen et al., 2010: 72-74).
The National Science Foundation recently funded a University of Alaska study to assess
the differences between Yup'ik and other populations in drug metabolism, as well as in
vulnerability to metabolic syndrome (development of risk factors for coronary disease, stroke,
and diabetes). This study will consider the relevance of dietary differences and resulting long-
term physical adaptation, including genetic adaptation. In a separate study, researchers from the
Center for Alaska Native Health Research (CANHR) are assessing how a subsistence diet affects
the vulnerability of Yup'ik people to disease (O'Brien et al., 2011). In a 2009 study whose
results strongly support the validity of red blood cell deltaN as a biomarker of eicosapentaenoic
acid (EPA) and docosahexaenoic acid (DHA); the researchers state, "the omega-3 (n-3) fatty
acids derived from fish, eicosapentaenoic acid (EPA; 20:5n-3) and docosahexaenoic acid (DHA;
22:6n-3) are associated with a reduced risk of cardiovascular disease and other chronic diseases
(O'Brien etal, 2009:913).
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While the amounts of salmon and other fish consumed varies from village to village, and
from one season to the next, the demonstrated importance of these foods in the diet is consistent
with the traditional knowledge shared by Yup'ik Elders and culture bearers, as presented above.
As discussed below, the salmon-dependent diet of the Yup'ik and Dena'ina benefits their
physical and mental well-being in multiple ways, as well as encouraging high levels of fitness
based on practices involved in subsistence activities.
3. Nutrition
The dietary habits of Yup'ik and Dena'ina living in the villages of the Bristol Bay region
shows regular dependence on several species of wild salmon which they sometimes consume
several times a day as the interviews attest. Yup'ik and Dena'ina primarily prepare and eat two
species of Pacific wild salmon, Coho (red) and Chinook (king) in different ways, including fresh,
salted, pickled, canned, dried, and smoked. Salmon and other traditional wild foods comprise a
large part of the villagers' daily diet throughout their lives, beginning as soon as they are old
enough to eat solid food (Interviews, 2011).
In addition to salmon, villagers also regularly consume other wild fish species, such as
humpback whitefish, Arctic char/Dolly Varden, Arctic grayling, rainbow trout, and northern
pike, the wild ungulates caribou and moose, and, to a smaller extent other mammals, birds, and
bird eggs. Wild plants, including blueberries, crowberries, salmonberries, ferns, and other
species, add fiber, vitamins, and minerals (Interviews).The Yup'ik and Dena'ina continue to
harvest certain plants with medicinal values (cf P. Kari 1995). It is important to recognize that in
addition to providing a wide range of valuable nutrients and protein sources, the subsistence diet
provides a year round workable harvest schedule with adequate time for preparation and storage.
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While subsistence technologies have changed and are now supported in part by the cash
economy that commercial fishing provides, enabling purchases of snow machines, rifles and
other equipment, the basic subsistence seasonal schedule has been approximately the same for
hundreds and probably thousands of years. The implications for population sustainability within
the environment, and co-evolution of the human population with environmental food availability
mean that hypotheses about the risks of significant changes to the salmon population are
important, and change in dependence on local wild salmon could have far-reaching impacts on
Yup'ik and Dena'ina physical and psychological health, including at the genetic level.
Villagers in the study area also eat store-bought foods, but do not prefer them (Interviews
2011). Like other northern subsistence cultures, the Yup'ik and Dena'ina consider their
traditional foods to be healthful and satisfying, in addition to providing strength, warmth, and
energy in ways that store-bought food does not (Hopkins, 2007:42-50). Hopkins' study on health
and aging also provides an insight into women's views of the importance of the subsistence diet.
Eating subsistence foods was an overwhelming theme among all participants.. They generally
viewed market or kass 'aq (white person) food as unhealthful (Hopkins, 2007:46). Hopkins
quotes one of the participants, describing the importance of the subsistence diet for Elders: "In
years back, before I was born, I know there were elders that were very healthy and strong
because they have their food, their native food, not mixed up with the kass 'aq food. Although
they have a hard life, they were healthy, strong, because of their native food. Seal oil, dried fish"
(Hopkins 2007:46-50).
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4. Fitness
Yup'ik and Dena'ina dependence on subsistence foods has the additional health benefit
of providing opportunities and incentive for physical fitness, since engaging in subsistence
harvesting improves fitness and fitness, in turn, enhances the efficiency of subsistence
harvesting. Subsistence hunting, fishing, and gathering demands stamina to endure long periods
of physical activity and strength to handle meat, large quantities offish and heavy fishing gear .
Hopkins (2007:45-46) quotes from the response of one study participant, over sixty years of age:
"I think today most of the women are healthy for activity, physical activities. When they go berry
picking, they're working using their bodies everything. When we are cutting fish, we are using
everything, our muscles, lifting things."
The fitness needed for and resulting from subsistence is part of other aspects of village
life, as well. Throughout the winter the Yup'ik villagers, from youth to middle-aged, play
basketball and other sports regularly competing in vigorous games. Researchers watched in New
Stuyahok as a team of middle-aged men defeated a youth team in an intense, hour-and-a-half
game, then went to church services for an hour and returned to play another game of equal
length. In several Yup'ik villages, including New Stuyahok, the physical activity of traditional
dancing, is making a comeback. As described in Section III.E., this cultural activity is based on
dance as story-telling, which both values and elaborates on traditional cultural practices, such as
fishing.
While in New Stuyahok, researchers observed that Elders, including the oldest present, at
around age 86, frequently walked to locations within the village. According to Hopkins, walking
was the primary physical exercise identified in that study's interviews. "The participants referred
to walking as an important component of health, both physical health and mental well-being.
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Walking is believed to keep the body strong, promote energy, and is a basic physical activity in
gathering subsistence foods" (Hopkins 2007:46).
The apparent overall fitness of the village population in New Stuyahok gave researchers
present at the Elders' Conference the impression of frequent exercise, and led to the hypothesis
that the practices of subsistence food gathering, in addition to the food itself, create higher levels
of fitness, and act to prevent and reduce health risks from more sedentary lifestyles. For Alaska
Natives, as for other Native Americans, the high risk of diabetes and subsequent health
consequences is serious enough to make the hypothesis an important one to test.
5. Disease Prevention
Beyond the Yup'iks' own personal conceptions and cultural knowledge about the
importance of wild foods in their diets, many studies also confirm the remarkable health benefits
of omega-3 fatty acids and the other nutrients found in high percentages in subsistence foods
such as wild salmon, and the combination of salmon, wild greens, blueberries and other berries
for preventive health among the Yup'ik. These studies particularly underscore the importance of
salmon-rich diets for the prevention of maladies, including cardiovascular diseases and type 2
diabetes. O'Brian et al. (2009:913; see also O'Brian et al 2011; O'Harra 2011), for example,
concluded that "the omega-3... fatty acids derived from fish... are associated with a reduced risk
of cardiovascular disease and other chronic diseases."
In a cohort study of Yup'ik from the Yukon-Kuskokwim area (Boyer et al., 2007:2535-
2540), the Center for Alaska Native Health Research (CANHR) found that metabolic syndrome
is uncommon in that population relative to others, occurring at a prevalence of 14.7% in the
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study population, compared to 23.9% in the general U.S. adult population. The study population
also had significantly higher high-density lipoprotein (HDL) cholesterol levels and lower
triglyceride levels than the general U.S. adult population.
In another related study of the same population, the Fred Hutchinson Cancer Research
Center, in collaboration with the CANHR, found that Yup'ik Eskimos consume 20 times more
omega-3 fatty acids from fish than the average American and display a much lower risk of
obesity-related disease despite having similar rates of being overweight and obesity (Makhoul et
al., 2011; Fred Hutchinson Cancer Research Center, 2011). Lead author, Zeina Makhoul, said:
Because Yup'ik Eskimos have a traditional diet that includes large amounts of
fatty fish and have a prevalence of overweight or obesity that is similar to that of
the general U.S. population, this offered a unique opportunity to study whether
omega-3 fats change the association between obesity and chronic disease risk....
It appeared that high intakes of omega-3-rich seafood protected Yup'ik Eskimos
from some of the harmful effects of obesity.... While genetic, lifestyle, and
dietary factors may account for this difference, it is reasonable to ask, based on
our findings, whether the lower prevalence of diabetes in this population might be
attributed, at least in part, to their high consumption of omega 3-rich fish
(Makhoul quoted in Woodward 2011).
Compounds derived from their subsistence diet, including omega-3 fats from wild salmon
consumption, may also benefit mental health in Yup'ik populations. Lesperance et al. (2010), for
instance, report that omega-3 fats can help prevent depression. Another study showed greater
improvement in symptoms for patients with chronic depression who consumed omega-3 fats
with their medication compared to those receiving only a placebo with their medication. After
four weeks significantly reduced symptoms of depression occurred in six often patients
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receiving E-EPA while reduced symptioms only occurred in one often receiving a placebo
(Nemets et al. 2006).
Other subsistence foods, such as wild greens have nutritional elements associated with
better mental health, including folic acid and Vitamins A and C. Other factors associated with a
subsistence lifestyle, including time spent outdoors and the physical fitness resulting from
subsistence activities, may also benefit mental health. It is interesting to note that several Elder
interviewees (Interviews 2011) said that, 20 years ago, no one in their villages knew anything
about Alzheimer's disease; it was not an illness they had seen before, but it is appearing now.
6. Local Wild Fish
The Yup'ik population of Bristol Bay Region has an interdependent relationship both
ecologically and nutritionally, and possibly evolutionarily, with the local wild salmon
populations. It is clear that the benefits, and particularly the long term fit between the human and
fish populations, depends upon maintaining the local wild salmon for subsistence fishing. While
it would be easy to assume that any salmon would provide a similar quantity and quality of
omega-3 fats, a Norwegian study showed that farmed salmon, fed a typical farmed salmon diet,
did not have the omega-3 fats in beneficial quantities, in contrast to the wild salmon which did
(Sincan, 2011).
It is important to underline that if a human population has adapted to particular
environmental dietary elements with a genetic modification in their population, that modification
is based on a relationship to the genetics of specific regional species, and subspecies. The fit
between environment and population may not be transferable to other places.
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Thus the elements of the subsistence diet, in particular wild salmon, provide several
substantial health and fitness benefits to the Yup'ik of the Bristol Bay region. According to
recent studies at CANHR led by Andrea Bersamin, "Diets emphasizing traditional Alaskan
Native foods were associated with a fatty acid profile promoting greater cardiovascular health
than diets emphasizing Western foods" (Bersamin et al., 2007: 266; see also Bersamin et al.
2008). The loss of the local wild salmon as a large component of the Yup'ik diet would result in
risks to the physical and psychological health of the population, including greater risks of
cardiovascular disease, type II diabetes, and depression.
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D. Traditional Ecological Knowledge
1. Voices of the People
But, I think, when they 're spawning, that's where they hit the spring waters, where it doesn 't
freeze. It's always open, even in the dead of the winter. It's always open; you got to be careful
there. Especially up in Lake Clark, around Kijik. It's, man, 30 below zero, and it's still open
water. M-29, 8/17/11
Our societies are not different than other societies we have special people that know fishing
inside and out, we have people in our society that know weather inside out, that know plants
inside out, and that know animals inside out. M-61, 9/16/11
...they drop last year 'sfish in the middle of the river and we do the same thing here. We put king
salmon remains on a string tied to a rock and go out with a boat to the middle of the river and let
it sink. That makes king salmon go on both sides [near the banks where they can be netted with
set nets.] M-26, 5/19/11
When the fish first come up here we don 'tput our nets out here before a bunch of them go by for
the people who live at the end of the river up in Nondalton and all those guys. They start calling
up then maybe middle of July [to tell us they have fish, and then] we start putting our nets out.
We just kind of watch the salmon go by for the people who live upstream from us. M-54, 8/20/11
They [the fish] are like us, when we want to know something we ask. The fish are the same way.
As we were talking about earlier he mentioned that the fish have souls. Every living creature has
a soul. All the animals have souls. They are sensitive, very sensitive. If you put something bad in
the water the fish will sense it. They will probably not go up the river, they will go somewhere
else. If they spawn here and they notice something different they will move to another spot. The
fish are very sensitive. M-20, 5/18/11
What they used to say, was the first time, when they first moved down to fish camps, then this
wild celery, I don't know if you know what that is, but we eat those. They go up on the
mountainside and pick lots of that, and then they peel it, they peel the peelings offandwe eat the
inside part. So we have big parties with that. We just really enjoyed the fresh salads that we just
had. it was already tall enough to eat. So when we get done with that, then the Elders would tell
us, take all the leaves and the skin and everything off of this plant, take it out in the river and
throw it in, and they would do that. Then we started asking why we were doing this. This fresh
salad plant and the skin will meet with the salmon, and let the salmon know that they are already
good to eat, and they need to hurry up and come up because we are hungry. F-28, 8/17/2011
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In the winter not only salmon, we do a lot of ice fishing, and my uncle you met this morning [a
man in his 90s], he has a trout net he puts out. F-35, 8/18/2011
2. Introduction
Anthropologists and other scientists have used different terms to describe the knowledge
of indigenous peoples, including "cultural knowledge," "indigenous knowledge," "traditional
knowledge," and "local knowledge" (Berkes 1999:8). Fikret Berkes and others working in this
area of ethnoscience use the term, "traditional ecological knowledge" or TEK. Berkes defines
TEK as "a cumulative body of knowledge, practice and belief evolving by adaptive processes
and handed down through generations by cultural transmission, about the relationship of living
beings (including humans) with one another and with their environment" (Berkes 1999:8). TEK,
as Berkes describes it, includes spirituality and social relations, as well as a wide range of
cultural beliefs and behaviors related to surviving in a particular landscape, because of the
holistic nature of culture itself.
Early studies of TEK depended on comparisons between taxonomies and species lists
drawn up by Western scientists and those created by indigenous peoples (Knott, 1998). More
recently, however, it has become clear to anthropologists, geographers, biologists, and others
working with indigenous peoples that their knowledge is far more ecological in scope and
requires more than species lists to document. Therefore, a number of scientists working with
indigenous peoples have come up with a diverse range of tools to collect and document
indigenous knowledge. These research tools include, but are not limited to:
Maps of local hunting, fishing, and gathering areas
Maps of sacred sites and other special use areas
Traditional Place Names mapping
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Species lists
Collection of stories, songs, and dances of interactions between animals, humans and
other species, humans and the natural environment, or allegorical animal stories
Studies of subsistence technologies
Animal life histories and their interactions with other plant and animal species including
humans, told as information by locals
Plant life histories and their interactions with other plant and animal species, including
humans, told as information
Stories of human mistakes made, and lessons learned, about interactions with nature and
the environment, including storms, earthquakes, floods, ice, glaciers, changes in nature
Advice in the form of rules, proscriptions against certain behaviors, prescriptions for
other behaviors, and guidelines for management of animal and plant harvests
Uses for animal and plant species, including recipes for foods and medicines
Observations shared, often about the complex interactions and ecological relationships in
the landscape where the people live, hunt, fish, and gather.
Local descriptions of long term trends for species numbers and migration patterns,
weather patterns, climate, and other natural events
Linguistic, biological, and archaeological evidence.
And finally, at a broader level, the values, beliefs, social systems and spiritual practices
that have developed over thousands of years through the cumulative application of TEK.
It requires months and years of patient work with indigenous groups to elicit and
document in-depth TEK. Researchers must spend adequate time in the field to understand the
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landscape and local ecosystems as well as the local cultures. More important, local people need
time to develop trust in the researchers' methods and personal qualities before they will be
willing to share such important confidential knowledge as hunting sites or productive fishing
locations. Fortunately, while the months afforded to this project were not sufficient to develop
new in-depth TEK studies with local populations in the villages, there are several existing
studies, both in the Nushagak and Kvichak River watersheds, and in the Lake Clark and Diamna
Lake area, that cover TEK in detail. Among the Nushagak studies is one by the Nushagak-
Mulchatna Watershed Council (NMWC) (2007) and among the Kvichak studies are those by
Stickman et al. (2003) and Fall et al. (2010). These long-term studies have focused on the Yup'ik
and Dena'ina TEK in the Bristol Bay region and have provided sufficient information for our
Bristol Bay TEK characterization, which we summarize in Sections a through c below.
To supplement those long-term studies, we focused interview sessions on the broader
levels of TEK, including the values, beliefs, social systems, and spiritual practices of the Yup'ik
and Dena'ina that have developed over thousands of years through their cumulative application
of TEK. During those sessions we learned much from the Elders and culture bearers about TEK
and the cultures as a whole. We also heard some specific examples of ecological insights,
prescriptions and proscriptions, and management guidelines for several species.
a. Nushagak-Mulchatna Watershed Conservation Plan
Over a two-year period [dates unspecified], the NMWC conducted interviews with
Elders, residents, and others who use the watershed to create a database of the TEK of the
Nushagak and Mulchatna drainages (NMWC 2007:3). The NMWC used the data to create an
overall plan for protecting the waters and natural resources of the watershed. The interviews
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helped with the development of maps to identify areas critical to protection of subsistence
resources and habitat. The plan identified 12 fish, 6 mammal, and 12 bird species important
for subsistence and mapped 125 traditional use areas and 153 traditional area names. The
flora and fauna considered most integral to traditional subsistence use were all five species of
Pacific salmon, whitefish, winter freshwater fish, moose, caribou, waterfowl, and edible and
medicinal plants (NMWC, 2007:19).
The study also identified probable threats to the watershed in the next fifty years, and,
based on the TEK information collected, developed four strategic actions:
1. Reserve adequate water flow for the Nushagak River and tributaries under existing
laws for in-stream flow reservation.
2. Maintain the vegetative complex that supports moose, fish and other species within
and adjacent to the floodplain.
3. Maintain water quality standards that protect wild salmon and other fish.
4. Prevent habitat damage that could result from mining. (Nushagak-Mulchatna
Watershed Council, 2007:3)
What is at stake includes habitat, and wildlife including terrestrial mammals, birds, fish,
and the subsistence way of life, along with the unique cultures it supports. The report states:
"The Nushagak River system is the fifth largest river in Alaska by volume of
water discharged. The drainage supports at least 13 anadromous species, 16
resident species, and four species offish restricted to estuaries. The Nushagak
River and its tributaries host five species of Pacific salmon and provide significant
habitat for Bristol Bay sockeye salmon - the largest runs in the world. The
Nushagak river hosts the largest sport fishery for Chinook salmon in the United
States, with the third-largest Chinook run in the country. In addition there are
significant numbers of rainbow trout, grayling, Arctic char, Dolly Varden,
northern pike, lake trout, and non-game species (NMWC, 2007:8).
The flora and fauna considered most integral to traditional subsistence use includes the
following. Fish: 1. Sockeye, Chinook, and Coho salmon; 2. Pink and Chum Salmon; 3.
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Whitefish; 4. Winter Freshwater Fish. Mammals: 5. Moose; 6. Caribou. Other: 7. Waterfowl;
and 8. Edible and Medicinal plants. The Elders and other knowledgeable individuals also
identified critical habitat for the species of concern and their harvest locations. The
conservation plan used this information to delineate the watershed into conservation target
areas, in terms of habitat types important for traditional use species (NMWC, 2007:20).
Salmon are the keystone species in the region, and provide enormous amounts of marine
derived nutrients to the ecosystems described above.
In the present study interviewees identified potential threats to the area including
recreation, recreational subdivisions, commercial lodge development, community
development, mining, roads, high seas salmon fishing, oil and gas development, and habitat
shifting and alteration. Interviewees in Pedro Bay during the fall of 2011, for example,
confirmed the high earthquake activity and expressed concerns about new road construction
and its potential impacts on their streams and community, based on their long-term ecological
knowledge.
b. K'ezghlegh: Nondalton Traditional Ecological Knowledge of Freshwater Fish
K'ezghlegh: Nondalton Traditional Ecological Knowledge of Freshwater Fish is based on
interviews with 18 Nondalton residents in 2001 and focused on their current and past
subsistence use of sockeye salmon and other fish in the Lake Clark/Newhalen River drainage
(Stickman et al. 2003: 8). Interview questions related to fishing practices, geographic
locations, and Dena'ina place names. The questions were presented in semi-directed
interviews, with USGS quadrangle maps of the Lake Clark Newhalen River area used to plot
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information. Answers revealed that the summer months, from mid-June through August, are
traditionally devoted to harvesting sockeye salmon that are returning to Sixmile Lake and
Lake Clark. Fish camps used to be set up around the outlet of Kijik Lake, but now are
primarily at the outlet of Sixmile Lake but also along the shores of the Newhalen River,
Sixmile Lake and Lake Clark (Stickman et al., 2003:11).
The interviewees listed nearly a dozen places as the most important locations for sockeye
fishing and eighteen as primary locations for harvesting spawned-out sockeye or "redfish."
Residents described in detail how and where they get salmon, listed 36 separate places where
sockeye spawn, and gave descriptions of several areas where they have noticed reduced
spawning activity, particularly Kijik Lake, which used to be well known as a very productive
area. This area in particular has reduced spawning activity due to beaver dams that seem to be
blocking the entry of the salmon into the Kijik River, and preventing fish from moving
upstream to spawning grounds in and around Kijik Lake. The study also asked about harvest
methods and detailed the use of seines, spears, and fish traps. Seining is no longer allowed
under State of Alaska fishing regulations and fish traps were banned in 1959. People do use
commercially manufactured gill nets.
It was important to the residents that they were respectful of the fish and cared for them.
"Everyone interviewed reported that they generally stop fishing once they have caught the
number offish they need" (Stickman et al., 2003:23). Residents also disapproved of people
leaving their nets out too long unattended. Andrew Balluta, one of the residents interviewed,
said, "They used to say if you don't use what you are catching in your net, don't leave your net
out there" (Stickman et al., 2003:24). The study also elicited descriptions of putting up fish.
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The remaining sections of this report document residents' comments about change:
observed change in salmon over time, observed environment changes, human-induced change;
and finally the importance of salmon to the population as documented in the observance of the
fish camps and the First Salmon Ceremony. A separate section documents the use of other
freshwater fish, including rainbow trout, Dolly Varden, whitefish, grayling, northern pike,
burbot, candlefish, sucker, and lake trout, and their relative abundance. Residents also noted
significant changes in the number offish returning in the five to ten years prior to the 2003
report. "Each person interviewed reported fewer fish than in the past, and all indicated that they
first noticed the change in abundance between five and ten years ago." (Stickman, 2003:26).
While Stickman et al. describe numerous possible reasons for the reductions in numbers, as well
as changes in timing of the runs, the report also noted that flows in the Newhalen River in 2001
exceeded the level observed to prevent or delay sockeye migration into the lower river (Stickman
et al., 2003:27-28 citing C. Woody).
c. Tanaina Plant!ore: Dena'ina K'et'una
Two important TEK studies were conducted largely by Priscilla Russell Kari. The first is a
study of Dena'ina (also known as Tanaina) plant lore describes the seasonal cycle in the
Dena'ina use of plants, as well as detailing the gathering, processing, and preserving of the most
important plants (P. Kari, 1995). She also covers Dena'ina beliefs concerning plants and the
Dena'ina plant classification system. Her study, based on long-term work in several Dena'ina
communities, with a wide range of Dena'ina women, documents more than 150 plants that the
Dena'ina depend on for foods, medicines, and other uses (P. Kari, 1995). The second was done
with George West (Russell and West 2003) and details Dena'ina use of birds.
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E. Social Relations
1. Voices of the People
I feel good, proud [to share]. And when our friends give us back, way proud. M-60, 9/16/11
We share with the Elders first, then with family. Usually how I do it, if someone goes with me we
go 50-50 and he can decide who to share his fish with, and we do the same. It's not decided by
one person, usually me and my wife decide. M-26, 5/19/11
It makes me feel good when I give salmon to somebody. F-47, 8/20/11
It makes you feel good inside because you are sharing. M-53, 8/20/11
It's a good feeling, because we know other people want it. It's a good thing to give away, it's
healthy. F-3Q, 8/17/11
Oh, it makes you really feel good [to give salmon], because I know we enjoy it, and people that
can't get it that were almost raised on it.... That 'sjust the way the whole village is; they share.
F-38, 8/18/11
In our culture here you share with everybody. When I got my first moose, I had to give it to
people; when my grandson got his first moose, you give it to people. You share it. That is one
thing good about the community of Bristol Bay; we still hold on to our cultural values really
strong. Sharing is a very important component to our culture. If somebody is handicapped and
unable to provide for themselves, you find some Young Turk or young family to go help them out.
You don't expect pay. M-60, 9/16/11
You know, I was having a hard time, and her husband [gestures] brought me a whole truckload
of food, and I damn near cried.... Now, when somebody needs help, we do the same thing. If
someone needs help, I try to help as much as I can; we always share. When we give something, it
feels good, and when we are having hard times and get something, it feels good. M-43,8/19/11
[Reference to a woman's] mom was blind, and she couldn 't do certain things, so my mom always
made sure she shared with her. That is one of the things she told me about sharing. She thought
it was good to share with people who couldn't do things for themselves. But, she was always
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doing nice things for us, too. She [the blind woman] made us string to hang fish and things like
that. She was really a nice person, her mom. F-44, 8/19/11
Yeah, we always share. Holidays, we share, and if somebody passes away, after burial we have a
potlatch; we share. We share with people; that is the way we are brought up. F-41, 8/19/1 1
We share with people here and in Anchorage ....I like to go fishing, so if we run out of freezer
space, I will ask people [who can't fish in the village, e.g. Elders] if they want fish, then I'll go
out and catch some fish if they want. M-70, 9/18/1 1
Me, I share it with my younger sisters who never do subsistence. Like, some can 't work anymore.
They [gesture] share it with their parents. Me, I share it with my younger sisters or my son, my
faw. F-23, 5/18/11
Me and my daughter always share after we fish for all summer, but she always tries to give me
lots more, but I say, "No, you 've got more kids. " Sometimes we give [fish to] our daughter-in-
law. F-22
I think, with us, during potlatch times, during hard times or Russian Christmas, or, if we gather
together, everybody brings out their dry fish or their jarred fish or their salt fish. Nobody goes
hungry; there 's always sharing. We would be greedy if we kept it all to ourselves, but there 's
always a sense of sharing with the community or sharing with relatives. F-32, 8/18/1 1
The people up there [Kvichak River villages in the 1990s] were not meeting their subsistence
needs /allegedly due to ADF&G management decisions/. They weren 't screaming about the cost
of gas or the price of lights. They were screaming that they didn 't have fish. There were people
from over here that were shipping fish over there for people to meet their subsistence needs. M-
60,9/16/11
You are a very rich per son if you share. If you don 't share, you are nobody.... I have to go share
food with my grandkids, great grandkids; it doesn 't matter. I don 't care if someone comes in and
eats with us; I like to share. That 's the way we were brought up. Anybody that is in the house,
come and eat with us; you are welcome. F-46, 8/20/1 1
You know, when I was working down in Seattle, my mom used to send me pieces of dried fish all
the time. You know, that mail was slow back then. When I would get it, man, it was just like
candy. No, but one time she sent me mixed berries. You make it with lard; we call it "agutak. "
She sent me those, and by the time it got there, it wasn 't good. Salmon doesn 't spoil when it is
dried M-53, 8/20/11
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We catch moose and caribou and give it away; it ensures good luck back. Even beaver, you give
the whole beaver away after you skin it. After you skin the beaver, you give it away; give the
whole beaver away. That animal that you give away...give[s] you back in return good luck. M-
54, 8/20/11
[My wife] and I have been doing it for thirty some years, doing the fish camp, and putting up
fish for the winter. When the kids were small, we were down therefor them too, and hopefully,
they will have a family, too, and carry on the tradition. M-33, 8/18/11
Some of the salmon we put up at my fish camp even goes all the way down [to] the states. My
friend [name] comes in here, and she puts up fish, and she cans salmon.... [My daughter] and
her friend... they also can fish and dry fish.... [My grandson] was here all summer. F-27, 8/17/11
The parents, their sisters, their aunties, their grandparents, their great grandparents. Everybody
is there [at fish camp], you know, telling them [the children] how to do this....Everybody does it
at their own camps, fish camps.... Everybody is living in different fish camps, so all these
families that are together, that's how they taught the younger kids. F-28, 8/17/11
He [five-year-old grandson] went fishing with us once; now, he went and seined with us. That's
...how we learn, that's how we teach our kids [fish camp]. I mean, it's togetherness. F-30,
8/17/11
One of the things we were taught and we are teaching our kids andgrandkids are that you do not
waste. Boy if they let the fish get rotten boy they would be disappointed in us really bad. So we
teach and pass that on, don't waste nothing. M-29, 8/17/11
We usually get our subsistence foods, salmon, and a wealthy person, years ago, was when he had
a lot of dry fish for his dogs, salt fish, smoke fish. The women had their wooden kegs full of
berries for their Eskimo ice cream. Maybe the father was fishing commercially and made enough
to buy a few groceries form the store, enough [rifle] shells. That was a wealthy person. I think
today a lot of people still think the same way. M-62, 9/16/11
Yeah, I think growing up in a small village wealth was defined by what you provided for your
family. If you were a highline fisher, you were very wealthy, both physically, as well as mentally.
If you were a good hunter, that in itself was very wealthy. Or a good trapper, good provider. M-
61,9/16/11
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Salmon is one thing. They make you feel rich, because you have something to eat all winter.
Smoked salmon, sun-dried spawned-out fish, all of those make you feel good, because you grew
up with it; it is in your body. M-53, 8/20/11
As long as we have a lot offish and meat and stuff, they are wealthy. We don't believe in...
having lots of money. The wealth to us is having more fish put away for the winter, and meat;
that's our wealth. F-27, 8/17/11
In this Western society of living in the city, everybody is for themselves. Everybody is worried
about "Joe Blow " next door, who has a bigger TV or a bigger car; they are worrying about
money, money, money! It just brings on the sickness of worrying. Here, we run a healthy life,
because we have everything we need here; everything we could possibly want is right here. F-32,
8/18/11
They don't learn that at school [proper attitudes toward salmon]. [Laughter]. Elders teach them,
Elders are teachers and pass it down to younger generations. They learn it and pass it down to
their children. Right down to grandchildren, great grandchildren. M-53, 8/20/11
2. Introduction
Though each has a different cultural social organization going back to pre-contact times,
today there are many similarities between the Dena'ina and Yup'ik of the Nushagak and Kvichak
River watersheds. Among them are the importance of sharing subsistence foods, fish camp as a
social and educational as well as economic institution, gender and age equity, and the concept of
wealth.
3. Sharing and Generalized Reciprocity
The Yup'ik and Dena'ina cultures center on belonging to community and on sharing food
as a means of creating and maintaining the living bonds of relationship. The focus on sharing
functions as the elemental ordering factor in sustaining the culture and the long-term health of
the communities. The practice of sharing is elemental in both indigenous and other cultures both
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from a material and a social standpoint (Counihan, 1999:13). Interviewees indicated that the
sharing, preparation, and consumption of food together has created opportunities for efficient and
sometimes ritualized teamwork, as well as social bonding and building of networks. The Yup'ik
and Dena'ina of the Nushagak and Kvichak River watershed villages, as traditional cultures,
continue these practices through harvesting, preserving, and preparing food together and sharing
food through traditional practices and ritual celebrations. They continue to experience the social,
spiritual, and nutritional benefits from sharing food, especially salmon, the staple food, up to the
present.
Sharing remains a fundamental institution within Yup'ik and Dena'ina cultures today,
according to interviewees, and the importance of sharing food, especially salmon, cannot be
overemphasized. Among the Yup'ik, for example, elaqyaq means "those of the same stomach"
and refers both to sharing food and being biologically related. Oscar Kawagley noted a similar
linguistic reference: "The Yupiaq [Yup'ik] term for relatives is associated with the word for
viscera, with connotations of deeply interconnected feelings" (Kawagley, 2006:11). As Langdon
indicates, the time people spent together in subsistence activities is extensive: "The Yupiit
[Yup'ik] enjoyed the bounty of some of the world's richest salmon fisheries. Large quantities of
fish were harvested and processed through relentless hours of work in order to sustain families
and their dogs throughout the long winters" (Langdon, 2002:41).
Yup'ik and Dena'ina sharing is "generalized reciprocity," because the time and place of a
return gift is not specified. In general, interviewees indicated that people do not expect a return
gift when they share salmon or other subsistence foods with someone else, particularly an Elder,
but a return gift of food always seems to appear, whether that month, that year or sometime in
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the future. The altruism is part of social solidarity. Villagers do not consider sharing to be an
obligation, but a way of life, as the Voices of the People at the beginning of this section indicate.
Interviewees universally indicated that giving or receiving salmon or other subsistence foods
makes them feel good. The altruism of sharing food expresses social solidarity between the
participants. Almost universally, Dena'ina and Yup'ik seem to have small jars of salmon
available for visitors to take with them.
Villagers particularly recognize some Elders who cannot participate in the rigors of
subsistence harvesting as people with whom to share salmon and other subsistence foods. The
informal first salmon sharing, for instance, always includes Elders (see Section III.F.5).
Sharing salmon and other subsistence foods with family living in Anchorage or even farther
away is an important bond to home, family, and place. Interviewees consistently talked about
how much they appreciated a gift of canned or jarred salmon from home when they were away
from the village. They also talked about how important it is for them to send a part of the place
to family and friends living away from Bristol Bay.
The Dena'ina believe that tangible items can take on aspects of the owner. This
personification is called beggesha if the aspects are positive and beggesh if negative (Boraas and
Peter 2008: 215-9). Artifacts or places can have beggesha or beggesh depending on events
associated with them. A place, something someone made, such as a birch bark basket, or salmon
someone prepared take on beggesha. The term does not easily translate into English, so today
people talk about giving "love" when giving a gift of something they made or prepared.
Conversely, one receives "love" when receiving a similar gift. This perspective is one of the
reasons that Alaska Native foods, especially salmon, are served at all gatherings such as potlucks
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and potlatches. Preparing and giving food is a tangible act of love. Recipients appreciate non-
Native foods, but they are not from the place, were not made by the giver and, consequently, are
not an expression of love when gifted.
4. Fish Camp
Both the Dena'ina and Yup'ik have a long tradition of going to fish camp to harvest
salmon. As interviewees indicate, the villages of the Nushagak and Kvichak River drainages
harvest salmon either at or very near town, and fish camp may be only a short boat ride or four-
wheeler trip to a traditional fishing locality where they may or may not camp out (cf. Fall et al.
2010). Many villagers, however, still travel to a traditional place, set up camp, and live for
several weeks catching and putting up salmon. Villagers from Kokhanok, for example, travel to
fish camp on Gibraltar Lake, while residents of New Stuyahok, Ekwok, and Koliganek stay at
various camps on the Nushagak River, downstream of the villages primarily at Lewis Point, and
villagers from Nondalton go to camps on Sixmile Lake and Lake Clark. Generally, the
interviewees indicate the fish camp consists of an extended family, with three or more
generations, but close friends may also participate (Fall et al. 2010).
Families typically view fish camp as a good time when they can renew bonds of
togetherness by engaging in the physical work of catching and processing salmon. Family
members who don't live in the villages often schedule vacation time to return home to fish camp,
not just for the salmon, but for family. The importance of sharing in vigorous, meaningful work
cannot be overestimated. It creates cross-generational bonds between children, their parents,
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aunts, uncles, and/or grandparents that, today, are rare in Western culture because there are so
few instances in which meaningful, multi-generational work occurs (Interviews, 2011).
Fish camp is a time when children and teens learn not only the practice of how to
properly catch, clean, and process fish, but the values that are an integral part of harvesting
salmon and interacting with nature. As such, it is a primary educational institution (Fall et al.
2010). Young people learn from their parental generation and, particularly, from their
grandparents, their Elders, about the Yup'ik or Dena'ina way. The primary value passed on at
fish camp is respect for nature and, particularly, respect for salmon. As discussed in Section
III.F.4., showing this respect involves using everything and disposing of what little is left over in
a respectful manner. Fish are not disparaged, bragged about or made fun of. Catching salmon
with a good attitude is the first step in imbuing it with the beggesha or love discussed in the
previous section.
5. Steam Baths
In many villages, informal gender-specific groups meet several times a week for steam
baths in small wooden buildings heated with wood-fired barrel stoves and share stories, the
advice and wisdom of the Elders, and cultural connections. In some ways, these steam baths, or
maqi as the Yup'ik call them, have taken the place of the men's traditional house, qasgiq, and the
women's house, ena, where the transmission of cultural values and knowledge traditionally
occurred, as well as much entertaining talk. Among Dena'ina the traditional word for steambath
is neli which traditionally was a spiritually powerful place as well as a place for healing
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(Kalifornsky 1991:48-50; 218). Today the Dena'ina neli has many of the social aspects of the
Yup'ik magi.
Figure 8. Firewood sled (foreground) and Steambath (background). New Stuyahok,
January, 2012.
6. Gender and Age Equity
Gender equity among subsistence families is balanced and has many of the characteristics
of a traditional family farm or family-run business. Both men's roles and women's roles are
equally valued, and it is common that men can do most "women's" activities (cook, clean fish,
etc.), while women can do most "men's" activities (shoot a moose, run a boat, etc.) (Interviews
2011).
Traditionally, Elders are important members of village society, seen both as sources of
values and storehouses of traditional knowledge, and they are valued in child-rearing, village
decision-making, and life guidance. A common saying in the villages is: "When an Elder dies,
we lose an encyclopedia."
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7. Wealth
When asked their perception of wealth, only 3 of 53 interviewees, all from the same
village, indicated that they measure at least part of their wealth in terms of money, material
items, and potentially high-paying jobs (see Section III.B.8.). The remaining interviewees who
commented responded that wealth is measured in terms of one, or more, of three themes: food in
the freezer, family, and/or freedom.
To the majority of interviewees, stored subsistence food means a family is wealthy or
rich as noted in Section III, B. Various entities attempt to monetize this value, but to the people,
subsistence is priceless. It means you won't starve; it means you will have among the healthiest
diets in the world; it means you will be able to actively engage in the sharing networks described
above; and it means shared, activity that enhances family and/or village togetherness. A full
freezer (or freezers, as is often the case), a well-stocked pantry and a full wood bin are the
primary symbols of wealth in the Nushagak and Kvichak River villages. Most villagers, of
course, recognize that money is a necessity, but money is not the singular measure of wealth.
Money is necessary for the tools for subsistence, gas and oil for boat and house, and occasional
travel, and locals generally acquire it through part-time jobs or commercial fishing that still
allows time for subsistence activities. By Western materialist standards most of the villages are
poor; by their own standards Nushagak and Kvichak River villagers are rich, and it is the people
who live a non-subsistence lifestyle who are poor (summarized from interviews, 2011).
Interviewees indicate that wealth also derives from having a large, extended family,
particularly one that is closely knit by subsistence activities. Having an extended family means
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having people you can count on if need be, and it means having people to whom you can give
your love and assistance. This tradition of alliance through marriage has its origin in pre-contact
Yup'ik and Dena'ina culture (see Sections II.B.3 and II.C.2).
Few interviewees spoke with fondness of living in Anchorage or other urban places they
have lived or visited. Though hunting and fishing require abiding with ADF&G regulations,
most villagers see those activities as involving a degree of freedom that does not often occur in
non-subsistence work settings. As described in many interviews, with subsistence as your job,
you don't have to punch a clock, you only follow nature's clock; you don't have a boss, you are
your own boss, and you either suffer the consequences if you do not perform well or reap the
benefits if you do. During our May visit to one village on the Nushagak River, two young men in
their early twenties left on a 17-day subsistence trip upriver into the Mulchatna area, one of the
most remote places in North America at any time of year, but virtually deserted in spring, when
snow was still present. They were on their own, and apparently all who were connected to the
endeavor embraced that freedom. As they left, for example, the mother of one of the boys simply
said, "Be careful," just as a parent living on Alaska's road system might say to a son embarking
on a trip to Anchorage. This view comes from villagers having knowledge of and ranging over a
vast territory, almost all of which is in a natural state. Consistently, people are thankful to live in
a place where they can live off the land in the manner of their ancestors, and don't want to live
anywhere else (Interviews, 2011).
8. Suicide in the Study Area
Tragically, suicide is one of the primary indicators of individual loss of identity and
breakdown of society (anomie). Alaska has one of the highest suicide rates in the nation and that,
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sadly, is due in part to very high rates in rural Alaska. However, as indicated by data from the
Alaska Bureau of Vital Statistics (see Table 8), those high rates are not spread equally
throughout rural Alaska. In the Northwest Arctic census area the age adjusted suicide rates per
100,000 are four times the Alaska rate (22.7 in 2009) and six times the national rate (11.5 in
2011) (see Table 14). Similarly suicide rates for the Bethel area north of the study area indicate a
similarly grim picture.
Table 14. Suicide Rates in the Study Area compared to Alaska and Other Selected Areas.
Data from "Alaska Bureau of Vital Statistics, Detailed causes of Death in
Alaska, http://www.hss.state.ak.us/dph/bvs/death statistics/Detailed Causes Census/frame.htm
2010
Population
2007-2009
2006-2008
2005-2007
2004-2006
2003-2005
2002-2004
2001-2003
2000-2002
1999-2001
Alaska
698,473
*per
100,000
22.7
22.6
20.9
21.0
21.0
21.5
19.4
19.6
18.3
Dilllngham Census
Area
4,933
*per
100,000
42.4
-
-
-
-
-
-
-
-
Actual
Number
6
2
2
2
4
4
3
1
2
Lake and Peninsula
Census Area
1,488
*per
100,000
-
-
-
-
-
-
-
-
-
Actual
Number
0
0
0
0
0
1
1
3
2
Bethel Census Area
17,236
*per
100,000
61.6
50.1
38.3
48.1
56.9
50.8
32.7
27.6
23.8
Actual
Number
30
25
19
24
29
26
17
13
11
Northwest Arctic
Census Area
7,208
*per
100,000
67.5
93.0
81.9
79.4
66.1
74.8
78.4
74.5
62.2
Actual
Number
15
21
18
18
15
17
17
16
13
* Rate is Age-Adjusted per 100,000 calculated at the 95% confidence interval
~ Rate per 100,000 not calculated because the incidence is too low to be within the 95% confidence interval
The suicide rates for the study area including the Dillingham census area which includes
the Nushagak drainage villages of Dillingham, Ekwok, Koliganek, and New Stuyahok as well as
five other villages outside the study area are comparatively much lower. In only one two year
period was the Age-Adjusted rate per 100,000 even calculable at the 95% confidence level
because the number of suicides was so low (see Table 14). Suicides were even lower for the
Lake and Peninsula Census area which includes the study area villages of Igiugig, Iliamna,
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Kokhanok, Levelock, Newhalen, Nondalton, and Pedro Bay in the Kvichak drainage and 10
other villages outside the study area. While any suicide is a horrible loss for family and
community, especially in small rural villages, statistics indicate suicide is not of epidemic
proportions in the study area it is in other parts of Alaska.
While suicide is complex, one of the chief reasons is a debilitating feeling of hopelessness.
The 2011 Alaska Federation of Natives panel on suicide identified specific factors including
historical trauma, substance abuse, sexual abuse and family violence (DeMarban 2011). It is also
not easy to determine why suicide rates are much lower in some parts of rural Alaska such as the
Nushagak and Kvichak drainage. One reason is that Orthodoxy is generally strong in these
villages and Orthodoxy considers suicide to be a sin and a violation of the fifth commandment
"Thou shall not kill" (Morelli n.d.). Resident priests with close ties to the village no doubt
provide effective remediation, in some cases, to those in despair who might be contemplating
suicide. The cultural strength of a subsistence lifestyle cannot be discounted as a second effective
antidote to suicide. Eating a healthy, natural diet, engaged in vigorous outdoor activity with
family and friends and the village support of those friends and family, and having a measure of
independence and therefore feelings of control of one's destiny, and living in a cultural
continuum that goes back thousands of years on the landscape of one's ancestors no doubt
truncates the despair that can lead to suicide before it ever gets to a critical state.
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F. Spirituality and Beliefs Concerning Water and Salmon
1. Voices of the People
Respect and Thanks
Yes, they do [streams have a spirit], like everything else, all living things. Before Russian
Orthodox came here, that is what we worshipped. We worshipped all the living things, even the
air, the sky, the moon, the sun, snow, rain. It is in every aspect of our lives, how we are made up,
what we believe in, why are we still here? M-33, 8/18/11
They say everything on Earth has a spirit, like we have a spirit. So everything has spirits, the
streams, the waters, the lakes, the mountains, trees, birds; everything has a spirit. To me, I think,
that's why we have to pray, and you have to keep the streams clean, not pollute it. F-27, 8/17/11
/ think that, if you treat animals disrespectful, that they are not going to show up again. F-32,
8/18/11
That is why we are so clean around here... they [outsiders] don't know if we camped around here
or not, because we clean up our garbage, and we hardly leave any evidence that we were there.
M-36, 8-18-2011
Yes, like all other things you are granted [by God], you give thanks for [salmon]. F-69, 9/18/1 Ib.
First Salmon Ceremony
The first salmon, it's still tradition to share with everybody. You do say a prayer. F-47, 8/20/11
When we catch the first king salmon, about this month [May], maybe next week, we share that
king salmon, cut in little pieces, to give to them to cook, especially to the Elders, because they
always want fresh fish. F-22, 5/18/11
First catch is shared with all of the Elders. Elders first, always the priority, Elder, because they
cut it in pieces, you know, if you catch a king, you share, instead of eating the whole fish by
yourself. The first catch. M-20, 5/18/11
Traditionfirst salmon, the very first salmon you catch you boil everything, everything. You
don't waste anything then you eat it too. I mean, even the liver, if it's a male the sperm sac,
everything. M-29, 8/17/11
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Every year, when I first catch a king salmon, I usually pray to God and thank Him for it. A lot of
people do the same thing, because he is the one giving us these wild foods. M-63, 9-18-11
Figure 9. Russian Orthodox Church, Koliganek. September 15, 2011.
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Great Blessing of the Water
There are a lot of folks along the Nushagak, down to Dillingham, and along the chain that are
Orthodox because of the Russian influence. They actually have three ceremonies in the church
that deal with the salmon. The first one is the Blessing of the Water in the winter time. You have
probably seen the newspaper articles about the priest that goes out there and blesses the water.
It can be minus 40 or minus 50 [degrees Fahrenheit], and you seem them running that cross in
the water, and they never freeze. That in itself is a miracle, I think. The other thing that happens
is that, just prior to fishing, the church has a special service of the blessing of all the resources.
The third thing is the blessing of the fishing boats. The individual fishermen, when they get done
with all their nets and all their gear, they can ask the priest to come and bless their boats. M-81,
9/16/11
They do it every year at Theophany.... It's very important to us; it's a blessing of the water,
blessing the river so the fish come in. It's an Orthodox religion ceremony.M-20, 5/18/11
The Holy water is so pure. We believe it is healing, has healing powers. When you are sick or
have a cold, have just a little tiny bit. F-69, 9/18/11
And over on the Iliamna side, they will do the same thing that Father will do over here with the
water, make holy water. People will come down there too with either buckets or jugs and fill
them up. M-65, 9/18/11
I used to live in Portage where there is no clinic. That is the only thing I could give my kids [holy
water, when they were sick]. You know pray upon them and let them make the sign of the cross
and let them have a taste of the holy water. F-72, 9/19/11
That holy water is strong. To be honest with you people, I would not be talking with you right
now [if not for holy water]. A long, long time ago, before I become a lady, we were upriver with
my mom and dad. My mom was sick too, my grandparents and dad, too, and uncle [name]. In
night time, I guess I almost go [die] you know. But my dad, he prayed for me. If you 're really
true, praying really hard, I guess he 'II answer you. My dad tell me I have no more breathing, no
more pulse. And when I come to, my dad was holding me like this, up you know, feeling my
heartbeat. As soon as I opened my eyes my dad said 'you get up'. I said yeah, I told him I was
going to sleep, how come you woke me up? I was going to go to Big Church [heaven], and my
dad said 'you can't go to Big Church' When he tell me that, I told him holy water/ call Native
way, malishok, holy water, malishok [Yup 'ikj 'give me holy water to drink'. He did, my dad, he
did. A little bit you know. I opened my mouth, I swallowed, the water was going down into my
stomach ...I closed my eyes, pretty soon I come through. My dad was up, my momma was
sleeping, she was sick too upriver [Yup 'ikplacename]. I go but I came back. Almost going to
that Big Church. My dad he tell me not to go into the church, come back, that's why I become a
lady. It's true, I tell you guys the truth, better not forget that. Holy water is strong, that is what
made me come back. F-66, 9/18/11
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2. Introduction
Most of the residents of the interior villages of the Bristol Bay drainage are Russian
Orthodox Christians, and the Orthodox Church, along with the public school and the tribal
structure, is among the dominant institutions in the small villages. Many of the villages have a
resident priest or priests; for others, clergy visit periodically on a scheduled basis. In some
villages Protestant churches have formed: Port Alsworth, and Dillingham have Protestant church
buildings, the latter in addition to an Orthodox church.
Beliefs concerning streams and salmon, in those villages where Orthodoxy is the
dominant religion, involve a syncretism merging traditional beliefs with Russian Orthodox
practice. Dena'ina writer Peter Kalifornsky (1991:249) described syncretism when writing about
his great-great-grandfather's nineteenth century message to the Dena'ina people after his
conversion to Orthodoxy: "Keep on respecting the old beliefs, but there is God to be believed in;
that is first of all things on earth." Russian Orthodoxy itself has a syncretic tradition of melding
Middle Eastern-derived Christianity with spirituality influenced by the northern environment.
Billington (1970:18-19, and 403) points out that, though Orthodoxy moved north from Greece and
Asia Minor into Russia in the ninth century A.D., its long history in the northern forest has shaped
the belief system to interpret and interact with aspects of the subarctic taiga. Billington writes, "God
came to man not just through the icons and holy men of the Church but also through the spirit-hosts
of mountains, rivers, and above all, the forests" (Billington 1970: 403). Consequently, many
Russian Orthodox rituals involve interaction with nature. The mystical aspects of Orthodoxy fit
well with traditional Dena'ina and Yup'ik beliefs, many of which related to interacting with the
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landscape on which their survival depended (Boraas, in press). For the Dena'ina and Yup'ik
living in the Nushagak and Kvichak River drainages, beliefs regarding pure water and the return
of the salmon, discussed below, ritually and spiritually express the meaning of life as people of
the salmon.
3. Great Blessing of the Water
The "Great Blessing of Waters" takes place during the Feast of Theophany, a major event in
the Orthodox Church calendar and is celebrated on January 6th of the Julian calendar, the calendar of
Orthodoxy (January 19* in the Gregorian calendar). While all church rituals are important,
Theophany can be considered to be the third most important church ritual after Christmas and
Easter to the Orthodox of the Nushagak and Kvichak watersheds (personal communication, Fr.
Alexi Askoak, St. Sergis Russian Orthodox Church, New Stuyahok, January 19th, 2012). A
theophany is an event in which God reveals himself to humans and the Great Blessing of the Water
marks the baptism of Jesus by John the Baptist. After Jesus' baptism God appears saying, "this is
my son whom I love, with him I am well pleased," (Matthew 3:17, New International Bible). As
explained by Fr. Alexi Askoak (personal communication, January 19, 2012), in the Orthodox view,
baptism both redeems sin and brings the Holy Spirit to the recipient. Orthodoxy believes in the
triune God, consequently Jesus is God and without sin. So Orthodoxy transfers the ceremony to one
of God's most important creations, water, and one of the creations most important to the people of
the Nushagak and Kvichak since salmon and related wild foods are dependent on clean water. An
evening church service is held on the eve of Theophany in preparation for the blessing the next day.
The two-day ritual is a liminal event with believers moving into a deeply spiritual mental state. At
the service I (Alan Boraas) attended, 211 villagers of New Stuyahok were present filling the small
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church in New Stuyahok. The next morning a communion service was held and, as the sun rose, the
people led by the priests went out onto the frozen Nushagak River where an Orthodox cross had
been cut into the ice and a small hole had been made to withdraw holy water (Figure 10). There a
baptism service was held purifying and sanctifying the water of the Nushagak River. At the moment
in the service when the priest dips the cross through the hole in the ice into the water for the third
time, God is believed to sanctify the water making it holy. According to Father Michael Oleksa the
Great Blessing of the Water is done to "reaffirm the Church's belief that the natural world is sacred
and needs to be treated with care and reverence" (Orthodox Church in America, n.d.). The Orthodox
Saint John Maximovitch (n.d.) wrote:
... when we bless waters of lakes, rivers and streams, we ask God to send His blessings
upon the waters of His creation so that even though humanity has spoiled the world
through sin and abused the environment over many generations, God has not forsaken the
world. He sends His spirit to cleanse and sanctify His creation.
"Sin" in the form of human-caused pollution and other contaminants are ritually removed from the
water and it is now considered pure and holy (personal communication, Fr. Alexi Askoak, January
19, 2012). In New Stuyahok, and other villages where the ceremony is performed, the now blessed
water is removed in containers for personal spiritual use and a large container is taken back to the
church for use as holy water.
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Figure 10. Great Blessing of the Water, Fr. Alexi Askoak, St. Sergis Church, New Stuyahok.
January 19,2012.
Holy water from the sanctified rivers is believed to have curative powers for both physical
and mental illness and is drunk or put on the affected part (Fr. Alexi Askoak, personal
communication, January 19, 2012). Several interviewees shared very personal incidents of the
power of holy water to cure. Fr. Alexi told the story of one bitterly cold Theophany when he frosted
his face during the ceremony. When they returned to the church one of the parishioners rubbed holy
water on his face and he subsequently did not blister or suffer any ill effects other than one little spot
the water had missed which left a mark for several years. Fr. Alexi believes God healed him through
the holy water. A young interviewee in Koliganek movingly told of a time when her children were
gravely ill and there was no doctor, health worker, or suitable medicine available. She said, "all I
had was holy water." She had the children drink the holy water and in a few days they recovered.
She attributes their recovery to the power of the blessed water. An elderly woman movingly told the
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story of being brought back from near death when she was a child by holy water. Both stories are
recounted in the "Voices of the People" at the beginning of this section.
From a secular standpoint, the question is not whether or not holy water has healing
efficacy, but how the Great Blessing of the Water ceremony and holy water reflect values of the
people. People elevate to the sacred those things that are most meaningful or critical in their lives.
As described in section II. E. 4 the Dena'ina word for water, vinlni, has sacred overtones and water,
itself, is sacred. Since the word predates Christianity in south-central Alaska, we can assume sacred
water has long been a part of the salmon cultures of the Nushagak and Kvichak watersheds because
clean water and salmon are fundamental to life itself. The Great Blessing of the Water ceremony is
an obvious extension of that very old concept, rendering in Christianity that water is sacred to life.
The antiquity of the Great Blessing of the Water in Alaska is apparently as old as
Orthodoxy. Hegumen Nikolai was an Orthodox missionary priest briefly stationed in the Nushagak
area in 1846 and then transferred to be the first permanent priest in Kenai where he served from
1846 to 1867 (Znamenski 2003:15-18). In his travel journals Hegumen Nikolai describes
conducting the Great Blessing of the Water in Kenai in 1862 and 1863 on January 6*, Julian
calendar. (Znamenski 2003: 94, 108) (Travel journals, official church documents missionary priests
were required to submit to the diocese yearly, have not been translated for earlier years for
missionary priests operating in the Dena'ina or Yup'ik areas of the Nushagak and Kvichak
watersheds.)
4. Respect and Thanks
Water and salmon play additional roles in modern Orthodoxy in the study area as derived, in
part, from traditional subarctic spiritual practices. Describing traditional Dena'ina beliefs,
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Kalifornsky (who was also a devout Orthodox Christian) writes (1991:362-363) that, after
putting out his net, "^uq'a shegh dighelagh" or "a fish swam to me," indicating that the spirit of
the salmon had a will and would allow itself to be taken for food if the net-tender had the correct
attitude. Today, all interviewees that commented on it still believe that salmon have a spirit or
soul and that soul is a creation of God. Further, all interviewees who responded report offering a
prayer of thanks when they catch salmon, particularly the first salmon as noted in the "Voices of
the People" at the beginning of this section. That prayer may be a humble "in one's mind"
statement or it may be spoken thanking God for the salmon.
Interviewees also still believe in treating all animals, including salmon, with respect.
Several modern practices reflect this belief, for example, using the entirety of a fish for food,
except the entrails, which villagers return to the water along with the bones that remain after
consumption. Another example, interviewees report, is never allowing fish or meat to spoil.
Interviewees repeatedly stressed the importance of giving salmon and all subsistence animals
respect. This attitude echoes the pre-contact beliefs that animals had a will and, if not treated
properly, would not allow themselves to be taken for food, leading to dire consequences for the
people (Boraas and Peter 1996:190-192).
5. First Salmon Ceremony
The First Salmon Ceremony is a world renewal ceremony which, like other world
renewal ceremonies, recognizes the cyclical onset of the most important yearly event in the
culture. As mentioned in Section II.C.2, the First Salmon Ceremony was described by
ethnographer Cornelius Osgood (1976:148-9) and was practiced in pre-contact times and is
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based on a mythical story that merges people and salmon. Because of the importance of salmon
in the lives of the Bristol Bay villagers, interviewees report they continue to mark the return of
salmon in the spring by a special observance. The actual practice varies, but involves a prayer of
thanks to God for the return of the salmon and sharing the first salmon caught in the spring with
Elders and others in the community. Typically, according to interviews, each receives a small
piece, and there is a general feeling of happiness that the salmon have returned and the cycle of
the seasons has begun again and nature will provide the people with sustenance. In some places
the First Salmon Ceremony takes place at fish camp, where extended families and others present
share the first salmon they catch with one another, including the Elders. In at least one village,
New Stuyahok, the ceremony includes sharing the first salmon with "the underground," by
placing a small piece of it under the forest mat at the cemetery, symbolically sharing salmon with
the deceased ancestors buried there.
Figure 11. Kvichak River and Lake Iliamna at Igiugig. May 16, 2011.
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G. Messages From the People
At the conclusion of the interviews we asked interviewees if there was anything else they
wanted to say, anything we had not covered, and/or any message they wanted the Environmental
Protections Agency to hear. The following reflect those comments:
1. Voices of the People
/, myself, get very emotional when the topic of the Pebble Mine comes up. I don't even want to
think about it. In the future I don't want to think about total ruin of our way of life. It really
saddens me. F-69, 9/18/11
For quite a few years there when we were building up the king salmon run we didn 't even fish in
June. It was just to buildup those runs. It is kind of ironic that the kings we built up are on the
Koktuli River where that mine is going to go. It is almost a whole decade that we sacrificed to
build up that run. We built it up and now it might go away. M-61, 9/16/11
You don't see Bristol Bay having troubles because our ecosystem is whole and not damaged. We
are very appreciative of what we have. In relationship to the mine the place I work up here is the
Bristol Bay Economic Development Corporation and... one of the companies we bought is
Ocean Beauty Seafoods which is one of the largest salmon producers in Alaska. We put up
161 million pounds of commercially caught goods in a year. Sol talk to the people and if there is
a mine that goes in like pebble and we have copper coming out and affecting our fish, are you
interested in buying our fish? These are customers we sell 300-400 thousand pound lots to. No,
we are not interested.... We don't want ourselves and our kids to eat contaminated foods. M-60,
9/16/11
It is clear, good water to drink. This is what we protect our good water to drink. F-48, 8/20/11
We can't even fathom somebody hurting the salmon. When the pebble mine folks first came in
they said they were going to pump the tailings right into the middle of the lake. We said you are
going to kill the lake. They said you guys got no say so....We said no you 'II kill the lake. We
couldn 't fathom it. We said you kill the lake and we will go to war. M-60, 9/16/11
Since the Pebble Mine started their exploration, I speak for everyone around here that we have
not had the big caribou herds that come through here anymore. F-69, 9/18/11
That is our greatest fear about the mine. The size of the hole and the tailing pond they are going
to build. You know you see our KDLG water tower up here and the size of the walls are going to
be greater than that and if we get a spill we are done. What we say is that we can't afford the
risk. The mine might be safe but there might be an earthquake and pollution happens. We can't
afford the risk. M-60, 9/16/11
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In Easter they went up to Koliganek the next village up. He said people up there caught white
fish and pikes. He said the water is good upriver, it's not like down here. I think it's the water
that is coming down from upMulchatna. He thinks it's from them working on that pebble up
there [pebble mine]. F-23, 5/18/11
There's open water all over. They got drilling rigs that are sitting on open water. You can't walk
up there with knee boots you got to have hip boots there is so much water this year. The ground
is saturated. M-60, 9/16/11
[Translator of 80+ year old Yup'ik-only speaking Elder] He is only worried about the Pebble,
right now. If the Pebble starts, the water is going to get effected before anything else. That's
what he is worried about.
M-21, 5/18/11 We feel that EPA is very important around here to give us a fair shot at examining
this.... [reference to specific individuals deleted] You know they [state officials] are all for this
economic development. You know economic development up in that mine they are going to bring
in outsiders they are going to destroy the culture up there like you wouldn 't believe. Most of the
outsiders will, most of the jobs will go to outsiders and we will be left with the pollution. M-60,
9/16/11
They [Salmon] would not go there [where water is contaminated] They are also very sensitive to
temperature. They have a really keen sensory acuity, not only them, but all the critters, all the
birds. ... They are so sensitive in every aspect of that word. ...It's relying on the renewable
resources for our people have been going on for a long time. The respect for it, it is still therefor
those of us who do respect it. We have been sharing it with everybody. Nobody was jumping up
and down, hollering about one group or another, until the Pebble people came. We took all
these resources just for granted. We did not know anything about open pit mine or mining. I
realize as human beings we need mines. I have to buy bullets now and then. I have to buy a prop
for my outboard motor. I have to go buy bearings for my Honda. This is not a place to have that.
They cannot have that here. There is no balance there. They talk about coexistence, that is
not... that's coming from the other side. That stuff can't coexist with salmon. Are you going to
compare coal to copper? Copper is a thousand times more devastating that coal. [M-33,
8/18/11
The drill wells are making all the noise. We were over there, my wife and I were over there last
spring, and when we went over there to check out the Pebble, there /we/ saw three other
helicopters right in the same area, and that's lots of traffic. We have not had caribou meat
around here ever since. Haven't had caribou meat caught here in probably the last six years.
M-68, 9/18/11
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Bristol Bay is renowned for what it has to offer. Like I was saying earlier, this region had a very
good working agenda before the Pebble people came. M-33, 8/18/11
[Name] went with her and she is about 88 years old [mother and daughter on an Outside mine
visit]. They went out to look at mines and [name] cried at every mine she looked at, she couldn 't
believe that man would be that disrespectful of the earth. She said literally cried... like her
brother, mom or dad died. She represents us all, we can't see destroying the earth like that.
We 're not greenies you know we are far from green but we can you know. Without EPA we are
sunk. ...We know it is just a matter of time. All of us have had a few cocktails and drove, one of
these times we are going to have a few cocktails and get in a car wreck. It is just a matter of
time. Just like that mine. We really feel helpless with the state government. It is like we are
dispensable out here and it is better for the big boys to come in. that is what the mine people are
telling us. Right guys? When they first started coming? You got no say, so we are coming. M-60,
9/16/11
And what is going to happen when this mine closes up? Our great-great-great grandchildren are
going to end up paying for it. If they are fortunate enough to still be living in Bristol Bay if the
salmon, the streams are not contaminated and sustained. I hate to think of the future if this mine
goes through. The long haul it is going to be devastating. M-62, 9/16/11
We are very rich. With this new mine coming up, I would never trade my fish for money or a
new house, or whatever. I'd like to have all that, but I would not trade what we have every year
for how many centuries. F-35, 8/18/2011
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IV. CONCLUSIONS
As described in Sections II and III, the Yup'ik and Dena'ina cultures of the Nushagak
and Kvichak River watersheds practice a subsistence lifestyle that developed over several
thousand years of living in the area and depends primarily on salmon. At the same time the
people have incorporated modern technology, political participation and educational standards
into a successful transition into the modern world. As illustrated by the Elder and culture-bearer
interviews, this lifestyle has built strong, connected networks of extended families and a culture
based on sharing, traditional knowledge, and respect for the environment.
Most of the villages have schools (except Pedro Bay where children are home schooled),
city government or tribal council, a health clinic, post office, small store, church, airstrip, and
electricity and running water in most homes. Homes have radio and satellite TV and many are
being connected to high-speed fiber-optic internet. Basketball games in the school gym and
bingo at the council building, and sometimes Yup'ik and Dena'ina dancing, and communal
sweatbaths are popular in the evenings. Four-stroke outboards on large skiffs, four wheelers, and
snow-machines are everywhere. These changes are recent, however; up until about sixty years
ago, traditional dog sleds and kayaks provided the transportation, and caring for dog teams took
much time and effort. The availability of material goods from beyond the villages was limited,
modern housing was nonexistent and formal education was mainly offered through boarding
schools. The villages of the study area grew dramatically between 1980 and 2000, probably due
to post-ANCSA changes in land-ownership (Fienup-Riordan 1994:39) and the population is now
holding steady although there is local village variability.
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These changes have resulted in some loss of traditional cultural practices; for instance,
people no longer openly practice the Bladder Festival, Kelek or Petugtaq, although essential
elements of these can be found in more informal practices, and in some cases transformed
through corollary rituals in the churches (see Section III.F). Other changes have been more
severe and have both made the communities more vulnerable to changes in their environment
and placed them at higher risk for further cultural and individual losses. Examples of such
changes include loss of control over traditional use areas, loss of community members to
Western diseases and outmigration of young people, for either employment or education, the
latter of which included, in the past, the involuntary placement of children in distant boarding
schools, removed from the traditional culture (Interviews, 2011).
Some interviewees expressed a fear of the future that a traditional prophecy of "bad
times" told by Elders might be coming true due to economic development resulting in cultural
loss characterized as "anomie," the loss of meaningfulness, sense of belonging, and direction in
life. The cultural and social impacts associated with Westernization have been described as
anomie. Merton (1938: 682) gave a classic definition of anomie where he writes, "At the
extreme, predictability virtually disappears and what may be properly termed cultural chaos or
anomie intervenes." Anomie, the loss of meaningfulness, sense of belonging, and direction in life
has occurred among all Alaskan Native cultures to one degree or another. Anomie increases
cultural and individual risk for social ills such as depression and suicide, alcoholism and drug
abuse, domestic violence, and aggressive behaviors. Healing practices can include those used for
trauma and post-traumatic stress disorders, including traditional practices that reconnect the
individual to society and the natural environment through meditative rituals. Traditional
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drumming, singing, and dancing have been shown to be effective in treating trauma and post-
traumatic stress. Culture camps and other methods of cultural revitalization (see Section III.E.4,
5) can be both preventative and healing for children and adults of indigenous cultures. It is
critical to assess future risks and vulnerability, and take appropriate measures to reduce both.
Despite colonial disruptions to indigenous peoples in Alaska, the underlying cultures
have so far endured among the Yup'ik and Dena'ina people of the study area because of a strong
subsistence base. Wholesale changes to the ecosystem that supports their subsistence resources,
however, whether they come from large-scale development, including mine development,
climate change, high-seas overfishing, and/or declines in the ecological integrity of the North
Pacific Ocean such as acidification, carry with them the risk of substantially altering the
subsistence lifestyle and the fabric of Yup'ik and Dena'ina cultures. Among the specific
potential risks associated with diminishment in either the quantity or quality of subsistence, and
especially salmon, resources are:
Degradation of nutrition and physical health due to diminishment of subsistence foods
and lifestyle.
Loss of political power due to becoming a minority in one's own homeland, if there is an
influx of outsiders to the region due to extractive resource development.
Deterioration in mental and emotional health due to the loss of traditional culture and
meaning for life.
Loss of language and traditional ways to express relationships to the land, one another,
and spiritual concepts.
Loss of meaningful work by extended families operating together as a cohesive unit.
Reduction of gender equity resulting from loss of important economic activities and
social networking opportunities, due to the potential diminishment of subsistence foods
harvest and preparation, and replacement of this work with jobs that are typically more
accessible to men (e.g. mining) or to fewer women (such as those who do not have small
children).
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Loss of the means to establish and maintain strong social networks though sharing of
subsistence foods.
Impact on belief systems that revere clean water and a clean environment.
Increased discord within and among villages between the majority and the minority over
development issues within the villages has the potential to create long term rifts within
the villages and between them.
In summary, salmon and clean water are foundational to the Yup'ik and Dena'ina
cultures in the Nushagak and Kvichak watersheds. The people in this region not only
rely on salmon for a large proportion of their highly nutritional food resources; salmon is
also integral to the language, spirituality, and social relationships of the culture. Because
of this interconnection, the cultural viability, as well as the health and welfare of the local
population, are extremely vulnerable to a loss either quality or quantity of salmon
resources
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V. APPENDIX 1. METHODOLOGY and TRIBAL LETTER OF INTRODUCTION
April 11,2011,
Revised April 25, 2011,
Revised May 24, 2011
Purpose:
The purpose of this qualitative study is to describe the subsistence, nutritional, social,
linguistic, and spiritual importance of salmon to the Yup'ik and Dena'ina of the Nushagak and
Kvichak River drainages of Bristol Bay. This information will be integrated into a larger study,
called the Bristol Bay Assessment, coordinated by the Environmental Protection Agency to be
used to determine to proceed with a Section 404c review of the Clean Water Act. This action was
requested by nine tribes/villages of the Bristol Bay region. If approved, 404c designation would
prohibit any discharge into, fill, or similar modification of a stream or river in the region or other
actions that would impact the subsistence fishery.
Design:
The product of this study consists of two parts.
A. Summary of existing research: One part of this assessment consists of a literature and
gray literature search and summary of the culture history, linguistic, subsistence and
other aspects of cultural lives of the traditional and cultural lives of the Nushagak and
Kvichak drainage people as it relates to streams and fishery subsistence, particularly
salmon
B. Elder and Culture Bearer Interviews: Second, this study will incorporate Elder and
culture bearer interviews to ascertain the importance of salmon and other stream-
related resources and places in the ideal culture of the people. Ideal culture is a
standard to aspire to and thus is a measure of values and ideology that form the core
of the people's contemporary identity. We are not undertaking a statistical sample of
attitudes reflecting everyone in the culture, but listening to culture bearers who have
the status of expert witnesses and act as spokespeople for their respective cultures.
The remainder of this methodology will describe the Elder and culture bearer interviews.
Selected Villages
Both time and money prohibit interviews in all villages in the region. Since this is not a
statistical study, nor a hearing, we believe that a self-selected group of Elders and culture bearers
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can best represent the perspective of the region. We intend to interview Elders and culture
bearers from six villages.
Semi-Structured Questions:
The interview format will be semi-structured, meaning the same questions will be asked
of each of the Elder/culture bearers. The questions are intentionally open-ended and intended to
elicit narrative responses. If an Elder/culture bearers wishes to provide additional information or
talk about subjects beyond the scope of the question, that, of course, will be recorded.
Interview Questions
Draft Interview questions will be formulated in the following categories:
Subsistence
Nutrition
Language and Stories
Place names and Special/Spiritual places
Social Factors
Spirituality related to streams and fishery
The draft interview questions will be distributed for review by
Village councils or similar authority
E.P.A. personnel
Selected anthropologists
and reformulated and condensed as needed.
Self-Selection
Village councils, traditional councils, or similar entity will be asked to select
Elders/culture bearers to be interviewed. We anticipate this will involve about three men and
three women in each village.
Release
Interviewees will be asked to sign a consent form allowing the interviewers to use the
recorded and transcribed interviews in a written document. In addition the village councils will
be asked to sign a release form for the village to permit photographs and video both of
individuals or the village to be taken and potentially used in the final product. Restrictions will
be respectively adhered to.
Recording and Transcription
Interviews will be recorded either individually or in small groups. A digital recording and
transcription will be made. Elders may wish to speak in Yup'ik in which case we ask a translator
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provide a summary at the time of the interview. Elders and culture bearers will be paid according
to current standards for village/Elder interviews. The interviews will be approximately two-hours
and conducted at a comfortable place.
The interviews will be transcribed into MS Word documents and both the recording and
transcription be archived either at the National Park Service Alaska or suitable repository. Copies
of recordings and transcriptions will be sent to tribal councils.
Coding
Word document interviews will be coded. Key words will be set up for use in identifying
the subject of the paragraph of the transcribed recording. For example, through sophisticated
searches everyone who responded to or used the term "sharing salmon" will be electronically
listed and some or all of these responses either quoted or paraphrased in the final document.
Confidentiality
According to Institutional Review Board standards, names of interviewees will not be
revealed in the final document. Each interviewee will be asked to sing a consent form that
includes the voluntary nature of the interview, confidentiality, and that there is no known or
perceived risk in granting the interview.
Peer Review
Both drafts and a final document will undergo peer review. For the purpose of this study
anthropologists, EPA reviewers, other scholars, and Village Elders or Culture Bearers are peers.
Community Review
The final draft will be sent to communities who have participated in this study for their review.
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KENAITZE
INDIAN
TRIBE
March 1, 2011
To Whom It May Concern:
The purpose of this letter is to formally introduce our friend and honorary Kenaitze Tribal
member. Dr. Alan Boraas. Dr. Boraas has worked with and on behalf of our Tribe for over
30 years. We have found him to be ethical, fair, and responsive to our requests for
confidentiality. He respects our Dena'ina culture, traditions, and values, and lives them.
Dr. Boraas asked for this letter of introduction in observance of tribal protocol and
because he values and respects our rights to sovereignty and self determination. We
have no doubt that you will find him to be a man of integrity who shares our love for our
waters and lands.
Please feel free to contact me if you have any questions or concerns.
§
Sincerely,
Jaylerve fferefrson Nyren
V-/ V
Executive Director
Kenaitze Indian Tribe
o
==
d
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