STATE OF WASHINGTON    

Status Report for the

Tufted Puffin

Thor Hanson and Gary J. Wiles

January 2015

 The Washington Department of Fish and Wildlife maintains a list of endangered, threatened, and sensitive
species (Washington Administrative Codes 232-12-014 and 232-12-011, Appendix B). In 1990, the
Washington Wildlife Commission adopted listing procedures developed by a group of citizens, interest
groups, and state and federal agencies (Washington Administrative Code 232-12-297, Appendix B). The
procedures include how species listings will be initiated, criteria for listing and delisting, public review
standards, the development of recovery or management plans, and the periodic review of of listed species.
The first step in the process is to develop a preliminary species status report. The report includes a review of
information relevant to the species’ status in Washington and addresses factors affecting its status. The
procedures then provide for a 90-day public review opportunity for interested parties to submit new scientific
data relevant to the draft status report and classification recommendation. At the close of the comment
period, the Department incorporates new information and prepares the final status report and listing
recommendation for presentation to the Washington Fish and Wildlife Commission. The final report and
recommendations are then released for public review 30 days prior to the Commission presentation.
The draft status report for Tufted Puffins was reviewed by researchers and state and federal agencies. This
was followed by a 90-day public comment period from September 12–December 11, 2014. All comments
received were considered during the preparation of the final status report. The Department intends to
present the results of this periodic status review to the Fish and Wildlife Commission for action at the
February 2015 meeting.

This report should be cited as:
Hanson, T. and G. J. Wiles. 2015. Washington state status report for the Tufted Puffin. Washington
Department of Fish and Wildlife, Olympia, Washington. 66 pp.

Cover photos by Peter Hodum showing an adult Tufted Puffin and seabird burrows at Destruction Island, Washington
Inside cover photo © Charles Bergman; back cover photo © Bill Hoglund

 Washington State Status Report for the Tufted Puffin

Tufted Puffin at Tatoosh Island, Washington (Photo © Charles Bergman)

Thor Hanson
The SeaDoc Society, 942 Deer Harbor Road, Eastsound, WA 98245
Gary J. Wiles
Washington Department of Fish and Wildlife, 600 Capitol Way N, Olympia, WA 98501-1091

Washington Department of Fish and Wildlife
Wildlife Program, Diversity Division
600 Capitol Way N
Olympia, WA 98501-1091
January 2015

 TABLE OF CONTENTS
ACKNOWLEDGMENTS .............................................................................................................................................. iv
EXECUTIVE SUMMARY .............................................................................................................................................. v
INTRODUCTION ............................................................................................................................................................ 1
TAXONOMY ..................................................................................................................................................................... 2
DESCRIPTION ................................................................................................................................................................. 2
GEOGRAPHICAL DISTRIBUTION .......................................................................................................................... 3
World ................................................................................................................................................................................ 3
North America ................................................................................................................................................................ 3
Washington ...................................................................................................................................................................... 3
NATURAL HISTORY...................................................................................................................................................... 4
Daily Habits ..................................................................................................................................................................... 4
Diet and Foraging ........................................................................................................................................................... 5
Reproduction, Longevity, and Survival ....................................................................................................................... 6
Migration and Dispersal................................................................................................................................................. 8
Molting ............................................................................................................................................................................. 8
Ecological Relationships ................................................................................................................................................ 9
Diseases and Parasites .................................................................................................................................................. 10
HABITAT REQUIREMENTS ..................................................................................................................................... 10
POPULATION STATUS AND TRENDS ................................................................................................................ 11
Global Status and Trends ............................................................................................................................................ 11
Washington Status and Trends ................................................................................................................................... 13
HABITAT STATUS ........................................................................................................................................................ 17
Past .................................................................................................................................................................................. 17
Present and Future ....................................................................................................................................................... 17
CONSERVATION STATUS ........................................................................................................................................ 18
RESEARCH, MONITORING, AND RESTORATION ........................................................................................ 19
FACTORS AFFECTING CONTINUED EXISTENCE ....................................................................................... 20
Adequacy of Existing Regulatory Mechanisms........................................................................................................ 21
Reduced Prey Availability ............................................................................................................................................ 21
Climate Change Effects ............................................................................................................................................... 24
Environmental Contaminants..................................................................................................................................... 26
Introduced Species ....................................................................................................................................................... 28
Commercial and Tribal Fishing Bycatch ................................................................................................................... 29
Human Disturbance ..................................................................................................................................................... 30
Bald Eagle Predation .................................................................................................................................................... 31
Other Factors ................................................................................................................................................................ 31
CONCLUSIONS AND RECOMMENDATION .................................................................................................... 32
LITERATURE CITED................................................................................................................................................... 34
PERSONAL COMMUNICATIONS AND SOURCES OF UNPUBLISHED DATA .................................... 46
Appendix A. Counts of Tufted Puffins at breeding sites and other locations in Washington, 1886-2013. ...... 47
Appendix B. Washington Administrative Codes 232-12-011, 232-12-014, and 232-12-297 and Revised
Codes of Washington 77.15.120 and 77.15.130. ...................................................................................................... 57
Appendix C. WDFW responses to public comments received during two public review periods .................... 63

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 LISTS OF TABLES AND FIGURES
TABLES
Table 1. Name, location, and occupancy status of documented Tufted Puffin breeding sites in
Washington……...…………………………………………………………………………………. 4
Table 2. Population trends of Tufted Puffins in North America …………….……………………………. 12
Table 3. Changes in the numbers and sizes of breeding colonies of Tufted Puffins in Washington
from the late 1800s to 2007-2010….………………………………………………………………. 15
FIGURES
Figure 1. Hourly attendance patterns of Tufted Puffins at Tatoosh Island, Washington, from
June 1 to August 24, 2005–2008, averaged across years ...…..……………………………………… 5
Figure 2. Population trends (abundance or density) for Tufted Puffins in Washington from four time
series datasets: Westport offshore surveys, 1972-2001; coastal offshore surveys from Cape Flattery
to Point Grenville, 2001-2012; Tatoosh Island colony counts, 1983-2001; and Protection Island
colony counts, 1973-2001…………………………………………………………………………. 15
Figure 3. Locations and historical maximum population estimates for 44 Tufted Puffin breeding colonies
documented in Washington. Panels A-C depict colony occurrences along the outer coast from
northwestern Clallam County to northern Grays Harbor County during the periods from
1886-1977, 1978-1984, and 2007-2010. Panels D-F depict colony occurrences in the San Juan
Islands and eastern Strait of Juan de Fuca during the periods from 1886-1977, 1978-1984, and
2007-2010………………………………………………………………………………………… 16

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 ACKNOWLEDGMENTS
Much of the preparation of this report was funded by the SeaDoc Society, a program of the Wildlife Health
Center, UC Davis School of Veterinary Medicine (www.seadocsociety.org). This work would not have been
possible without financial support from private citizens, especially K. and R. McDowell, G. Georges, E.
Snyder, R. Lundeen, K. and G. Keeler, A. Azous, R. and P. Henigson, and K. Allen. J. Gaydos provided
important organizational support.
Internal and peer review comments were kindly provided by S. Pearson, D. Lowry, B. Tweit, H. Allen, P.
Becker, B. Hoenes, D. Stinson, J. Gaydos, D. Noviello, C. Niles, L. Wargo, A. Kettle, P. Hodum, D. Bertram,
L. Antrim, S. Thomas, S. Horton, S. Gremel, L. Slater, S. Stephensen, and L. Leschner. S. North offered
substantive comments during the public review period. We also thank the many biologists and other
professionals who contributed background information on various topics, including S. Pearson, S. Thomas,
D. Stinson, I. Vilchis, A. Kettle, L. Slater, P. Hodum, J. Parrish, K. Fresh, P. Arcese, K. Stick, M. Hipfner, T.
Scruton, M. Rodway, K. Koski, R. Lowe, C. Niles, L. Slater, S. Stephensen, S. Mills, J. Dolliver, L. Wargo, S.
Thiesfeld, H. Reed, C. Burns, K. Henry, and others. J. Jenkerson maintained the WSDM database for Tufted
Puffins at WDFW and created the breeding colony maps used in Figure 3. N. Alboucq helped significantly
with the literature review.

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 EXECUTIVE SUMMARY
With its striking plumage and brilliant orange bill, the Tufted Puffin (Fratercula cirrhata) is an iconic seabird
well known to native peoples, fishermen, and coastal communities throughout its range in the temperate and
sub-arctic North Pacific. Though pelagic in winter, puffins gather on islands and headlands during spring and
summer to breed and raise their young. They are members of the auk family, with stocky bodies adapted to
“flying” underwater as they dive in pursuit of a wide range of fish and invertebrate prey. Nesting Tufted
Puffins range up to 100 km from their breeding colonies to forage for their nestlings, and are famed for
carrying 20 or more small fish at one time, neatly lined up and carried crosswise in their large, brightly colored
bills.
Historically, 44 Tufted Puffin nesting colonies were documented in Washington and the bird was considered
common in the San Juan Islands, the Strait of Juan de Fuca, and particularly along the outer coast of the
Olympic Peninsula. The population along the outer coast was conservatively estimated at about 25,000
individuals in the early 1900s, and the statewide population remained in that range for much of the 20th
century, with a 1978-1984 minimum estimate of 23,342 birds at 35 known sites. Recent surveys, however,
found nesting birds at only 19 sites in 2007-2010 and resulted in a minimum outer coastal population estimate
of 2,958 individuals in 2009. Maximum estimates once ranked nine colonies at 1,000 or more individuals
each, with two reaching 10,000 or more, but no current breeding sites now hold more than a few hundred
birds. Significant average annual rates of decline have been recorded on pelagic surveys west of Westport
(13.6%, from 1972 to 2001) and coastal surveys from Cape Flattery to Point Grenville (8.9%, from 2001 to
2012). Taken together these studies strongly suggest that Tufted Puffins in Washington have undergone an
order of magnitude population decline, which is ongoing, and a decrease in the number of occupied breeding
colonies of 57% since 1886-1977 and 46% since 1978-1984. This declining trend corresponds with a broader
geographic pattern of range contraction, population decline, and breeding colony disappearance noted
throughout the southern portion of the species’ distribution, including California, Oregon, and Japan.
Causes for the decline are unknown, but potentially include a number of historical and recent factors such as
reduced prey availability, changing oceanic and climatic conditions, entrapment in fishing nets, mortality from
oil spills and chemical contaminants, human disturbance of breeding colonies, impacts from introduced
species, and increased Bald Eagle predation. The largest known mortality event in Washington was the 1991
Tenyo Maru oil spill, which killed an estimated 9% of the state’s Tufted Puffin population. Rising ocean
temperatures and other shifts associated with climate change are expected to reduce ocean productivity and
forage fish populations throughout the North Pacific, creating a challenging environment for piscivorous
birds like the Tufted Puffin.
The current conservation status and legal protections for Tufted Puffins include no specific management or
recovery requirements for Washington’s population. Puffins in Washington are considered a species of
concern by the U.S. Fish and Wildlife Service and have been petitioned for listing under the federal
Endangered Species Act, but an evaluation to list the species will not begin until 2016 or 2017. At the state
level, Tufted Puffins have been a candidate for listing since 1998 and have been identified by Audubon
Washington as a species at high risk from the impacts of climate change.
Given the rate of recent population decreases, widespread colony abandonment, ongoing threats from
multiple factors, and the challenging ocean conditions expected for piscivorous seabirds in the years ahead,
Tufted Puffins are likely to continue declining in Washington. If the current 8.9% annual rate of decline
continues, the state’s population could become functionally extirpated within about 40 years. For these
reasons, it is recommended that the Tufted Puffin be listed as endangered in Washington.

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 INTRODUCTION
Tufted Puffins (Fratercula cirrhata) are an iconic seabird species. With their brilliant orange bills and white
faces framed by golden plumes, they adorn the covers of field guides and feature regularly in wildlife films,
photo collections, and animal calendars around the world. Their likeness can also be found on souvenir tshirts, postcards, posters, figurines, stuffed animals, and jigsaw puzzles, as well as the label of an awardwinning red wine from Washington state. Aquariums from Tacoma to Tokyo maintain popular live puffin
exhibits, and one Northwest coastal community organizes an annual “Puffin Watch” as an alternative to
fireworks on the Fourth of July.
Inhabiting a wide geographic zone across the temperate and subarctic North Pacific, Tufted Puffins spend
much of the year foraging singly or in small groups far out to sea. They are considered the most pelagic and
far-ranging member of the auk family (Alcidae) (Piatt and Kitaysky 2002). During spring and summer,
however, the birds congregate to breed on rocky coastal islets and headlands from Japan north to Siberia, and
from Alaska south as far as California. It is their habits during the breeding season that have long brought
them into contact with human cultures. Salish, Tlingit, Haida, and other native peoples have hunted Tufted
Puffins at their breeding colonies for at least 1,600 years (Bovy 2007, Moss 2007), consuming their meat and
stitching their feathered skins into garments (Bent 1919). Puffin eggs, too, were an important traditional food
source throughout the coastal Northwest, while the birds’ bright bill plates provided colorful decorations for
dance rattles and the fringes of ceremonial robes (Piatt and Kitaysky 2002).
Following European settlement, early commercial fishermen came to regard the species as a wily adversary.
Trollers in Southeast Alaska reported Tufted Puffins following close behind their skiffs, plucking herring and
other baitfish from lines as deep as 15 fathoms (Heath 1915, Willet 1915). With the advent of gillnet and
driftnet fisheries, the deep-diving birds often became ensnared and drowned in nets. In the mid- to late 20th
century, Tufted Puffins ranked among the most commonly caught seabirds in the salmon and squid net
fisheries, with tens of thousands or even hundreds of thousands of birds killed annually (DeGange and Hay
1991, DeGange et al. 1993). These bycatch losses, combined with the birds’ high vulnerability to oil spills,
brought Tufted Puffins to the attention of wildlife managers and conservationists in the 1970s. At the same
time, biologists began recognizing puffins and other fish-eating seabirds as important indicators for the health
of marine systems (Piatt et al. 2007). Feeding high on the food chain and breeding in dense colonies, they
provide a visible and highly sensitive means for gauging a range of trends, from plankton productivity to
ocean warming to climate change (e.g., Hatch and Sanger 1992, Hunt et al. 2002, Hyrenbach and Veit 2003).
Because their natural history is relatively well known, Tufted Puffins can signal deeper ecological patterns
through subtle changes in their diet, behavior, and breeding success. Following the Exxon Valdez oil spill, for
example, Tufted Puffin populations were studied as a potential measure of the recovery of Prince William
Sound (e.g., Agler et al. 1994, 1999, Piatt et al. 1997).
While the late 20th century saw increasing public and scientific appreciation for Tufted Puffins, it also marked
a steep decline in populations throughout the southern portions of their range. Puffins decreased
dramatically or disappeared entirely from dozens of colonies in California, Oregon, Washington, and Japan
(Piatt and Kitaysky 2002). Despite the species’ prominence, information on the rate and cause of the decline
remained largely anecdotal.
In Washington, concern for Tufted Puffin populations intensified in the 1990s following the abandonment of
former breeding colonies in the San Juan Islands and the steady downward trend in outer coastal populations
and offshore sightings (Speich and Wahl 1989, Wahl and Tweit 2000). Tufted Puffins were added to the
Washington Department of Fish and Wildlife’s list of candidate species in October 1998 to be reviewed for
potential listing as endangered, threatened, or sensitive (per Policy 5301 [formerly Policy 6001]).

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 Per Washington Administrative Code (WAC) 232-12-297, the purpose of this status report is to collate the
available scientific data on Tufted Puffins so that the Washington Department of Fish and Wildlife can
prepare an informed state listing recommendation of endangered, threatened, or sensitive to the Washington
Fish and Wildlife Commission. The report presents all known data on the historical and current status of the
puffin population and its habitats in the state, as well as threats to the population.

TAXONOMY
Tufted Puffins are members of the auk family (Alcidae), a group of diving seabirds that includes puffins,
auklets, murrelets, guillemots, Razorbills (Alca torda), and Dovekies (Alle alle). Alcids share the order
Charadriiformes with the shorebirds and gulls. The 24 extant alcid species inhabit temperate waters of the
northern oceans, with 15 species known from Washington. Alcids are stout-bodied divers specifically
adapted for underwater pursuit foraging. Diversity within the family is considered a classic example of
adaptive radiation, with bill and body-sizes specialized for plankton feeding, fish feeding or, in the case of
puffins, an intermediate strategy that takes advantage of both (Bédard 1969). Genetic and morphological
analyses divide the alcids into six major lineages, with Tufted Puffins found in a close-knit group made up of
their sister species, the Horned (F. corniculata) and Atlantic (F. arctica) Puffins, as well as the Rhinoceros Auklet
(Cerorhinca monocerata) (Friesen et al. 1996, Pereira and Baker 2008). Of these, Rhinoceros Auklets are also
regular breeders in Washington waters and often use the same nesting colonies as Tufted Puffins. The close
behavioral and ecological relationship between these two species has been demonstrated by exchange
experiments, where adults of each species successfully raised chicks of the other to fledging (Vermeer and
Cullen 1979).
The Tufted Puffin is classified in the genus Fratercula, from the Latin for “little brother” or “little monk,” and
with the specific epithet cirrhata, Latin for “tufted.” It shares this genus with the Horned and Atlantic puffins,
though until recently its larger size and distinctive plumage led taxonomists to place it in its own genus, Lunda
(AOU 1998, Sangster et al. 2011). No subspecies are recognized. Though its breeding range overlaps with
that of the closely-related Horned Puffin and Rhinoceros Auklet, no hybrids are known.

DESCRIPTION
General description . Tufted Puffins are among the largest alcids, measuring 35-40 cm (14-16 in) in length
and weighing on average 775 g (1.7 lb) (Piatt and Kitaysky 2002). They share the general alcid family traits of
short, stubby wings and legs set well back on a stocky body. Their characteristic rapid flight strokes and lack
of maneuverability in the air stem from high wing loading, and they often require a runway of clear water for
takeoffs and landings. Any awkwardness disappears beneath the surface, however, where diving puffins “fly”
rapidly and gracefully underwater in pursuit of prey. Tufted Puffins have large, triangular orange bills that are
laterally compressed and grooved on the upper mandible, and their orange feet are strongly clawed for aid in
digging, as well as providing traction on land. Males grow slightly larger than females, but the sexes are
otherwise indistinguishable in the field (Piatt and Kitaysky 2002).
Plumages and other features. The iconic image of a Tufted Puffin features the bird in its breeding
plumage: jet black body feathers, with a white face framed by long golden plumes that sweep backward and
down the neck. The breeding colors occur from April to September and are augmented by brilliant orange
bill plates, orange rictal bristles, and a crimson ring of bare skin around the eye. The non-breeding plumage
(October to March) is less familiar because the birds are usually far out at sea. Body feathers are brownish
black, with a dusky black face. The golden tufts and rictal bristles are gone, and the skin around the eye
becomes dark. The bright bill plates are also lost, making the bill appear smaller (though still orange).
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 Immature birds strongly resemble winter adults and do not develop breeding plumage until at least their
second year. Bill size is proportionally smaller than in adults.

Vocalizations. Tufted Puffins are generally quiet birds, but communicate with a range of low purrs and
growls at breeding sites and when gathered on the water. Although puffin vocalizations remain poorly
studied, at least four distinct calls have been identified in adults (Piatt and Kitaysky 2002), while the peeping
of chicks differs according to levels of hunger and stress (Gjerdrum et al. 2006).

GEOGRAPHICAL DISTRIBUTION
World
Tufted Puffins range throughout the temperate and sub-arctic North Pacific (Nettleship 1996, Piatt and
Kitaysky 2002). Though vagrants have been noted as far south as Laysan Island in the Northwestern
Hawaiian Islands (Clapp and Giezentanner 1979, Clapp 1986), they are generally restricted to the cool waters
above 30–34°N latitude (Gould and Piatt 1993). Like other alcids, as well as penguins, Tufted Puffins rely on
cold water to help dispel body heat produced by underwater “flight,” and their energy requirements depend
on the high prey densities found in high latitude oceans (Nettleship 1996, Van Buren and Boersma 2007).
Warm, low-productivity tropical waters present a dispersal barrier to both groups (Sparks and Soper 1987,
Pereira and Baker 2008).
Relatively little is known about Tufted Puffins during winter on the open seas, but individuals and small
groups have been sighted on surveys throughout the central North Pacific (Gould and Piatt 1993), where they
were also a common bycatch in late 20th century driftnet fisheries (DeGange et al. 1993). Sub-adults may
remain at sea year-round, but breeding birds congregate on rocky coastal islands from as early as March
through September. Breeding concentrations are highest around the Bering Sea, Aleutian Islands, and Gulf
of Alaska, but colonies also occur along the Asian coast as far south as Hokkaido, Japan (Brazil 1991, Osa
and Watanuki 2002), and in North America as far south as the Channel Islands in California (McChesney et
al. 1995).

North America
Of the 1,031 nesting colonies known worldwide in about 2000, 802 (78%) occur in North America (Piatt and
Kitaysky 2002). These are concentrated in Alaska, particularly in the Aleutian Islands and along the Alaskan
Peninsula, where some individual colonies host more than 100,000 birds. Tufted Puffins also breed in
significant numbers in Southeast Alaska and British Columbia, and less commonly in Washington, Oregon,
and California.

Washington
In Washington, Tufted Puffin breeding colonies lie mainly along the outer coast from Point Grenville north
to Cape Flattery (Speich and Wahl 1989, Wahl 2005). The species formerly bred in small numbers at sites
throughout the San Juan Islands (e.g., Edson 1908, 1935), but nesting colonies on inland marine waters are
now restricted to Protection and Smith islands in the eastern Strait of Juan de Fuca (P. Hodum, et al., unpubl.
data). Table 1 lists all known former and current nesting sites in Washington. No breeding colonies were
ever detected in Puget Sound.
During the winter months, Tufted Puffins migrate far offshore, but a few occur in Washington waters over
the continental shelf as late as October (Wahl 1975), and birds occasionally wash up along outer coastal
beaches after winter storm events (Lawrence 1892, Dawson and Bowles 1909). Puffins have been sighted
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 between 64 and 155 km offshore in January
(Wahl 2005). An early description of Tufted
Puffins as common winter residents around
Tacoma (Bowles 1922) was certainly in error, as
nearly all the birds are far out to sea at that time
of year. Casual winter sightings of Tufted
Puffins in Washington waters can usually be
attributed to misidentified Rhinoceros Auklets,
because the two species resemble one another
closely during winter when out of breeding
plumage.
The species occasionally wanders south from the
Strait of Juan de Fuca into northern Puget
Sound (e.g., Wahl 2005, Hunn 2012). A
specimen collected in the mid-19th century near
Fort Steilacoom in present-day Lakewood,
Pierce County, is the only confirmed occurrence
from southern Puget Sound (Jewett et al. 1953).

NATURAL HISTORY
Daily Habits
Few studies have quantified the time budget for
Tufted Puffins and very little is known about the
activities of birds wintering at sea. They are
diurnal, however, and during the breeding
season, birds show activity peaks in the morning
and evening hours. At Tatoosh Island,
Washington, for example, summer counts of
birds in and around the colony peaked at 11 a.m.
and 9 p.m., with mornings generally busier than
afternoons (Figure 1; P. Hodum et al., unpubl.
data). Colony attendance becomes more
consistent throughout the day as the season
progresses, though provisioning visits to chicks
remain concentrated in the morning (Piatt and
Kitaysky 2002). Provisioning trips may last
several hours, during which adults forage alone,
in small groups, or in mixed-species flocks.
Preening and resting occurs on land or on the
water, and birds generally sleep on the water
near nesting sites, though burrows may also be
used. During winter and for non-breeding
individuals, all activities occur on the water but
specific patterns are unknown. Birds winter
alone or in small groups at sea.

January 2015

Table 1. Name, location, and occupancy status of
known Tufted Puffin breeding sites in Washington.
Occupancy data for each site appear in Appendix A.
Colony name1
Bodelteh Islands (East)
Bodelteh Islands (Middle & West)
Cake Rock
Cakesosta
Carroll Island
Dhuoyautzachtahl (Petrel Rock)
Dohodaaluh
Hand Rock
Jagged Island (Wishalooth, Bald Is.)
James Island
Kochaauh
Seal Rock
Silver Sides
Table Rock
Tatoosh Island
Unnamed Rock 1
Unnamed Rock 2
White Rock
Erin’s Bride
Grenville Arch
Grenville Pillar
Point Grenville Cliffs
Puffin Rock (Erin)
Willoughby Rock
Smith Island
Alexander Island
Destruction Island
Ghost Rock
Half Round Rock
Protection Island
Rounded Island
Bare Island
Bird Rocks
Castle Island
Colville Island
Flattop Island
Hall Island
Matia Island
Puffin Island
Skipjack Island
South Peapod
Sucia Islands
Viti Rocks
Williamson Rocks
1

County2
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
G
G
G
G
G
G
I
J
J
J
J
J
J
SJ
SJ
SJ
SJ
SJ
SJ
SJ
SJ
SJ
SJ
SJ
Sk
Sk

Status3
(1982)
O
O
O
O
O
(1907)
(1915)
O
O
O
(1981)
O
(1978)
O
?(1978)
?(1982)
O
O
(1915)
(1980)
?(1980)
O
O
O
O
O
?(1981)
(1981)
O
O
(1978)
(1984)
(1928)
(1984)
(1949)
(1990)
(1984)
(1982)
(1936)
(1949)
(1886)
(1970s)
(1985)

Colony names conform to Speich and Wahl (1989). 2 C=Clallam,
G=Grays Harbor, I=Island, J=Jefferson, SJ=San Juan, Sk=Skagit.
3
O=currently occupied; dates in parentheses indicate year of last
confirmed occupancy; sites with question marks were not checked
during the 2007-2010 surveys of P. Hodum et al., thus current
status is unknown. Data are from Speich and Wahl (1989),
Osborne et al. (1998), USFWS (2010; unpubl. data), P. Hodum et
al. (unpubl. data), and T. Scruton (pers. comm.).

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 Figure 1. Hourly attendance patterns of Tufted Puffins at Tatoosh Island, Washington, from
June 1 to August 24, 2005–2008, averaged across years (P. Hodum et al., unpubl. data).
Data are instantaneous counts of all birds present (on colony, on water, in air).

Diet and Foraging
Tufted Puffins are diving, underwater pursuit foragers with bills adapted to take both fish and invertebrates
(e.g., Bédard 1969, Baird 1991, Davies et al. 2009). Their diet includes an exceptional range of prey – one
study found Tufted Puffins consuming 47 different species in the same habitat whereas the closely-related
Horned Puffin took only 20 (Piatt et al. unpubl., cited in Piatt and Kitaysky 2002). However, foraging
behavior and diet vary considerably from year to year, based on fluctuations in the availability of prey
(e.g., Baird 1990, 1991, Hatch and Sanger 1992). Diet may also vary between colonies located near one
another (Williams and Buck 2010). In general, wintering birds and breeding birds that forage over deep,
oceanic habitats exhibit a broader diet, including a higher proportion of invertebrates, particularly euphausiids
(i.e., krill, shrimp-like crustaceans), while breeders foraging near coastlines and over the continental shelf take
more fish (Baird 1991, Piatt and Kitaysky 2002).
Fish comprise the bulk of the nestling diet, >95% in some studies, and research consistently links the type
and availability of fish in nearby waters to the growth rates and fledging success of chicks (Baird 1990, Hatch
and Sanger 1992, Hipfner et al. 2007, Williams and Buck 2010). At Triangle Island, British Columbia, adults
exhibit a dramatic shift in habitat use and diet over the course of the nesting season. Early in the season they
forage on abundant pelagic zooplankton and squid of modest quality, but switch to coastal fish species of
higher nutritional quality later in the season when they are feeding growing chicks and keeping themselves
fortified for long provisioning flights (Davies et al. 2009). Studies in Alaska, however, suggest that adults
continue to support themselves on invertebrates while feeding fish to their nestlings (Piatt and Kitaysky
2002). Some evidence suggests that adults, particularly males, lose body mass sharply in lean years, depriving
themselves of food to keep their chicks fed (Williams et al. 2007). In times of extreme scarcity, however,
individual nestlings or even whole colonies may be abandoned mid-season (Vermeer and Cullen 1979,
Vermeer et al. 1979, Gjerdum et al. 2003). The fidelity of nestling diets to local fish stocks is well known and
has been recommended for monitoring as an indicator of forage fish availability (Hatch and Sanger 1992).
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 Though no dietary studies for Tufted Puffins have been conducted in Washington, research in British
Columbia and Alaska has documented a range of prey species that are available to puffins nesting in
Washington (e.g., Wehle 1983, Piatt et al. 1997, Gjerdum 2004, Williams and Buck 2010). Pacific Sand Lance
(Ammodytes hexapterus) and Pacific Herring (Clupea pallasii) rank among the most important prey species, but
the list also includes juvenile rockfish (Sebastes spp.), Eulachon (Thaleichthys pacificus), Pacific Sardine (Sardinops
sagax), and Northern Anchovy (Engraulis mordax), as well as various squid, euphausiids, and other
invertebrates. Puffins on the Washington coast have been spotted with bill loads containing sand lance,
herring, anchovy, and smelt (Cody 1973, Frazer 1975). A number of additional fish species in Washington
fall within the 20 to 200 mm size range preferred by puffins during the breeding season (Piatt and Kitaysky
2002), including Surf Smelt (Hypomesus pretiosus), Night Smelt (Spirinchus starksi), Popeye Blacksmelt (Bathylagus
ochotensis), Whitebait Smelt (Allosmerus elongatus), Pacific Saury (Cololabis saira), Kelp Greenling (Hexagrammos
decagrammus), and juvenile salmon (Oncorhynchus spp.). It is also likely that diets of puffins nesting on the outer
coast vary from those nesting in inland marine waters, a pattern that has been reported for other Washington
alcids. For example, the diet of Rhinoceros Auklets on Protection Island near the eastern end of the Strait of
Juan de Fuca is dominated by Pacific Herring and Pacific Sand Lance while the coastal diet (Tatoosh and
Destruction islands) is much more diverse and includes Northern Anchovy, Pacific Sand Lance, Pacific
Herring, Whitebait and Surf Smelt, juvenile rockfish, and Pacific Saury (Pearson et al. in review).
Details of Tufted Puffin foraging behavior derive largely from studies of birds at breeding colonies. For
breeding birds, the demands of a growing chick dictate their daily foraging routine. Evidence suggests that
puffins will forgo high quality foraging sites in favor of areas located close to their colonies (Sealy 1973,
Ostrand et al. 1998). Just what determines an individual’s foraging strategy remains unclear, but site choice
and target prey can vary dramatically between sub-groups within the same colony, with measurable impacts
on nestling development (Hipfner et al. 2007). It’s possible that individuals pattern their foraging habits after
those of other puffins foraging nearby, a flocking impulse that persists regardless of foraging success (Hipfner
et al. 2007). On the outer coast of Washington, adults fly an average of 4.67 km from breeding sites to feed,
but sometimes venture as far as 100 km from their colonies (Cody 1973). Tufted Puffins generally forage
singly or in small groups, and position themselves on the periphery when joining mixed-species flocks
(Hoffman et al. 1981, Tyler et al. 1993, Ostrand et al. 1998). Most prey are probably caught within 60 m of
the surface, but puffin dives to depths of 110 m have been recorded, ranking them among the deepest-diving
alcids (Piatt and Kitaysky 2002). Captive birds spend more time foraging at depth than almost any other
species (Duffy et al. 1987). When feeding themselves, prey items are consumed immediately, but puffins
often capture and hold multiple fish when provisioning their young. Bill loads vary, but generally include five
or more fish, and sometimes reach more than 20 fish (Wehle 1983). Foraging puffins on the water
sometimes grow accustomed to the presence of people and boats and have been known to steal bait from
fishing lines (Heath 1915, Willet 1915), and successfully “beg” for bait fish tossed to them from sport fishers
(L. Slater, pers. comm.).
Dietary information for wintering birds remains poorly known because individuals disperse too widely on the
open seas for practical observation. It’s generally believed, however, that birds outside of the breeding season
take a higher proportion of squid and euphausiids (Piatt and Kitaysky 2002).

Reproduction, Longevity, and Survival
Breeding season. The Tufted Puffin breeding season begins as early as March in California and as late as

June at the northern extent of the species’ range. In Washington, puffins begin returning to their nesting
colonies in mid-April (Frazer 1974, 1975, Burrell 1980). Egg laying peaks from May to early June and parents
remain in the vicinity of the colony provisioning their chicks until they fledge in mid-August or early
September, when all birds depart for the open sea (Frazer 1975, Burrell 1980).

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 Breeding behavior and territoriality. Tufted Puffins become gregarious during the breeding season when
they nest in colonies. Upon arrival at nesting colonies, birds spend a week or more re-establishing pair bonds
and occupying or excavating burrows (Frazer 1975). They display high mate fidelity, establishing pair bonds
that have been observed to last for consecutive seasons (Wehle 1980) and are thought to endure longer (Piatt
and Kitaysky 2002). Courtship behaviors include billing and displaying with nesting materials, as well as chase
and copulation sequences on the water where the male jerks his head rapidly while opening and closing his
bright bill (Piatt and Kitaysky 2002). Individuals and pairs compete for nesting sites and defend burrow
entrances, with fights and aggressive displays common during the early breeding season. Adults defend the
area around their burrow entrance, but given high nest densities at some colonies (e.g., Dawson and Bowles
1909), this can amount to an area of 1.5 m2 or less.
Nesting and brood rearing. Tufted Puffins excavate straight, curved, or branched nesting burrows in the

soil on offshore islands, inaccessible headlands, or occasionally in estuaries (Gill and Sanger 1979, Piatt and
Kitaysky 2002). Burrows generally range from 0.5–1.5 m deep and 13.5–19.0 cm in diameter, though depths
of up to 5 m have been reported (Piatt and Kitaysky 2002). Where surface material is thin or when
competition for choice nest sites is fierce, rocky crevices, overhangs, and even talus slopes may be used (e.g.,
Edson 1929a, Bailey and Faust 1981). Clutches consist of one whitish egg 71–74 mm long and 48–49 mm
wide (Piatt and Kitaysky 2002). A replacement egg may be laid within about 21 days if the first is lost (Wehle
1980). Adults take turns incubating for about 43–46 days (range 41–53 days) (Piatt and Kitaysky 2002).
Average hatching date at Destruction Island, Washington, is July 1 (range, June 21–July 24; Burrell 1980).
Adults remain with the hatchling for several days until it develops the ability to thermoregulate (Wehle 1980).
Both parents provision the growing chick until it fledges, together bringing an average of four bill loads of
fish per day (Piatt and Kitaysky 2002). Evidence suggests that adults adjust the number of provisioning trips
based on the development and corresponding nutritional requirements of the chick, and begin reducing the
amount of food sharply late in the nestling period to help induce chicks to fledge (Gjerdrum 2004). The
nestling period averages about 48 days (range 38-59 days) (Wehle 1980, Piatt and Kitaysky 2002).

Chick growth and fledging. Hatchling weights and growth rates vary by colony and season, but chicks
generally weigh 60–70 g at hatching and reach about 450–577 g at fledging (Piatt and Kitaysky 2002, Williams
and Buck 2010). Chick development appears to favor the most important physical characters, allocating
resources to wing length (necessary for the first flight to sea and subsequent foraging), and body mass (fat
reserves for survival during the critical post-fledging period, while learning to feed) (Gjerdrum 2004).
Fledglings with larger weights and longer wing lengths exhibit higher survival and earlier fledging dates
(Morrison et al. 2009). Puffins leave the nest after six to eight weeks, an age when they still lack the ability to
fly well, but are agile in the water and can make short dives in pursuit of prey (Piatt and Kitaysky 2002).
Fledging generally occurs under the cover of darkness to avoid predators, with chicks walking, hopping, or
flying from burrows to the water where they are immediately independent of the parents.
Fledgling survival varies markedly by year depending on food availability and corresponding condition of the
chicks. A survey of multiple studies at various sites found a mean (± SD) fledging success of 64 ± 28% and a
mean breeding success of 0.43 ± 0.22 chicks produced per pair (Piatt and Kitaysky 2002). On Tatoosh
Island, burrow occupancy ranged from 72-91%, hatching success from 47-90%, and chick survival from 3075% for the 19-25 burrows surveyed per year between 2005 and 2008 (P. Hodum et al., unpubl. data).
Overall fledging success ranged from 23-63%. The 2005 season was anomalous because of late spring
upwelling that resulted in widespread colony abandonment by other seabirds in the California Current (e.g.,
Sydeman et al. 2006). However, even without the 2005 data, fledging success for 2006-2008 ranged from 3563%, which is considerably lower than that reported by Piatt and Kitaysky (2002) for a range of sites, and
considerably lower than that of Rhinoceros Auklets (66-89%) nesting in burrows on the same colony during
the same years (2006-2008; P. Hodum et al., unpubl. data; Pearson et al. in review).

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 Longevity and survival. The lifespan of wild Tufted Puffins has not been determined (Piatt and Kitaysky
2002), but captive birds regularly live 15-20 years and occasionally longer than 30 years (S. Perry, pers.
comm.). Morrison et al. (2011) reported average annual survival rates of 95 ± 3% in adult females and 92 ±
4% in adult males.
Tufted Puffins comprise a small proportion of the beached (dead) seabirds found along shorelines in
Washington during surveys conducted by the Coastal Observation and Seabird Survey Team (COASST)
program (J. Parrish, pers. comm.). The vast majority of Tufted Puffins recorded from 2001 to 2012 were
counted along the outer coast, with very few or none noted in the Strait of Juan de Fuca, the San Juan
Islands, and Puget Sound. Along the outer coast, 0 to 7 beached individuals were found per year during this
period, except in 2012, when 82 birds were recorded between mid-January and mid-April. This mortality
event followed an extended period of stormy weather in December 2011 (S. Horton, pers. comm.). No
monthly pattern in beachings was evident during other survey years.

Migration and Dispersal
Migration. Tufted Puffins migrate annually between their coastal breeding colonies and wintering areas in

the North Pacific. The exact winter range of Washington’s population remains unknown, but nearly all birds
are absent from coastal and shelf waters off British Columbia, Washington, Oregon, and California from late
fall until spring (Tyler et al. 1993, Wahl et al. 1993). Tufted Puffins have long been considered the most
pelagic of the alcids (Bent 1919), but very little is known about the routes they use during migration or the
extent of their winter movements. Retreats and advances in pack ice determine migratory patterns for the
most northern populations, but little else is known definitively about puffin movements at sea. Attempts at
radio and satellite telemetry have led to altered behaviors, nest abandonment, and even mortality in subject
birds, and no tags have yielded data beyond the breeding season (Hatch et al. 2000a, 2000b, Whidden et al.
2007). Flight habits, foraging stops, flocking, and other migratory behaviors are not well understood, though
mass arrivals at breeding colonies have been observed (Piatt and Kitaysky 2002).

Dispersal. Information about the dispersal of young Tufted Puffins remains scant, though fledglings
probably depart for pelagic wintering areas soon after leaving their burrows (Piatt and Kitaysky 2002). Some
degree of fidelity to natal nesting colonies is assumed among adults and yearlings. Investigators have
speculated that large peaks in colony attendance early in the spring reflect the return of yearling, non-breeding
birds to their natal colonies (Frazer 1975). Shortages of suitable nesting sites point to overcrowding at some
colonies (e.g., Dawson and Bowles 1909), and seabirds are known to show higher reproductive success in less
crowded conditions (Hunt et al. 1986). An ecological incentive for dispersal therefore exists, and the reoccupation of former breeding colonies has occurred (McChesney et al. 1995), but the vital questions of how
often and by whom remain unanswered.

Molting
Like most other alcids, Tufted Puffins exhibit an intense and rapid molting of their flight feathers, where first
the primaries and then the secondaries are dropped in rapid succession, reducing wing area by up to 40%
(Bridge 2004). The rectrices (tail feathers) follow the secondaries and new growth of all three flight feather
types is completed at roughly the same time (Thompson and Kitaysky 2004). This molt occurs in the spring
for second-year birds (Thompson and Kitaysky 2004) and in the fall for adults (Piatt and Kitaysky 2002,
Bridge 2004). Though it reduces diving efficiency and renders the birds flightless for 32-45 days (Bridge
2004, Thompson and Kitaysky 2004), the rapid molt strategy is thought to reduce the overall period of flight
disruption. It is common in birds like puffins and other alcids, where high wing-loading makes the sequential
feather loss of a traditional molt a far more prolonged threat to their ability to fly (Ehrlich et al. 1988). In
contrast to their flight feathers, Tufted Puffins molt their body plumage over a much longer time frame twice
a year, once in the late summer and fall, and again from February to April (Piatt and Kitaysky 2002).
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 Ecological Relationships
Competition . Though the topic has received relatively little research attention, Tufted Puffins engage in a

range of competitive interactions, particularly in and around breeding colonies. They have been observed
displacing Horned Puffins at nesting sites and are thought to compete aggressively with other burrow-nesting
alcids as well (Piatt and Kitaysky 2002). On the other hand, Ancient Murrelets (Synthliboramphus antiquus),
Rhinoceros Auklets, Leach’s Storm-Petrels (Oceanodroma leucorhoa), and Fork-tailed Storm-Petrels (O. furcata)
have all nested successfully in side-tunnels off of Tufted Puffin burrows (Frazer 1975, Piatt and Kitaysky
2002). The nocturnal habit of these species presumably reduces direct competition with puffins.
In a study of six captive alcid species, Tufted Puffins were the most aggressive underwater, chasing and
stealing fish from Common Murres (Uria aalge), Rhinoceros Auklets, Cassin’s Auklets (Ptychoramphus aleuticus),
and Pigeon Guillemots (Cepphus columba), as well as conspecifics (Duffy et al. 1987). Puffins also regularly join
mixed-species flocks where their diving behaviors help concentrate fish prey schools, potentially benefiting
surface-feeding birds (Hoffman et al. 1981).
In general, competition among co-occurring alcids may be reduced through differences in foraging strategies
(e.g., distance from nesting colony) and nesting sites (burrows vs. crevices vs. cliffs) (Cody 1973). Still, a
meta-study of seabird colonies underscores the important role of intra- and interspecific competition for food
and nesting sites, because birds in smaller colonies fared better in a range of reproductive measures than
those breeding at large, crowded sites (Hunt et al. 1986). In California’s Farallon Islands, competition for
nesting sites is exacerbated by the presence of non-native European Rabbits (Oryctolagus cuniculus), another
burrowing species (Ainley and Lewis 1974).

Predation. A number of predators are known to attack puffins. Brown Bears (Ursus arctos) excavate Tufted

Puffin burrows and have been implicated in population declines or extirpations from several Alaskan
breeding colonies (Bailey and Faust 1981). Puffin remains have been found in Bald Eagle (Haliaeetus
leucocephalus) nests (Vermeer et al. 1976; see Factors Affecting Continued Existence – Bald Eagle Predation)
and in the pellets of Snowy Owls (Bubo scandiacus; Williams and Frank 1979), and several authors report
observing attacks by Peregrine Falcons (Falco peregrinus; Vermeer et al. 1976, Addison et al. 2007). River
Otters (Lontra canadensis) eliminated the last pair of puffins nesting on Mandarte Island, British Columbia (P.
Arcese, pers. comm.), and are known to take chicks and adults at East Amatuli Island, Alaska (A. Kettle, pers.
comm.). River Otters have been observed entering unidentified seabird burrows on Alexander Island,
Washington (S. Pearson, pers. comm.), which has one of the largest remaining colonies of Tufted Puffins in
the state.
Studies of Bald Eagle predation on another alcid, the Common Murre, have revealed that the indirect effects
of predation (e.g., less time spent defending eggs) can also have important consequences (Parrish et al. 2001).
Puffins alter their flight patterns and activity rates in the presence of eagles and falcons (Addison et al. 2007),
but the effects of raptor predation on population trends remains unclear.
The strongest documented impact of predation on Tufted Puffins comes from the introduction of non-native
mammalian predators to nesting colonies. The introduction of Arctic Foxes (Alopex lagopus) for fur farming
led to severe population declines on several Alaskan islands (Bailey 1976, Bailey and Faust 1981), while
introduced Red Foxes (Vulpes vulpes), Arctic Ground Squirrels (Urocitellus parryii), and Norway Rats (Rattus
norvegicus) have adversely impacted a number of colonies throughout the puffin’s range (see summary in Piatt
and Kitaysky 2002). Other documented predators of adults, nestlings, and/or eggs include Common Ravens
(Corvus corax), Glaucous-winged Gulls (Larus glaucescens), Steller’s Sea Eagles (Haliaeetus pelagicus), and Eurasian
Eagle Owls (Bubo bubo) (Piatt and Kitaysky 2002). Several other common predators known to prey on
burrow-nesting seabirds have yet to be confirmed depredating Tufted Puffins, including Black Rats (Rattus
rattus), Raccoons (Procyon lotor), and American Mink (Neovison vison).
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 Kleptoparasitism. The puffin habit of carrying whole fish crosswise in their bills makes them a favored
target of kleptoparasites. Glaucous-winged Gulls, Red-legged Kittiwakes (Rissa brevirostris), Black-legged
Kittiwakes (R. tridactyla), and Parasitic Jaegers (Stercorarius parasiticus) regularly harass puffins foraging in mixed
flocks, as well as during their approaches to colonies. Tufted Puffins appear to adjust their overflights and
approaches to reduce the risk of kleptoparasitism (Blackburn et al. 2009), a behavior confirmed in Atlantic
Puffins (Rice 1987). Glaucous-winged and Western (Larus occidentalis) gulls are the most common
kleptoparasites at breeding colonies in Washington (Cody 1973, Frazer 1975) and at Triangle Island, British
Columbia (St. Clair et al. 2001, Blackburn et al. 2009). A study at Triangle Island found that rates of
kleptoparasitism varied among years, but that general incidence was low and there was no relationship
between rates of kleptoparasitism and puffin breeding success (St. Clair et al. 2001). Investigators generally
agree that kleptoparasitism may contribute to the challenge of provisioning chicks during difficult years, but
probably doesn’t constitute a major ecological constraint (Piatt and Kitaysky 2002). Similar conclusions have
been reached about gull kleptoparasitism on Atlantic Puffins (Pierotti 1983, Rice 1985) and Rhinoceros
Auklets (Wilson 1993), although auklet chicks in gull-free areas on Protection Island, Washington, did grow
significantly faster than chicks from burrows located within a gull colony (Wilson 1993).

Diseases and Parasites
Pathogens and diseases affecting Tufted Puffins are poorly understood. No major mortality events have been
attributed to outbreaks of illness, but Tufted Puffins are known to carry antibodies to the Tyuleniy virus, a
tick-borne flavivirus related to West Nile virus, yellow fever, and dengue fever (Lvov et al. 1972). Parasitic
nematodes have been recovered from Tufted Puffins at high enough concentrations to cause ulcerated lesions
(Nagasawa et al. 1998), but the effects of worms on puffin health remain unknown. In general, Tufted
Puffins support fewer numbers and species of internal parasites than other alcids (Piatt and Kitaysky 2002).
Captive populations show susceptibility to a fatal form of avian mycosis, a fungal infection, but this has not
been studied in the wild (Monroe et al. 1994). Ectoparasites including ticks and lice have been noted in
several populations, but have not been studied in detail (Piatt and Kitaysky 2002).

HABITAT REQUIREMENTS
Nesting habitat. Tufted Puffins nest in burrows, rocky crevices, and occasionally in dense shrubbery on
isolated offshore islands and inaccessible headlands (Dawson and Bowles 1909, Nettleship 1996, Piatt and
Kitaysky 2002). Ideal habitat includes grassy slopes, bluffs, and plateaus with soil deep enough for burrowing
in areas that are free of introduced predators (e.g., foxes, rats) and human disturbance. Rocky areas and
thickets may be utilized, but are not preferred. Access to steep slopes, cliff edges, or elevated rocks is
important for taking flight. In British Columbia, colony locations are characterized by lower than average
rainfall intensity, lower than average air and surface water temperatures, and higher salinity in surrounding
waters than sites without colonies (Kaiser and Forbes 1992). Nesting Tufted Puffins are sensitive to
disturbance, generally avoiding inhabited areas (e.g., islands with manned light stations) and often abandoning
nests accessed for scientific observation (e.g., Frazer 1975, Pierce and Simons 1986, Hatch et al. 2000b; see
Factors Affecting Continued Existence – Human Disturbance).
Foraging habitat. During the breeding season, adult Tufted Puffins range as far as 100 km from their
breeding colonies, utilizing open water foraging habitats from the nearshore to the open sea (Cody 1973, Piatt
and Kitaysky 2002). Like many other seabirds, they often congregate where currents, coastline features, and
other conditions concentrate schools of prey (Piatt and Kitaysky 2002, Zamon 2003). Foraging habitat
during the breeding period requires access to adequate supplies of sand lance, herring, or other fish species
that are fed to the young.

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 Wintering habitat. Tufted Puffins spend the winter at sea in the North Pacific, though they also
infrequently use continental shelf waters and are occasionally spotted off of Washington’s outer shores or
washed up on coastal beaches (Lawrence 1892, Dawson and Bowles 1909, Alcorn 1959, Wahl 2005). Little is
known about their habitat requirements in the winter months. They have little contact with people during
this period, but are vulnerable to chronic oil discharges, ingested plastics, and illegal driftnet fisheries
(DeGange et al. 1993).

POPULATION STATUS AND TRENDS
Observers have estimated Tufted Puffin populations using various measures, including burrow counts,
burrow occupancy, colony counts from boats, on-water counts, aerial surveys, and general estimates of
colony attendance (Piatt and Kitaysky 2002). Each method comes with its own set of strengths, biases, and
limitations. The number of adults visible at a colony varies dramatically depending on the time of day (Figure
1), and daily activity patterns change over the course of the season. Count data are also influenced by weather
and sea conditions, equipment, and observer bias that, if unaccounted for in the sampling design, can make it
difficult to compare estimates across time or from different studies. Still, general patterns and trends emerge
from even the most cautious examination of the available data. This report draws on published studies from
throughout the range of Tufted Puffins, as well as an extensive compilation of historical and contemporary
population estimates for Washington (a full list of Washington population estimates is included in Appendix
A).

Global Status and Trends
Although the International Union for the Conservation of Nature (IUCN) considers the species globally
secure, noting its expansive range and relatively large population (BirdLife International 2012), the steep drop
in puffin numbers throughout the southern portions of their range suggests to many investigators that the
species is undergoing a major range contraction (McChesney et al. 1995, Piatt and Kitaysky 2002, Gjerdrum
et al. 2003). Published estimates put the total number of Tufted Puffins between 2.97 million (Piatt and
Kitaysky 2002) and 3.50 million (Byrd et al. 1993). These numbers represent compilations and extrapolations
of colony estimates, mostly from Alaska where >75% of the species breeds. Little historical data exist to
establish long-term global trends (Byrd et al. 1993), but site-specific surveys and occupancy records show a
dynamic patchwork of population gains and losses.
Even at the heart of their range, Tufted Puffins experience highly variable breeding success and
corresponding changes in local populations. This pattern is typical of colonial nesting seabirds, where raising
young to fledging is tightly linked to fluctuations in nearby food supplies and environmental conditions (Baird
1990). In the Gulf of Alaska, for example, Tufted Puffins declined >60% between the 1970s and 1990s,
probably as a result of changing ocean conditions and related declines in forage fish availability (Piatt and
Anderson 1996). Some colonies in the Aleutian Islands, however, increased by as much as 8.7% annually
over the same general time period (Piatt and Kitaysky 2002, Dragoo et al. 2010). Piatt and Kitaysky (2002)
assembled data from 17 published and unpublished colony surveys (Table 2) and concluded that populations
were stable or increasing in the northern portion of the range, but generally declining sharply throughout the
southern regions.

Regional range contraction. In California, only 13 breeding sites have been occupied in recent years

(McChesney and Carter 2008) and the population of the largest colony, the South Farallon Islands, decreased
from “thousands” in 1911 (Dawson 1911) to about 250 in 2012 (Warzybok et al. 2012). In Oregon, the
breeding population decreased by more than an order of magnitude in 20 years, from 4,858 at 49 sites in 1988
to 142 at 15 sites in 2008 (Kocourek et al. 2009) and 146 at 12 sites in 2009 (Suryan et al. 2012). Oregon’s
largest colony of puffins varied in size from 74 to 143 birds during 2010-2013 (Stephensen 2014). Puffin

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Washington Department of Fish and Wildlife

 Table 2. Population trends of Tufted Puffins in North America (adapted and expanded from Piatt and
Kitaysky 2002).
State/province
area/location
Alaska
W. Aleutians
Nizki I.
Buldir I.
C. Aleutians
Adak I.
E. Aleutians
Bogoslof I.
Aiktak I.
N. Gulf of Alaska
E. Amatuli I.
Pr. William Sound
Pr. William Sound
SE. Alaska
St. Lazaria I.

British Columbia
S. Queen Charlotte I.
Kerouard I.
N. Vancouver I.
Triangle I.
Washington
Strt. of Juan de Fuca
Protection I.
N. Outer Coast
Tatoosh I.
Various
S. Outer Coast
Westport

Oregon
Oregon Coast NWRC
Three Arches Rock
California
N. Coast
Castle Rock
Central Coast
SE Farallon I.

1Type

Type of
count1

No.
counts

Years

Shoreline
Burrow

4
5

1976-1998
1991-2001

Burrow
Burrow

8
16

Burrow

Burrow

7

1988-1995

Population change
Probability

+8.7
+6.8

<0.05
ns3

A
A

<0.001
ns

B
B

+17.9

<0.001

-2.9
-2.9
+3.9

<0.001
ns
ns

B
C
D

ns

E

G

1973-2005
1989-2007

+3.0
no trend

11

1996-2006

-5.9

Colony

2
6

1977-1986

1984-2009

negative
trend

<0.01

Colony

8

1973-2001

-13.9

<0.05

Burrow
Pelagic
Pelagic

Burrow

13
6
7

B

F

1983-2001

-13.6

<0.001
0.01

H
I

3
2

1979-2008
1979-1988

-12.4
-3.6

ns

K
L

6

1972-1999

-6.3

<0.05

M

17
24
19

Shoreline
Colony
Colony

Colony

-10.6

<0.05

A

-16.9
-8.9

Bird Plot
Offshore
Pelagic

1995-2007
1972-1993
1989-2000

Source2

%/year

37

1983-2001
2001-2012

1972-2012

+3.0

<0.001

-

J

N

of count: shoreline = bird count from colony shoreline; burrow = plot count of occupied burrows; pelagic =
bird count at sea; colony = whole colony count; bird plot = bird count within plots; offshore = bird count on
transects within 8 km of shore.
2Sources: A = J. Williams & G. V. Byrd, as cited in Piatt & Kitaysky 2002; B = Dragoo et al. 2010; C = Agler et al.
1999; D = Lance et al. 1999; E = Campbell et al. 1990; F = Rodway & Lemon 2011; G = U. Wilson, as cited in Piatt
& Kitaysky 2002; H = R. T. Paine, as cited in Piatt & Kitaysky 2002; I = S. Pearson, unpubl. data; J = Wahl & Tweit
2000; K = Kocourek et al. 2009; L = Hodder 2003; M = Jaques & Strong 2001; N = Warzybok et al. 2012.
3ns = not significant.
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Washington Department of Fish and Wildlife

 numbers in Washington have also declined dramatically, as discussed in detail below. Declines have been
reported from Hokkaido, Japan, where two-thirds of former breeding colonies are now unoccupied and the
population declined 9–15% annually during the 1970s to 1990s to an estimated 10 remaining breeding pairs at
two sites in about 2012 (Osa and Watanuki 2002, Ono 2012).
While range contraction is apparent, the causes underlying it remain unclear, and may be the result of a
combination of historical and current factors (see Factors Affecting Continued Existence). Population
declines during the 20th century have been attributed to oil spills, drift net and coastal gillnet fisheries (e.g.,
Ainley and Lewis 1974, DeGange and Day 1991), and dramatic population crashes of several important
forage fish stocks (e.g., sardine, anchovy, eulachon). Links have also been found between rising ocean
temperatures, food availability, and puffin breeding success (Golobuva 2002, Gjerdrum et al. 2003).
Determining the exact causes of population declines can be difficult, especially for long-lived species, because
there is often a lag between specific events or ecological changes and detectable population trends. This is
particularly true for seabirds when reproductive success is affected, because long-lived adults returning to
nesting colonies year after year can mask the lack of recruitment. A crash in Norwegian herring stocks in the
1960s, for example, led to large-scale reductions in Atlantic Puffin populations that weren’t fully realized until
the 1980s (Anker-Nilssen et al. 1997). It can also be challenging to determine causation if populations near
the edge of a species’ range (like Tufted Puffins in Washington) are bolstered by emigration from core areas.
In that case, factors affecting the core population and rates of emigration may be undetectable at the fringe.
Though it is also possible that reductions in Tufted Puffins in Washington, Oregon, and California may partly
be due to the northward movement of some individuals to colonies in British Columbia and Alaska, this
seems unlikely because of the strong fidelity to breeding sites and limited dispersal of immature birds among
most puffin species (e.g., Harris and Wanless 2011).

Washington Status and Trends
Historically, ornithologists considered Tufted Puffins among the most common seabirds on the Washington
coast and estimated the breeding population in the tens of thousands (e.g., Rhoads 1893, Dawson and Bowles
1909, Palmer 1927, Jewett et al. 1953, Larrison and Francq 1962, Wahl 2005). Dawson and Bowles (1909)
conservatively put the number of birds along the outer Olympic Coast at about 25,000 birds in the early
1900s, an estimate that was maintained for much of the 20th century (e.g., Jewett et al. 1953, Speich and Wahl
1989). Speich and Wahl (1989) estimated the 1978-1982 nesting population at 23,342 birds, although they
remarked that the population could be ≥50% larger than this number. Though colony declines and
abandonment were observed in the Strait of Juan de Fuca and the San Juan Islands (Jewett et al. 1953, Speich
and Wahl 1989), Tufted Puffins were still considered common in offshore surveys into the 1990s (Briggs et
al. 1992, Wahl et al. 1993). More recent studies, however, point to widespread colony abandonment and a
rapid order of magnitude population decline throughout Washington (e.g., Wahl and Tweit 2000; P. Hodum
et al., unpubl. data). This downward trend in puffin occurrence in Washington is evident, and its strength and
scale are consistent, among all three types of data used for this report: boat-based surveys, breeding colony
occupancy records, and breeding colony attendance counts. Declining trends were particularly evident from
the mid-1980s through the late 1990s, but continue to the present.

Boat-based surveys. Two multi-year boat-based studies contain data for Tufted Puffin populations in
Washington, and both show statistically significant declines in recent decades. From 1972 to 2001, Wahl and
Tweit (2000, unpubl. data) surveyed seabirds annually between July and October during more than 4,400
counts on more than 230 cruises from Westport westward to about 50 km offshore. In the early years of this
period, puffins occurred on 78.6% of counts (Wahl 1975), but sightings decreased dramatically during the
latter part of the study (Figure 2). The overall downward trend averaged 13.6% per year and no puffins were
sighted at all during 198 counts in 1997 (Wahl and Tweit 2000).
A second set of boat-based surveys conducted annually from 2001 to 2012 for Marbled Murrelets
(Brachyramphus marmoratus) and other seabirds from Cape Flattery to Point Grenville (made within 8 km of
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 shore from mid-May to late July and averaging 24 transects totaling about 1,000 km per year, see Raphael et
al. 2007, Falxa et al. 2011) also shows a declining trend in Tufted Puffin density, averaging a loss of 8.9% per
year (Figure 2; S. Pearson, unpubl. data). Using data from the 2009 survey for an expanded area between
Cape Flattery and the mouth of the Columbia River with 1,380 km of transects, S. Pearson (unpubl. data)
generated a minimum breeding season population estimate of 2,958 Tufted Puffins on the water for the outer
coast. This estimate does not account for individuals provisioning chicks or otherwise attending colonies, or
those farther offshore.

Site occupancy records. Forty-four historical breeding locations have been documented in Washington,
with 35 of them still occupied during 1978-1984 (Tables 1, 3, Figure 3, Appendix A; Speich and Wahl 1989;
USFWS, unpubl. data). While this later estimate represented a 20% drop from the historical maximum
number of colonies, most of the abandoned sites had previously held ≤50 birds, so the impact on the overall
population was relatively small. However, surveys from 2007 to 2010, conducted mainly by P. Hodum et al.
(unpubl. data), have revealed a much larger decline in site occupancy since then, with breeding birds found at
only 19 sites. This represents a 46% drop in site occupancy since 1978-1984 and a 57% decline since 18861977. Sites that remain occupied are overwhelmingly (89%, 17 of 19) those that held larger numbers (≥100)
of puffins during 1886-1977, whereas the vast majority (88%, 22 of 25) of those abandoned by 2007-2010
held fewer than 100 birds during 1886-1977.
Colony attendance counts. Multi-year land-based counts of Tufted Puffins exist for two breeding colonies
in Washington, with both showing statistically significant declines. Puffin numbers fell an average of 16.9%
annually at Tatoosh Island between 1983 and 2001 (Figure 2; R. Paine, unpubl. data) and an average of 13.9%
annually at Protection Island between 1973 and 2001 (Figure 2; U. Wilson, unpubl. data). These rates of
decline are similar to those recorded for the on-the-water population during boat-based surveys.
While time series datasets do not exist for other breeding sites, over 400 observations, counts, collections, and
other records of breeding Tufted Puffins have been documented in Washington (Appendix A). These
records represent a range of methodologies that do not allow for statistical comparisons; however, the data
are informative for documenting large-scale changes in population. At Carroll Island, for example, the puffin
population declined from estimates of 10,000 birds in 1907 (Dawson 1908a), to 2,270 birds in 1982 (Speich
and Wahl 1989), to 18-211 birds observed (total number of birds seen on the water, flying, or on the colony)
during repeated boat-based surveys between 2007 and 2010 (P. Hodum et al., unpubl. data). In the Strait of
Juan de Fuca, combined estimates at Protection and Smith islands fell from over 1,070 puffins in the 1950s
(Ainley et al. 1994) to about 60 birds in 2007-2008 (P. Hodum et al., unpubl. data). Of the 19 occupied sites
surveyed along the outer coast and in the Strait of Juan de Fuca in 2007-2010, only three (Cakesota, Erin’s
Bride, Rounded) continued to support populations similar in size to those reported in 1978-1982 by Speich
and Wahl (1989) (see Appendix A).
Nine Washington colony sites are estimated to have once supported ≥1,000 puffins each, with two (Carroll,
Jagged) reaching 10,000 or more individuals (Appendix A). Of the eight currently occupied colonies
(Alexander, Bodelteh Islands [all], Cake, Carroll, Dhuoyautzachtahl, Jagged, Puffin Rock, Silver Sides) that
formerly held ≥1,000 birds, the maximum count at any of these sites in 2007-2010 was 211 individuals.

Future population status. Future population trends for Washington’s Tufted Puffins likely depend on a
range of local, regional, and global factors. Populations will be negatively affected by many of the
considerations detailed below under Factors Affecting Continued Existence, some of which are beyond the
control of local regulations and management practices. If the current 8.9% annual rate of decline continues,
the species could become functionally extirpated from Washington in about 40 years (based on a current
estimated population of 3,000-4,000 puffins falling to 100 birds).

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 Figure 2. Population (abundance or density) trends for Tufted Puffins in Washington from four time series
datasets. These include results from two boat-based surveys (Westport offshore surveys, 1972-2001
[Wahl and Tweit 2000; T. R. Wahl and B. Tweit, unpubl. data] and coastal offshore surveys from Cape
Flattery to Point Grenville, 2001-2012 [S. Pearson, unpubl. data]) and two colony attendance surveys
(Tatoosh Island, 1983-2001 [R. Paine, unpubl. data] and Protection Island, 1973-2001 [U. Wilson, unpubl.
data]).

Table 3. Changes in the numbers and sizes of breeding colonies of Tufted Puffins in Washington from
the late 1800s to 2007-2010.
Colony size (no. of birds)

1-99
100-999
1,000-9,999
≥10,000
Total no. of occupied sites

January 2015

1886-1977
24
13
6
1
44

No. of colonies per survey period
1978-1984
18
11
5
1
35

15

2007-2010
10
9
0
0
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 Figure 3. Locations and historical maximum population estimates for 44 Tufted Puffin breeding colonies
documented in Washington. Panels A-C depict colony occurrences along the outer coast from
northwestern Clallam County to northern Grays Harbor County during the periods from 1886-1977, 19781984, and 2007-2010. Panels D-F depict colony occurrences in the San Juan Islands and eastern Strait
of Juan de Fuca during the periods from 1886-1977, 1978-1984, and 2007-2010. Count data for each
breeding site are presented in Appendix A.

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 HABITAT STATUS
This discussion of habitat status in Washington is limited to current and former areas used during the
breeding season, since Tufted Puffins generally do not winter in state waters. The few birds visiting the state
in winter are strays from pelagic regions of the North Pacific. The status of their winter pelagic habitat is
poorly known, but that habitat is subject to large-scale trends in climate, ocean conditions, and pollutants
(e.g., plastic debris).

Past
Historically, Native Americans visited Tufted Puffin colonies in Washington both to hunt birds and gather
their eggs (Jones 1908a, Bovy 2007). Small islands with nesting puffins and other seabirds were also used for
the rearing of dogs (Paterek 1996) and for the collection of camas (Camassia spp.) and other food plants
(Turner 1999). Burning after the camas harvest was a common practice to reduce woody vegetation (Boyd
1999) and records indicate that Native Americans performed burns on at least two islands in Washington and
British Columbia that once held nesting puffins (Waldron Island, Washington, and Mandarte Island, British
Columbia; Turner 1999, Sprenger and Dunwiddie 2011). It is unclear whether or how puffin nesting habitats
were affected by these activities.
Following European settlement, several important Tufted Puffin colonies were adversely impacted by
construction, human habitation, farming, and other human disturbances. Tatoosh, Smith, and Destruction
islands all supported manned lighthouse stations for many decades, where activities included the development
and maintenance of landing sites, lighthouses, outbuildings, and other infrastructure. Protection Island was
extensively farmed and at one time platted and partially developed for an 800-lot vacation home project.
Flattop Island experienced use as a training range for warplanes during World War II, and other islands
received occasional visits from fishermen, naturalists, and the general public until well into the 20th century.
European Rabbits and dogs were introduced to several islands with nesting puffins, but impacts of these
species are poorly known except on Destruction Island, where overgrazing by rabbits appears to have
changed the island’s grassy vegetation and caused significant soil erosion and land slippage, which reduced
burrowing habitat for puffins (S. Pearson, pers. comm.). By the 1970s, virtually all former and active
breeding sites in Washington had been protected by the establishment of national wildlife refuges and
wilderness areas, where public access is much reduced or prohibited altogether.
Foraging areas for Tufted Puffins include vast tracts of outer coastal and inland marine waterways where the
birds disperse to feed and find fish to feed their young. These areas are governed by maritime laws and
fishing regulations, and more nominally by laws concerning seabird protection (Harrison et al. 1992).
Following European settlement, puffin foraging habitat in these areas was adversely impacted by fishing,
environmental contaminants, shipping, and increasing boat traffic.

Present and Future
Nearly all documented former and current breeding locations for Tufted Puffins in Washington are now
included in the Washington Maritime National Wildlife Refuge Complex and associated wilderness areas.
This group of refuges includes six distinct management units: the San Juan Islands, Protection Island,
Dungeness Spit, Cape Flattery, Quillayute Needles, and Copalis national wildlife refuges. Additionally, many
sites in the San Juan Islands are included in the San Juan Islands Wilderness. In most instances, light stations
have been automated or decommissioned and other human activities are greatly reduced at these sites.
Caretakers reside on Protection Island, and other islands receive periodic visitation from researchers and
refuge managers, but the amount of human disturbance at current Washington breeding colonies is low and
likely to remain so. Some of the rocks and islands formerly occupied by puffins in the San Juan Islands are
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 not regularly patrolled and therefore may be subject to recurring human disturbance. Rabbit control
measures have been proposed for Destruction Island, and current refuge management plans include an
emphasis on promoting native plants and animals (e.g., USFWS 2007, 2010).
With shipping and recreational traffic on the rise, foraging habitat continues to be affected by vessel traffic,
particularly in the inland marine waters. Oil spills and other environmental contaminants remain a threat to
these habitats, and diminished populations of forage fish have likely reduced habitat quality in some areas.
Commercial fishing in Washington has declined sharply in recent decades. Though no Tufted Puffin
entanglements have been reported recently, Common Murres, Rhinoceros Auklets, and other diving seabirds
continue to drown in nets, albeit in reduced numbers (e.g., Hamel et al. 2009). Derelict fishing gear also
continues to be a threat in some areas (Good et al. 2009, 2010), but is currently the subject of a clean-up
effort (see www.derelictgear.org) with positive early results (June and Antonelis 2009).

CONSERVATION STATUS
Federal. The U.S. Fish and Wildlife Service (USFWS) considers the Tufted Puffin a species of concern, a

non-regulatory designation intended to encourage conservation action before situations become more serious.
Tufted Puffins are protected under the 1918 Migratory Bird Treaty Act (and subsequent amendments), which
prohibits the taking, killing, possession, transportation, and importation of migratory birds, their eggs, and
their nests. Federal management of seabirds falls under the jurisdiction of the USFWS, and certain
exceptions to the law that allow interaction with migratory birds (e.g., scientific, cultural) are governed by
strict permitting requirements. In general, the law is more strictly enforced with regards to nesting colonies,
but less so in terms of the “taking” of puffins as bycatch from fishing activities (Harrison et al. 1992). The
Fish and Wildlife Conservation Act of 1980 is also pertinent to Tufted Puffins, requiring the USFWS to
assess migratory nongame bird populations, determine the effects of human activities, and identify
populations likely to become candidates for listing under the Endangered Species Act. An Executive Order
implemented in 2001 (Executive Order 13186 - Responsibilities of Federal Agencies To Protect Migratory
Birds) essentially expands this mandate to every federal agency whose actions are likely to impact migratory
bird populations, requiring them to enter into a Memorandum of Understanding for management and
monitoring purposes with the USFWS.
National marine sanctuary (NMS) regulations (15 CFR 922 Subpart O, 152(a)), which apply to the Olympic
Coast NMS off the outer northwest coast of Washington, contain prohibitions on the taking and possessing
of any seabird in the sanctuary, except as authorized by the Endangered Species Act and the Migratory Bird
Treaty Act. The regulations also prohibit the disturbance of seabirds by aircraft flying below 610 m over
waters within 1.85 km of Flattery Rocks, Quillayute Needles, and Copalis national wildlife refuges or within
1.85 km seaward from the coastal boundary of the sanctuary, with certain exceptions.
In February 2014, the USFWS was petitioned to list Tufted Puffin populations in Washington, Oregon, and
California under the federal Endangered Species Act (NRDC 2014). Because of other listing priorities, the
USFWS will not begin its evaluation to list the species in these states until 2016 or 2017 (D. Lynch, pers.
comm.). This means that any listing decision would probably not be reached until sometime between 2018
and 2020.

Washington. Tufted Puffins are covered under several Washington laws and regulations. The species is
protected under the category of “other protected wildlife” in the Washington Administrative Code (WAC
232-12-011; Appendix B). This prohibits the hunting, malicious killing, possession, and malicious destruction
of Tufted Puffins and their eggs and nests, but does not protect the species from harassment. Violations of
this law are a misdemeanor offense (RCW 77.15.130; Appendix B), with penalties ranging up to 90 days
imprisonment, a $1,000 fine, or both. The species also receives protection under WAC 232-12-064, which
prohibits the capture, importation, possession, transfer, and holding of most wildlife in the state.
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 The Washington Department of Fish and Wildlife (WDFW) has considered the Tufted Puffin a candidate for
state listing since October 1998, though this designation does not infer any specific legal status or protections.
Audubon Washington considers Tufted Puffins as an “early warning” species of concern, and one at “high
risk” of negative impacts from changes in ocean and atmospheric circulation related to climate change
(Audubon Washington 2004, 2009).

Other states. Tufted Puffins are listed by the California Department of Fish and Wildlife as a Priority 1 Bird
Species of Special Concern (S2; McChesney and Carter 2008). The Oregon Department of Fish and Wildlife
lists Tufted Puffins as Sensitive - Vulnerable, while the Oregon Natural Heritage Program ranks the breeding
population as Critically Imperiled (S1B; ORBIC 2013). In Alaska, the species is considered Apparently
Secure (S4; NatureServe 2014) and is not listed by the state.

International. In Canada, Tufted Puffins are protected from harvest by the federal Migratory Birds
Convention Act and the province of British Columbia’s Wildlife Act. They are not listed under the Species at
Risk Act (federal legislation). The province has designated the breeding population of the species as being of
Special Concern (S3B) and includes it on its Blue List, meaning that this population is sensitive or vulnerable
to human activities or natural events (B.C. Conservation Data Centre 2014). However, the province’s overall
puffin population (including wintering birds) is considered Apparently Secure (S4N). In Japan, Tufted
Puffins have been listed as a national and local endangered species since 1993 (Osa and Watanuki 2002).

RESEARCH, MONITORING, AND RESTORATION
Research. Research on Tufted Puffins dates back decades and covers a wide range of behavioral and

ecological subjects. Breeding biology, diet, foraging habitats, and ecological relationships are now fairly well
understood (Piatt and Kitaysky 2002), and new tools are opening up additional opportunities to refine
existing knowledge. Stable isotope analysis has revealed the subtleties of seasonal dietary shifts (e.g., Davies
et al. 2009), while population and ecological genetic tools developed for closely related alcids (Rhinoceros
Auklet, Whiskered Auklet) show great promise for use in Tufted Puffins (Dawson et al. 2005, Hasegawa et al.
2005). Challenges to puffin research include the inaccessibility of many breeding sites, the dispersed nature of
puffin populations in winter, and the sensitivity of the birds to research-associated disturbance. Studies have
shown changes in adult and chick behaviors during nestling research (e.g., Hatch et al. 2000 a, 2000b,
Gjerdrum et al. 2006), and have found that the presence of observers can negatively affect fledging success
and nestling growth rates (Pierce and Simons 1986). Attempts to surgically implant satellite transmitters in
Tufted Puffins have met with behavioral alteration (nest abandonment) and high mortality, and the
attachment of small radio-transmitters has negatively impacted reproductive success (Whidden et al. 2007).
Further refinements are necessary for these techniques to be effective and safe (Hatch et al. 2000a, 2000b).

Research needs. Though Tufted Puffins are considered one of the better-studied alcids, many important
research questions remain. No data exist on gene flow or the genetic and ecological distinctiveness among
populations, which are significant considerations for conservation efforts for populations near the southern
edges of the species’ range. Similarly, puffin distribution and habits during the winter months are largely
unknown, factors that may take on increased significance if ocean conditions follow the trends predicted by
climate change (e.g., Hoegh-Guldberg and Bruno 2010, Doney et al. 2012). Habitat use, diet, and survival at
sea remain essentially unmonitored and there is no information on whether winter habits differ among
colonies or regions, which could be particularly important for declining populations like those in Washington.
More precise data on foraging range and habitat use during the nesting season would also be of great use in
identifying effective management strategies for the species. In Washington, specific information on dietary
habits and the relationships between breeding success and trends in forage fish remains largely speculative.
Long-term, location-specific data of the type being amassed at Triangle Island, British Columbia, would be
extremely pertinent for the management and conservation of Washington colonies, and it would be helpful to
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 analyze the relative influence of both top-down (e.g., Bald Eagle predation) and bottom-up (e.g., prey
availability) factors affecting the population. Additionally, more precise time series data on population trends
would be helpful and a thorough assessment of threats to Washington’s sites (e.g., introduced species) would
help set management priorities. The extent and impacts of gull kleptoparasitism at Washington colonies
should be investigated, particularly in light of increasing abundance of gulls.

Surveys and monitoring. Tufted Puffins are currently included in long-term monitoring of seabird colonies

in Alaska (Dragoo et al. 2010) and at Triangle Island, British Columbia (M. Hipfner, pers. comm.).
Expansion of such surveys and standardization of protocols have been called for (USFWS 2005) and would
be particularly relevant for Washington and other areas where the species is declining. Regular surveys or
monitoring programs specific to Tufted Puffins have never been conducted in Washington. The
investigation by P. Hodum et al. in 2007-2010 was the first statewide attempt to assess the status of puffin
breeding colonies in 25 years. The species is counted on the water during annual boat-based surveys focused
on Marbled Murrelets (S. Pearson, pers. comm.). It also has been surveyed during the Westport Seabirds
trips traveling west out of Westport, Washington, since 1972, although a change in survey protocols has
prevented meaningful analyses of count data since 2002. The species is also recorded during annual boatbased surveys in the San Juan Islands National Wildlife Refuge.

Restoration potential. Restoration of Tufted Puffins at current and former breeding sites in Washington

was suggested in the wake of the 1991 Tenyo Maru oil spill, but never acted upon (Tenyo Maru Trustees 2000).
Though stabilizing and increasing populations at current breeding sites should receive first priority,
restoration may offer a future means of expanding breeding opportunities and buffering the species against
decline. Restoration has never been attempted with Tufted Puffins, but the successful reintroduction of
Atlantic Puffins to islands off the coast of Maine (Kress and Nettleship 1988) indicates that it may be
possible. In a worldwide review of seabird restoration, Jones and Kress (2012) found a 60% success rate for
alcid species. Recent colonization of Mandarte Island, British Columbia, by Rhinoceros Auklets (P. Arcese,
pers. comm.) and increased numbers of forage fish in recent trawls in the San Juan Islands (K. Fresh, pers.
comm.) suggest that conditions around some former nesting sites in Washington may be opportune for either
natural re-occupation or reintroduction. Site selection would require careful analysis, however, to avoid
encouraging puffins to reoccupy poorly suited sites.

Projects to restore natural habitat conditions at current and former nesting islands should also be undertaken
or evaluated to conserve Tufted Puffins in Washington. These would include the removal of invasive plants
and animals (see Factors Affecting Continued Existence – Introduced Species) and the replanting of native
vegetation. Eradication of European Rabbits on Destruction Island is already under consideration (S.
Pearson, pers. comm.) and control of non-native plants at Protection Island has been proposed (USFWS
2010).

FACTORS AFFECTING CONTINUED EXISTENCE
The causes of the Tufted Puffin decline in Washington remain undetermined because of a lack of scientific
study. Throughout its range, the species is adversely impacted by a wide range of natural and human factors.
On the water, threats include net fisheries, oil spills, ingested plastics, and bioaccumulation of chemical
pollutants, whereas breeding colonies are at risk from disturbance, predators, introduced species, disease
outbreaks, and habitat loss (Tyler et al. 1993, Mahaffy et al. 1994, Piatt and Kitaysky 2002, Gill 2007).
Reduced prey availability ranks as a major concern, particularly in combination with ocean surface
temperature trends predicted by global climate change (Gjerdrum et al. 2003). The puffin habit of nesting in
aggregations makes them particularly vulnerable to many of these threats (Gill 2007), while small, declining
populations like those in Washington face increased risk of extirpation from stochastic events like disease
outbreaks or oil spills (Donald 2010). These and other threats are discussed in detail below, set in the context
of existing regulatory mechanisms and with particular attention to conditions in Washington.
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 Adequacy of Existing Regulatory Mechanisms
Federal protection. Federal laws and regulations relevant to Tufted Puffins help protect the birds from
harassment and hunting (see Conservation Status), but are unevenly applied, particularly with regard to
bycatch from commercial fishing (Harrison et al. 1992). The U.S. Fish and Wildlife Service has
recommended several research and management initiatives for the species to be conducted by the agency and
other entities, including the development of protocols for long-term monitoring and the installation of
observers on commercial fishing vessels (USFWS 2005, 2009). There are no federal regulations that require
proactive management steps, however, which could be important for declining populations like those in
Washington.
State protection. Current state protections for Tufted Puffins resemble those offered by federal law, but do

not protect the species from harassment (see Conservation Status). Results of this listing process could
afford Washington’s puffin population some additional legal protection under RCW 77.15.120 (Appendix B).
Classification as a state endangered species would (1) protect puffins from malicious harassment, (2) increase
the legal penalties associated with unlawful taking, (3) result in the preparation and implementation of a state
recovery plan having conservation actions that benefit the species, (4) make puffins a high priority
conservation target within WDFW, and (5) bring greater public recognition to the imperiled status of the
species in Washington.

Reduced Prey Availability
Reductions in prey abundance and the timing of prey availability rank as major concerns for Tufted Puffin
populations in Washington and elsewhere. A recent study found widespread reproductive failure in 14
seabird species (including Atlantic Puffins) when forage fish and krill populations were depleted below onethird of their observed maximum (Cury et al. 2011). Data on the status of prey in the puffins’ winter range is
limited, so the discussion below focuses on prey availability during the critical breeding period, when the birds
reside in Washington waters. Because the timing and abundance of prey populations are often dependent
upon ocean temperature and climatic conditions, this threat should be considered closely intertwined with
ocean and climate forcing factors.

Forage fish. Populations of forage fish can vary widely in response to large-scale changes in oceanic

conditions (e.g., El Niño, La Niña, decadal oscillations) (Baird 1990, Brodeur et al. 2005, Lindegren and
Checkley 2013, Santora et al. 2014), as well as fishing or other human pressures (e.g., Ainley and Lewis 1974).
Seabird abundance and reproductive success correlate closely with prey availability around nesting colonies,
and numerous studies have linked Tufted Puffin breeding success with the availability of forage fish in nearby
waters (e.g., Vermeer et al. 1979, Baird 1990, Golubova 2002, Gjerdrum et al. 2003). Nestling body mass,
growth rates, fledging rates, and the timing of the first molt have all been linked to quantity and quality of fish
species that adults provide their offspring (Vermeer and Cullen 1979a, Thompson et al. 2001). Reproductive
failure, on the other hand, has been noted in years of forage fish shortage (e.g., Vermeer et al. 1979), or when
the timing or size of prey fail to match the parents’ foraging requirements (Golubova 2002). Similarly, the
type of forage fish available has been correlated to chick growth and fledging success in the closely-related
Rhinoceros Auklet (e.g., Hedd et al. 2006, Thayer and Sydeman 2007), which may indicate that birds have a
particular efficiency for assimilating nutrition from their preferred prey (Niizuma and Yamamura 2004).
Though there have been no studies in Washington specifically focused on Tufted Puffin diet or the
relationships between puffins and forage fish, several inferences can be made from available data on fish
populations. During the breeding season, puffins generally feed themselves and provision their young with
fish of high nutritional quality within a 20 to 200 mm size range. In Washington, these would include several
well-documented puffin prey species: Pacific Herring, Pacific Sand Lance, juvenile rockfish, Eulachon, Pacific
Sardine, and Northern Anchovy. Additional species within the preferred size range include Surf Smelt, Night
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Washington Department of Fish and Wildlife

 Smelt, Popeye Blacksmelt, Whitebait Smelt, juvenile Kelp Greenling, Pacific Saury, and juvenile salmon.
Catch records, survey data, and other observations suggest that some of these species experienced significant
population declines and/or loss of habitat in Washington during the 1900s through to the present, as follows:
•

•

•

•

Pacific Herring – A number of herring stocks in the Georgia Basin are comprised of sizable migratory
components that move to feeding grounds off the continental shelf along the outer Washington coast
from March to July (Stick and Lindquist 2009), where they become available as prey for Tufted Puffins.
Of 19 herring stocks in greater Puget Sound, only nine (47%) were considered healthy or moderately
healthy in the latest analysis (Stick and Lindquist 2009), the lowest number of healthy stocks since
tracking began in 1994. The herring population at Cherry Point, once considered Washington’s largest,
suffered a 95% decline between 1973 and 2000 and has shown little sign of recovery (Gustafson et al.
2006, Stick and Lindquist 2009). Recent declines in herring spawning have also been noted in the Strait
of Georgia (Therriault et al. 2009). However, some relatively healthy stocks do remain in Puget Sound
and elsewhere in the Georgia Basin (Gustafson et al. 2006, DFO 2014) and it’s likely that both resident
and migratory herring remain an important food source for Tufted Puffins in Washington. Spawning
populations of herring also exist in Willapa Bay and Grays Harbor (Penttila 2007), but no long-term data
are available to establish abundance trends.
Pacific Sand Lance – Sand lance constitute a major part of puffin diets in British Columbia and Alaska
(summarized in Piatt and Kitaysky 2002) and are known to be an important potential food source of
other alcids in Washington (Leschner 1976, Wilson and Manuwal 1986, Schrimpf et al. 2012, Pearson et
al. in review). Though sand lance population trends have not been monitored in Washington, the species
is known to spawn in upper intertidal areas vulnerable to shoreline alterations (Penttila 2007). The
impacts of shoreline modifications (e.g., armoring, nearshore fill, wetland loss, and the construction of
breakwaters, jetties, and overwater structures) on sand lance abundance in Washington are unknown, but
it is safe to say that spawning habitat has declined in extensively developed areas, such as Puget Sound.
However, in the areas historically occupied by Tufted Puffins, shoreline modifications have been
relatively modest (in the Strait of Juan de Fuca and San Juan Islands; Simenstad et al. 2011) or minimal
(the outer Washington coast), thus sand lance populations in these regions may have remained relatively
stable compared to historical levels.
Rockfish – Juvenile rockfish can be a significant percentage of adult and nestling Tufted Puffin diets
during the breeding season (e.g., Vermeer 1979), and are known to be an important food source for the
closely related Rhinoceros Auklet at coastal sites in Washington (Wilson and Manuwal 1986). Rockfish
populations in the interior waters of Washington have declined markedly over the past three decades,
with 22% of stocks considered vulnerable or depleted, and 56% considered precautionary (reduced but
apparently stable, or information lacking) (Palsson et al. 2009). Additionally, seven coastal rockfish
stocks are considered depleted and have been put under severe fishing restrictions in both federal and
state jurisdictions (Palsson et al. 2009). Thirteen rockfish species in Washington’s inner waters are federal
species of concern and three were listed as endangered or threatened under the Endangered Species Act
in 2010. Thirteen rockfish species are designated as state candidates for listing (WDFW 2014).
Overharvest has been a primary factor in these declines. While the species composition of the “rockfish”
component of puffin diets has not been established, decreased adult abundance and stochasticity in
reproductive success, perhaps exacerbated in part by climate change, have very likely made juvenile
rockfish a less abundant food source for Tufted Puffins in Washington.
Eulachon – Precipitous declines in Eulachon (also called Columbia River Smelt or Pacific Smelt) led the
National Marine Fisheries Service to declare populations throughout Washington, Oregon, California,
and British Columbia as threatened under the federal Endangered Species Act in 2010 (NMFS 2009,
2010, ODFW/WDFW 2014). Two major production areas for the species occurred in the Columbia and
Fraser rivers and would once have been prolific potential food sources for puffins, but Eulachon
populations in both areas have greatly declined since 1993 (NMFS 2009). Minor improvements in the
Columbia River population may have occurred since 2011 (ODFW/WDFW 2014).

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 •

•

Pacific Sardine – Sardines were among the most abundant forage fish off the outer Washington coast in
the early 1900s and supported a large commercial fishery until stocks collapsed in the 1930s and 1940s
coincident with a change in ocean conditions and high harvest levels (Bargmann 1998, Zwolinski and
Demer 2012). The species made a partial comeback in the 1990s (Emmett et al. 2005), which has
supported a limited commercial harvest since 2000. Sardine abundance along the North American west
coast has shown several fluctuations since 2000, but has generally displayed a downward trend during this
period (Hill et al. 2011), including the disappearance of stocks off Vancouver Island in 2013. Though the
impacts of these population fluctuations on Washington’s Tufted Puffins are unknown, the mid-20th
century crash in sardine stocks has been suggested as a factor in coincident puffin declines in California
(Ainley and Lewis 1974).
Northern Anchovy – The historical abundance of anchovies off the outer Washington coast is poorly
known (Bargmann 1998), although the species typically is associated with warmer water temperatures and
weaker upwelling, which are ocean conditions unfavorable for Pacific Sardines and some other forage
fish (Chavez et al. 2003, Santora et al. 2014). Anchovy biomass off Washington declined during the
1980s, then increased during the 2000s (Litz et al. 2008; L. Wargo, pers. comm.). Stocks currently
support a limited commercial fishery primarily for the bait trade. Anchovies have also been considered
abundant at times in Puget Sound (Bargmann 1998). Studies at Destruction Island note anchovies as a
vital food source for nesting Rhinoceros Auklets (Wilson and Manuwal 1986, Pearson et al. in review),
and they are likely an important prey species for puffins as well. In Californian waters, anchovies
function as a primary food item of seabirds during periods of decline among other prey species (Santora
et al. 2014).

Natural fluctuations caused by changing ocean conditions are a major influence on the abundance of forage
fish in the areas of Washington inhabited by Tufted Puffins. Additionally, existing data and historical
accounts make it clear that the availability of some populations of forage fish (especially Pacific Herring,
Eulachon, and rockfish) used by Tufted Puffins in Washington has declined beyond what might be expected
due to changes in ocean conditions (Gustafson et al. 2006, NMFS 2009, 2010, Palsson et al. 2009, Stick and
Lindquist 2009, ODFW/WDFW 2014). Past overfishing has played a role in the declines of herring,
rockfish, and perhaps other species (Gustafson et al. 2006, Palsson et al. 2009, Stick and Lindquist 2009).
Commercial fisheries currently exist for herring, Surf Smelt, sardines, and anchovies (largely inactive), but are
much reduced from their historical peaks and, barring major increases in harvest, are likely to have only a
limited role in reducing current prey availability for Tufted Puffins. Pelagic fisheries for sardines and
anchovies are managed by the Pacific Fishery Management Council with a priority for maintaining their
ecosystem functions, primarily as a food source for seabirds, marine mammals, salmon, and other predators
(PFMC 2011).
Loss or degradation of key habitats and declining water quality are other major concerns for forage fish
populations, especially in highly developed regions like Puget Sound, where there is extensive industry,
agriculture, and urban growth, and where widespread shoreline armoring has greatly reduced intertidal
spawning habitats for species like Surf Smelt, Night Smelt, and Pacific Sand Lance (Thom et al. 1994, Rice
2006, Krueger et al. 2010). However, shoreline modifications and other coastal development have been
minimal along the outer Washington coast and relatively modest in the Strait of Juan de Fuca and San Juan
Islands (e.g., Simenstad et al. 2011).
In addition to the quantity, and thus availability, of forage species, prey quality also plays a role in the
breeding success of Tufted Puffins. The nutritional value of potential Tufted Puffin prey varies widely
among species and among age classes of the same species (Anthony and Roby 1997). Dietary shifts to less
nutritious species reduce nestling growth rates in captive studies (Romano et al. 2006) and have been
implicated in declines in the wild (e.g., Piatt and Anderson 1996). Recovery of seabirds, including Tufted
Puffins, in the wake of the Exxon Valdez oil spill in Alaska was thought to be limited by the availability of
high quality forage fish (sand lance, herring, and capelin; Anthony and Roby 1997). Studies of Rhinoceros
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 Auklets, for example, show high breeding performance when Pacific Sand Lance predominate the diet fed to
chicks (e.g., Wilson and Manuwal 1986, Hedd et al. 2006, Borstad et al. 2011). During warm water events,
when sand lance numbers generally decline, juvenile Pacific Saury often become more abundant in the auklet
diet, resulting in reduced chick growth rates (Thayer and Sydeman 2007) and fledging success (Hedd et al.
2006). Timing of reproduction that may lead to reductions or changes in the availability of appropriately
sized prey for nestlings can also be a threat. Golobuva (2002) observed mass starvation of Tufted Puffin
nestlings in the Sea of Okhotsk in a year when herring were abundant, but of a size class too large for the
chicks to ingest.

Marine invertebrates. No time series data exist for the populations of squid and large zooplankton that
likely make up part of the Tufted Puffin diet in Washington. A critical winter food source, invertebrates can
also be important adult prey items during the early breeding season (Davies et al. 2009), and some individuals,
both breeders and non-breeders, appear to use them throughout the summer (Piatt and Kitaysky 2002).
Though local population trends remain unstudied, zooplankton blooms in the Northeastern Pacific respond
directly to ocean surface temperatures and other climatic trends, and are in turn a primary factor controlling
fluctuations in forage fish numbers (e.g., Mackas et al. 2007). In general, increases in ocean temperature
result in declining zooplankton biomass and changes in community composition of zooplankton (King et al.
2011), a pattern that has already been documented for euphausiids. Research on Cassin’s Auklets found that
warm water years altered the productivity and timing of critical euphausiid species with direct negative
consequences for nestling survival and fledging weight (Bertram et al. 2001, Hipfner 2008, 2009). The
availability of marine invertebrates is probably closely linked to the trends in climate and ocean temperatures
discussed below.

Climate Change Effects
The locations of breeding colonies used by Tufted Puffins may make them particularly vulnerable to the
effects of climate change. Shifting weather patterns and rising sea temperatures predicted by climate change
models could have a dramatic impact on the productivity and occupancy of established puffin colonies, as
well as the establishment of new ones.

Warming ocean and prey availability. Climate change is expected to cause a long-term rise in global ocean
surface temperatures (Morgan and Siemann 2011, IPCC 2013), a trend generally associated with lower ocean
productivity (e.g., Behrenfeld et al. 2006), altered availability of plankton (Hipfner 2008, Batten and Walne
2011, King et al. 2011), and corresponding changes in high-trophic communities like seabirds (e.g.,
Hyrenbach and Veit 2003, Frederiksen et al. 2013). Increases in ocean surface temperature have already been
detected in much of the North Pacific (Thompson et al. 2012). Predicted effects for this region are widely
expected to include altered food webs and faunal distributions (e.g., Fields et al. 1993, Di Lorenzo et al. 2008,
Abdul-Aziz et al. 2011), and recent fluctuations in temperature and climate have been linked to prey
availability and breeding success in alcids (e.g., Francis et al. 1998, Gjerdrum et al. 2003, Hedd et al. 2006,
Hipfner 2008, 2009). These relationships have been called the most important factor in determining the
viability of Tufted Puffin populations (Gjerdrum et al. 2003).
Previously, periodic short-term temperature increases have been linked to lower productivity, shifts in prey
communities, and corresponding declines in seabird abundance, particularly for diving pursuit feeders like
puffins (Ainley and Hyrenbach 2010, Miller et al. 2013). In general, piscivorous birds are thought to have
greater breeding success during cold water regimes, when ocean productivity is higher and more forage fish
are available to them (Hunt et al. 2002). Rises in ocean surface temperatures during El Niño events have
been associated with widespread nesting failures for Tufted Puffins and other alcids (e.g., Lowe 1993, Piatt
and van Pelt 1993), though adult survival can remain high during these periods (Morrison et al. 2011).
Following a decadal shift in ocean temperatures and cycling in the 1970s, forage fish declined throughout the
Northeastern Pacific (Francis et al 1998). This change has been implicated in corresponding declines of
several piscivorous seabirds (Francis et al. 1998), including Tufted Puffins (Agler et al. 1999). Anderson and
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 Piatt (1999) also noted widespread declines in Tufted Puffins and other piscivorous seabirds in the Gulf of
Alaska following a shift to warmer ocean temperatures, which corresponded with fewer prey (capelin and
shrimp). Whether these changes represented a reduction in the overall puffin population or a shift in
distribution is unclear, but the link between local numbers and ocean temperatures appears strong.
Relationships between puffins, climate, and ocean temperature have been studied intensively at Triangle
Island, British Columbia, where conditions resemble those of Washington’s outer coast. Gjerdrum et al.
(2003) found that hatch date, chick growth, and fledging success were all affected by ocean temperature, and
there appeared to be upper and lower temperature thresholds outside of which breeding success was reduced
essentially to zero. The mechanisms behind these results have not been determined, but are likely related to
reduced copepod abundance in warm, nutrient-poor surface waters, which in turn lead to reduced forage fish
populations around nesting colonies (Gjerdrum et al. 2003). This can occur in response to widespread trends
in ocean surface temperature, or when the local upwelling of cold, nutrient-rich waters is poor or late. Links
between ocean temperatures, prey availability, and nesting failures have been noted previously at Triangle
Island (Vermeer et al. 1979), as well as for Tufted Puffin colonies in Alaska (Agler et al. 1999) and Russia
(Golubova 2002), and for Atlantic Puffins in Norway (Durant et al. 2003).
Though most studies find puffins faring poorly in warm water regimes, Kitaysky and Golubova (2000)
documented increased reproductive success during high surface temperature periods in Tauyskaya Bay,
Russia. They postulated that in that system, abundant meso-zooplankton in warm water attracted larger
numbers of pelagic forage fish into the bay. It may be that in some cases local effects are ultimately more
important than global and regional trends in the fate of individual colonies. At Triangle Island, for example,
recent research on Rhinoceros Auklets suggests that large-scale weather patterns greatly complicate the link
between ocean temperatures and the availability of sand lance (Borstad et al. 2011). The timing of the annual
spring shift from winter to summer wind patterns determines when deep-water upwelling will bring planktonrich waters to the island (Borstad et al. 2011). If winter storms delay that transition beyond a critical period,
the plankton bloom is poor or late, juvenile sand lance fare poorly, and the birds that feed on the sand lance
suffer accordingly. A large-scale study of Cassin’s Auklets at sites from British Columbia to Baja California
also identified highly local conditions, rather than regional or global trends, as the best predictor of colony
breeding success (Wolf et al. 2009).
Thompson et al. (2012) recently reported an increased occurrence of Tufted Puffins at sea during February
cruises (but not during those in June and August-September) along a 1,425-km transect west of Vancouver
Island. These results suggest the possibility of regional shifts in puffin wintering distribution in the North
Pacific in association with warming ocean surface temperatures.

Sea level rise. Sea level rise caused by climate change is expected to negatively impact the breeding and
foraging habitats and prey populations of many coastal waterbird species (Clausen and Clausen 2014). For
Tufted Puffins, it may affect the abundance of some forage fish species by altering the intertidal and subtidal
habitat conditions (e.g., water depths) preferred for egg deposition and refuge (Penttila 2007, Krueger et al.
2010). This problem may be especially acute where shoreline armoring has reduced or eliminated the ability
of the beach face to erode landward, a situation referred to as “coastal squeeze” (Glick et al. 2007, Krueger et
al. 2010). Physical attributes of beaches within a given locale may vary significantly, leading fish to deposit
proportionally more eggs on “preferred beaches” (Quinn et al. 2012). As a result, losses of beach faces at
specific tidal heights at a relatively small number of spawning beaches might have a large negative effect on
egg abundance (Quinn et al. 2012).
Sea level rise, combined with increasing severity of storms, also has the potential to damage or destroy some
Tufted Puffin colonies through increased erosion of soil and unconsolidated bluffs used as nesting areas
(Miller et al. 2013). Examples of sites in Washington that may be vulnerable to this threat include Protection,
Smith, and Destruction islands (S. Pearson, pers. comm.). Jewett et al. (1953) previously described the loss of
occupied nesting burrows on Smith Island caused by wave erosion.
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 Ocean acidification. In addition to its predicted impacts on ocean temperatures and weather patterns,
climate change is also expected to alter the acidity of seawater, particularly at higher latitudes where Tufted
Puffins reside. As the ocean absorbs increased levels of atmospheric carbon dioxide (CO2) from human
activities and natural processes, acidity will rise and impair the ability of marine organisms to form calcareous
shells and skeletal structures (Fabry et al. 2008). This change could shift planktonic and benthic communities
away from calcium-dependent species, a low-trophic re-ordering with unpredictable but potentially profound
implications for marine food webs (Fabry et al. 2008, Branch et al. 2013). Where rising ocean surface
temperatures are expected to lower the abundance of zooplankton (and the forage fish that consume them),
rising acidity may change the composition of these communities as well (Roemmich and McGowan 1995).
The consequences for high trophic feeders like Tufted Puffins remain unknown.
Other effects. The interplay between changing climate, ocean conditions, plankton blooms, and forage fish
creates a dynamic environment with significant implications for Tufted Puffin reproduction. The prediction
that climate change will bring increased frequency of high intensity storms (IPCC 2013) is also a
consideration for Tufted Puffins during the breeding season, when fledglings have been known to die by the
thousands from severe weather and wave action (Reagan 1910). Some experts also predict climate change
will increase the frequency of harmful algal blooms, or “red tides” (Peperzak 2003), which can have a range of
negative effects on seabirds (see below).

Environmental Contaminants
Oil spills. Oil spills have long been considered one of the major anthropogenic threats to Tufted Puffins
(Ainley and Lewis 1974, Tyler et al. 1993, Piatt and Kitaysky 2002). Alcids often make up the majority of
seabirds killed during spills, and may also suffer when spills adversely affect their prey populations (Irons et
al. 2000). Most relevant to Washington is the particular vulnerability of Tufted Puffins during spring and
summer when their populations are concentrated around breeding colonies. The Exxon Valdez oil spill in
1989, for example, occurred in March when puffins and other alcids were just arriving at breeding sites. It
killed an estimated 250,000 seabirds, including as many as 13,000 Tufted Puffins (Piatt and Ford 1996,
Denlinger 2006). Another well-documented case occurred in Washington and adjacent British Columbia, at
the entrance to the Strait of Juan de Fuca. The 1991 Tenyo Maru oil spill took place in July at the height of the
breeding season, less than 30 km from the Tufted Puffin colony at Tatoosh Island. Oil slicks affected
Tatoosh and other shorelines all along the Washington coast, killing an estimated 9% of the statewide puffin
population (Tenyo Maru Trustees 2000). Because spills are more likely to occur during severe winter
conditions, puffins are probably less vulnerable during this period because of their pelagic distribution. For
example, a February 1986 spill off the coast of California resulted in thousands of alcid deaths, but Tufted
Puffins were only a minor component (Page et al. 1990).
In addition to the risk oil presents to birds at sea, it is important to consider that an oil spill need not reach
the islands where nesting colonies occur to inflict damage to eggs and nestlings. Oil encountered by foraging
adults at sea up to 100 km from the colony can be carried back to the nest, where eggs exposed to as little as
5 µl of oil can experience damage to developing embryos (Albers 1978, King and Lefever 1979).
The number and volume of oil spills increased markedly in the North Pacific during the later decades of the
20th century, corresponding with increased oil development, shipping, and industrialization of the Pacific
Rim (Burger and Fry 1993). As a shipping and oil-refining hub, Washington lies at the heart of this
commercial vessel traffic and experienced six major oil spills ranging from 0.1-2.3 million gallons along the
outer coast and the Strait of Juan de Fuca between 1964 and 1991 (Neel et al. 2007). Increased safety
measures and prevention programs since the 1990s have helped reduce the number and scale of vessel spills
globally (Anderson et al. 2012, Ramseur 2012), as well as in Washington, where no spills exceeding 100,000
gallons have occurred since 1991 (Etkin and Neel 2001, Neel et al. 2007). However, the sheer volume of
shipping traffic makes oil spills a persistent threat in the state. Shipping routes for major ports in Seattle,
Tacoma, and Vancouver, B.C., as well as several major oil refineries and the third largest naval base in the
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 U.S., all traverse waters near the largest puffin colonies in Washington (Serra-Sogas 2008). Marine vessel
transits in Washington numbered 7,100 in 2013, with hundreds of tank ships and tanker barges annually
transporting more than 15 billion gallons of crude oil, fuel, and other chemicals (Etkin and Neel 2001, Puget
Sound Action Team 2005, Neel et al. 2007, WSDOE 2014). Tanker traffic from ports in British Columbia
and possibly Washington, as well as oil transport by train, is expected to increase greatly in the next several
decades as oil and natural gas production expands in the interior of North America and as offshore oil and
gas development off Vancouver Island likely begins (NMFS 2013, O’Neil 2013). This is expected to increase
the risk of spills in Washington (Van Dorp and Merrick 2014). Barges, freighters, container ships, ferries,
naval vessels, and large fishing and recreational craft also carry oil and fuel in volumes large enough to
produce a significant spill. Places where spills are most likely to occur include the Strait of Juan de Fuca and
the outer coast, where puffin breeding and foraging sites are concentrated.
Among the safety measures instituted to prevent marine oil spills in Washington since the 1990s is the
establishment of an Area to Be Avoided (ATBA) off the northwestern coast, which encourages large vessels
to stay well offshore during transit along the coast (WSDOE 2014). Additional risk mitigation is provided by
a rescue tug stationed in Neah Bay that is able to respond to vessels with impaired maneuverability near the
entrance to the Strait of Juan de Fuca. Use of single-hull tanker vessels, including barges, was completely
phased out in the U.S. in January 2015.
Although lower in profile, the chronic discharge of waste oil at sea (e.g., from illegal releases of oily bilge
water and tank flushings) represents another pervasive and continuing threat to puffins (e.g., Jewett et al.
1953). Mortality from oil discharged in shipping lanes was implicated as a factor in the early 1900s decline of
Tufted Puffins at the Farallon Islands, California (Ainley and Lewis 1974). Lighthouse keeper logs and
naturalist reports from the period note that oiled puffins and other birds were a common occurrence
(Dawson 1911, Ainley and Lewis 1974). Near Westport, Washington, small numbers of Tufted Puffins were
found with bilge oil on their feathers in 1959 (Alcorn 1959). Surveillance of small-volume oil discharges from
1997 to 2006 in British Columbia indicates that the Strait of Juan de Fuca (including the entrance and the area
around Cape Flattery, Washington) and Canadian waters near the San Juan Islands are areas of relatively high
chronic oil pollution and some associated seabird loss (Serra-Sogas et al. 2008, O’Hara et al. 2009).

Ingested plastics. Plastic pollution has steadily increased throughout the North Pacific in recent decades

(e.g., Day and Shaw 1987, Robards et al. 1995), with some forms of plastic being most common in coastal
waters (e.g., microplastics; Desforges et al. 2014). Among alcid species, those that feed extensively on
zooplankton (i.e., puffins, auklets) may be most at risk of consuming plastic debris (Avery-Gomm et al. 2013).
Tufted Puffins ingest both consumer debris and industrial pellets (Blight and Burger 1997). While the
specific effects of plastics on puffins have not been studied, there is evidence that large plastic loads interfere
with feeding and digestion in other seabirds and can release toxins directly into the bloodstream (Derraik
2002, Teuton et al. 2009). A study comparing data from two time periods (1969-1978 and 1988-1990) in
Alaska found that the frequency of plastics in Tufted Puffin stomachs had increased from about 15% to 25%
and the number of particles per bird had doubled (Robards et al. 1995). The most recent data are from
coastal Oregon, Washington, and British Columbia, where 89% of the Tufted Puffin stomachs sampled
contained plastics (Blight and Burger 1997). All studies to date have been of adult birds – chicks would need
to acquire plastic from prey dropped by adults and this has not been assessed. Wehle (1982), however, found
significantly higher plastic loads in the stomachs of subadult puffins and suggested that subadults may be less
capable of differentiating between plastics and prey items.

Organochlorines, heavy metals, and other contaminants. During the 20th century, chlorinated

hydrocarbons, also called organochlorines, were associated with population declines of numerous marine
birds in the North Pacific (reviewed in Elliott and Noble 1993). Organochlorines include a wide range of
chemical compounds used in industry (e.g., polychlorinated biphenyls [PCBs], dioxins) and agriculture (e.g.,
DDT, other pesticides, herbicides). Many of these contaminants persist in the environment for very long
periods, entering the marine ecosystem through atmospheric transport, runoff, and point source pollution.
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 Organochlorines accumulate in fatty tissues and become amplified at higher trophic levels, where predators
amass and store the contaminants ingested and assimilated by their prey. Toxicity and effects vary with
species and exposure, but can include eggshell thinning, embryo mortality, skeletal deformities, wasting, and
impaired chick rearing and incubation behaviors (Fry 1995). Tufted Puffins feed at a relatively high trophic
level, particularly as nestlings and during the breeding season when their diet is mainly piscivorous. It follows
that they are vulnerable to the bioaccumulation of organochlorines, as well as heavy metals and other
compounds, and studies have confirmed generally higher levels of contaminants in puffins than in lower
trophic feeders living in the same system (Burger and Gochfeld 2009a, 2009b).
Organochlorines and heavy metals have been detected in several populations of Tufted Puffins, but there is
little information on whether contaminants have adversely impacted the species’ health, reproduction, or
behavior. Research in the Aleutian Islands has documented mercury, chromium, and other metals in Tufted
Puffin tissue samples, but generally at low levels (Ricca et al. 2008, Burger and Gochfeld 2009a, 2009b).
Though Honda et al. (1990) detected high levels of cadmium in puffins from the Aleutian Islands, it was
unclear whether the metal was natural or anthropogenic in origin, or whether it was harming the health of the
birds. Sublethal levels of PCBs and other organochlorines have also been found in puffins sampled in the
Aleutian Islands and at sea (Tanaka 1989, Ricca et al. 2008). While some of these contaminants were
attributed to point sources (e.g., a former military base), there was strong evidence that oceanic and
atmospheric patterns, combined with the birds’ wide-ranging pelagic habits, made puffins vulnerable to
distant sources, including pesticide applications (Ricca et al. 2008).
The only published information on contaminant levels in Tufted Puffins in Washington comes from a single
adult female found dead near Smith Island in August 2009 that had extremely high levels of organochlorines
and polybrominated dipenyl ethers (PBDE) (Good et al. 2014; S. Pearson, pers. comm.). Based on pollutant
concentrations in the prey of Rhinoceros Auklets in Washington (Good et al. 2014), it is likely that puffins
nesting in the eastern Strait of Juan de Fuca carry higher contaminant loads than those inhabiting the outer
coast. In Oregon, organochlorine levels were low in puffin eggs collected in 1979 and no discernible shell
thinning was observed (Henny et al. 1982).
In analyses of bone and muscle tissue from Tufted Puffins nesting on Amchitka, an Aleutian Island exposed
to underground nuclear testing from 1965-1971, no levels of radionuclides were found above the minimum
detectable activity (Burger and Gochfeld 2007, Burger et al. 2007). These tests took place more than 30 years
after the last underground detonation suggesting, in this instance at least, that puffin populations were little
affected by point-source or global fallout radionuclide pollution (Burger and Gochfeld 2007).

Introduced Species
Introduced species rank as one of the chief threats to birds that nest on islands (Gill 2007). They have been
implicated in scores of extinctions since 1500, not to mention an untold number of local extirpations and
population declines (Donald 2010). Island birds tend to be naive to non-native predators, leaving themselves,
their eggs, and their nestlings vulnerable to attack. Introduced animals can also compete for nesting space
and food resources, alter habitat structure, or bring with them exotic parasites and diseases, while introduced
plants can alter the composition and structure of nesting habitats.
Historically, Tufted Puffin populations have suffered from a range of introduced mammals, including Arctic
Foxes, Norway Rats, Arctic Ground Squirrels, and European Rabbits (Piatt and Kitaysky 2002). Several
islands in Alaska saw puffins decline or disappear completely following the introduction of foxes for fur
farming (Bailey 1976), only to return after the foxes were removed (Piatt and Kitaysky 2002). In California’s
Farallon Islands, rabbits competed directly with puffins for nesting sites and are thought to have contributed
to rapid puffin declines during the 20th century (Ainley and Lewis 1974). In the Queen Charlotte Islands,
Raccoons introduced for their trapping potential have spread through much of the archipelago, crossing
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 distances of up to 950 m of open water and putting an estimated 80% of the archipelago’s burrowing seabirds
at risk, potentially including Tufted Puffins (Hartman and Eastman 1999).
Though introduced species have never been thoroughly assessed at Tufted Puffin colonies in Washington, it’s
known that Native Americans raised dogs on Tatoosh Island (Paterek 1996), and that European Rabbits were
introduced to a number of islands, including Destruction, Smith, Colville, Flattop, Matia, and Skipjack (Couch
1929, Aubry and West 1984). The impacts of dogs and rabbits were never studied at these sites, but puffins
have disappeared from Colville, Flattop, Matia, and Skipjack islands. European Rabbits remain a particular
concern because they persist on Destruction Island, which otherwise offers excellent habitat and supported
hundreds of nesting pairs as recently as 1975. Rhinoceros Auklets on this island have declined by an
estimated 50% since the 1970s (measured by burrow occupancy), in spite of higher hatching and fledging
rates (Pearson et al. in review). Though direct evidence is lacking, Pearson et al. (in review) have seen rabbits
occupying Rhinoceros Auklet burrows there, thus rabbits may have contributed to the declining trend of this
species through direct competition for burrow sites, particularly in prime grassland areas. A study in Iceland’s
Westman Islands found that Atlantic Puffin burrow occupancy and density was markedly lower in areas
occupied by rabbits (64% of puffin burrows inactive) when compared to an unoccupied area (only 4% were
inactive) (Vigfusdottir 2007). The presence of rabbits and competition for burrow sites is also thought to
have played a role in the decline of Tufted Puffins at California’s Farallon Islands (Ainley and Lewis 1974).
The impact of rabbits on burrowing seabirds has also been documented on various islands, including Ile
Verte, Kerguelen Islands, South Indian Ocean, and Santa Clara Island, Juan Fernández Islands, Chile, where
European Rabbits were removed in 1992 and 2003, respectively. Following their eradication, burrow
occupancy by breeding pairs of Blue Petrels (Halobaena caerulea) increased 8-fold on Ile Verte (Brodier et al.
2011) and of Pink-footed Shearwaters (Puffinus creatopus) from 43% to 60% on Santa Clara Island (Hodum
2007).
Beyond the direct effect of competition, rabbits also contribute to indirect effects, including hyperpredation
and reduced habitat quality for nesting. Hyperpredation can arise when the introduction of a prey species
adapted to high predation pressure (like the European Rabbit) results in higher predator densities and
increased predation pressure on less resilient endemic prey species (Smith and Quin 1996, Courchamp et al.
2000). On Destruction Island, overgrazing by rabbits also appears to have changed the island’s grassy bluff
plant communities (where nearly all puffin burrows are located) from primarily perennial bunch grasses to
mainly annual grasses (S. Pearson, pers. comm.; S. Thomas, pers. comm.). This has resulted in significant soil
erosion and land slippage through both the close cropping of grasses by rabbits and because annual grasses
have smaller root masses to hold the soil, which in turn has caused the loss of burrowing habitat for puffins
and other breeding seabirds on the island (S. Pearson, pers. comm.). No non-native animals other than the
European Rabbit are currently known to occur at puffin breeding sites in Washington, but their potential
introduction remains a threat, particularly for sites subject to visitation by people.
Invasive non-native plants can negatively affect seabirds by reducing or degrading nesting and roosting
habitat, preventing access to nest burrows, and entrapping chicks. Introduced plant species are present at
many seabird nesting sites in Washington, particularly in the San Juan Islands and Strait of Juan de Fuca. At
Protection Island, for example, non-native grasses (e.g., Bromus tectorum) and shrubs (e.g., Cytisus scoparius) are
common on sandy bluff habitat used by nesting Tufted Puffins and Rhinoceros Auklets (USFWS 2010).
Control measures and restoration goals for up to 8 ha of bluff habitat were included in a recently adopted
management plan for the site (USFWS 2010).

Commercial and Tribal Fishing Bycatch
Fisheries bycatch can be a major cause of mortality in seabirds (Žydelis et al. 2013). Historically, Tufted
Puffins were a major part of the bycatch of large-scale drift net fisheries in the North Pacific, with tens of
thousands or even hundreds of thousands of birds killed annually (DeGange and Hay 1991, DeGange et al.
1993). Though banned by international treaty since 1991, some illegal drift netting continues in the North
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 Pacific (e.g., Rosen 2011), with undocumented levels of seabird bycatch. Coastal gillnet fisheries also impact
puffin populations. In Japan, Tufted Puffins made up nearly 20% of the avian bycatch in the salmon fishery
in the 1970s and 1980s, with yearly totals numbering from about 10,000 to more than 30,000 birds (DeGange
and Day 1991). More recently, observers at Kodiak Island, Alaska, noted an estimated 110 Tufted Puffins
killed during the 2002 salmon gillnetting season (Denlinger 2006).
In Washington, bycatch from fishing activities has been suggested as a factor in the decline of Tufted Puffins
(Smith et al. 1997, Wahl 2005). Current commercial fishing in Washington waters occupied by puffins
includes net fisheries for salmon, sardines, shrimp, herring (in Puget Sound only), and anchovy (southern
outer coast only), as well as hook-and-line and/or trawl fisheries for salmon, Pacific Halibut (Hippoglossus
stenolepis), Albacore Tuna (Thunnus alalunga), and various groundfish (TCW Economics 2008). On the outer
coast, net fisheries for salmon exist in Grays Harbor and Willapa Bay, which are less frequented by puffins
than areas farther north that are closer to nesting colonies. Commercial gillnet fisheries also occur in some
bordering areas of southern British Columbia where puffins from Washington may forage. Smaller tribal
fisheries exist in Washington for some of these same species, including salmon gillnetting at Mukkaw Bay and
in parts of the Strait of Juan de Fuca, trolling for salmon, longlining for halibut and Sablefish (Anoplopoma
fimbria), trawling for other groundfish, and purse seining for sardines. There is also the potential for smallscale ceremonial and subsistence harvest of forage fish by tribes.
No comprehensive Tufted Puffin bycatch data are available, but a recent report on West Coast ground
fisheries found no cases of Tufted Puffin mortality in the at-sea Pacific Hake (Merluccius productus) fishery, nor
in several other coastal groundfish fisheries between 2002 and 2009 (NFSC 2008, Jannot et al. 2011). A study
of seabird strandings associated with gillnetting activity in Washington’s inland marine waters also did not
document any Tufted Puffin mortalities between 1969 and 2007 (Hamel et al. 2009). Tufted Puffins were
scarce in the study area, however, and Hamel et al. (2009) noted that other alcids dominated the bycatch,
which added an estimated 0.2% and 2.9% to the annual mortality rates for Common Murres. Tufted Puffins
also were not detected among the seabird bycatch associated with commercial purse seine fisheries for salmon
in the San Juan Islands and southernmost Strait of Georgia from 2001-2012 (WDFW 2001-2012).
Some commercial fisheries in Washington operate at levels much reduced from their historical peaks in
foraging habitat for Tufted Puffins along the outer coast and in the Strait of Juan de Fuca and San Juan
Islands (e.g., McShane et al. 2004). Nonetheless, even minimal levels of fishing-related mortality pose a
potential threat to the diminished Tufted Puffin population in Washington. Similarly, while puffins have not
been documented in derelict fishing nets recovered in northern Puget Sound and the Strait of Juan de Fuca
(Good et al. 2009, 2010), recovery data are considered an underestimate and even low levels of mortality may
be harmful to puffin populations.

Human Disturbance
Historically, many nesting colonies of Tufted Puffins have suffered from disturbances and/or habitat loss
associated with human activities including hunting, egg collection, fur farming, lighthouses, development,
recreation, and habitation (Piatt and Kitaysky 2002). Studies have also demonstrated that the species is
sensitive to even the small-scale disturbances associated with research activities (Frazer 1975, Pierce and
Simons 1986, Hatch et al. 2000). Human disturbance (including researcher presence, recreational visitation of
nesting islands, and low flying aircraft; Frazer 1974) has been, for the most part, addressed at Tufted Puffin
breeding sites in Washington. All of these sites are currently under some kind of habitat protection. Puffin
colonies along the Washington coast now receive only occasional visitation from scientists and federal or
state land managers, and are not currently considered threatened by development. Former nesting sites in the
San Juan Islands are also protected (USFWS 2010), but may be subject to more visitation because of the
much greater human activity in the area and limited enforcement.

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 Bald Eagle Predation
Bald Eagle numbers have steadily increased in Washington since the early 1980s (Stinson et al. 2007),
resulting in more potential opportunities for interactions with nesting seabirds. A long-term study at Tatoosh
Island attributes the decline in Common Murre numbers to direct and indirect effects associated with
increased Bald Eagle predation (Parrish et al. 2001). Eagles attacked the murres directly, but also created
disturbances that allowed increased predation by Glaucous-winged Gulls (Larus glaucescens) and Northwestern
Crows (Corvus caurinus) (Parrish et al. 2001). These factors contributed to complete nesting failure for the
Common Murre colony in 2007 and 2008 (Schrimpf et al. 2012). A similar dynamic between surface nesting
seabird populations and White-tailed Eagles (Haliaeetus albicilla) has been reported from northern Europe
(Hipfner et al. 2012).
Bald Eagles regularly prey upon Tufted Puffins (Vermeer et al. 1976, Piatt and Kitaysky 2002) and puffins are
known to alter their flight behavior in the presence of eagles, reducing or synchronizing their colony arrivals
and departures to reduce the risk of attack (Addison et al. 2007). Whether or not the frequency of eaglepuffin interactions is changing or adversely affecting puffin populations in Washington remains unknown, but
puffins with full bill loads appear reluctant to enter burrows when eagles are present (S. Pearson, pers.
comm.). The density of eagles at puffin colonies can be high. For example, during visits to Smith Island (15
ha) in 2007 and 2008, P. Hodum et al. (unpubl. data) observed densities of 2.2 eagles/ha and 2.3 eagles/ha,
respectively. Parrish et al. (2001) pointed out that the secondary effects of predation can be subtle, but that
even small behavioral changes can have important implications for population trends.

Other Factors
H armful algal blooms. Historically known as “red tides,” harmful algal blooms result from rapid,
temporary increases in local populations of particular dinoflagellates, protists, or other phytoplankton.
Harmful algal blooms have been increasing globally in recent decades and while their underlying causes are
complex and poorly understood, some laboratory experiments predict increased occurrence with climate
change (Anderson 1997, Peperzak 2003, Glibert et al. 2005, Lewitus et al. 2012). Negative impacts on
seabirds include mortality and morbidity from the ingestion of algae-produced toxins (e.g., domoic acid), as
well as from the fouling of feathers with associated proteinaceous foam produced by the dinoflagellate
Akashiwo sanguinea (Jessup et al. 2009). Seabirds are also known to alter their movements and foraging
behavior during blooms (Shumway et al. 2003), a particular concern during sensitive breeding periods. Some
ornithologists suspect that harmful algal blooms are an important and underreported cause of seabird
mortality worldwide (Shumway et al. 2003).
Though the effects of harmful algae on Tufted Puffins have never been studied directly, large blooms in
recent years have led to mass strandings and mortality of seabirds in Monterey Bay, California (Jessup et al.
2009), and off the outer Washington and Oregon coasts (Du et al. 2011, Phillips et al. 2011). The
Washington bloom reached waters in and around major puffin breeding colonies and peaked in September at
the end of the breeding season. No puffin mortalities were recorded in the aftermath, but an estimated
10,000 Surf Scoters (Melanitta perspicillata), White-winged Scoters (M. fusca), and other seabirds died when their
feathers became fouled with proteinaceous foam (Welch 2009, Phillips et al. 2011). This was the first
recorded bloom of A. sanguinea in Washington, but in 1942 over 2,000 dead seabirds stranded on the coast
coincident with a bloom of Alexandrium catenella (McKernan and Scheffer 1942, Landsberg 2002). Other
blooms that have occurred in the region include Pseudo-nitzschia (which produces domoic acid) and multiple
species producing potent neurotoxins (saxitoxins), which are known to affect piscivorous birds by
contaminating their prey base (Landsberg 2002, Lewitus et al. 2012).
Based on the increased frequency and severity of harmful dinoflagellate blooms in recent years, Jessup et al.
(2009) hypothesized that the negative effects from these events could become more common in the future. If
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 this occurs and the blooms overlap with critical breeding and fledging periods, their effect on puffin
populations could be severe.

Pathogens and disease. Though the impacts of particular pathogens and diseases on Tufted Puffins remain
poorly understood, the potential effects of these are likely to increase when birds are under stress from other
environmental factors. The physical condition of adult male puffins, for example, is known to decline when
attempting to provision offspring during periods of prey shortage (Williams et al. 2007) and could make some
birds more susceptible to disease. Also, nesting in dense colonies increases the potential consequences of a
major disease outbreak (Gill 2007).
Egg collection. Historically, seabird breeding colonies in Washington experienced hunting and egg

collection by Native Americans, and later by Euro-American settlers, fishermen, and natural history
collectors. Tufted Puffins were included in these annual hunts, and Jones (1908a) recounted Quileute egg
collectors preceding his and Dawson’s 1907 visit to islands near La Push: “Fortunately for our purposes, the
seas prevented a landing at Carroll Islet, but the adjoining island of Wishalooth was as nearly cleaned of eggs
as it was possible for them to clean it.” Later confiscated by a game warden, the collectors’ haul included
over 250 eggs of Glaucous-winged Gulls, Common Murres, and Tufted Puffins. The impacts of such
harvests were never quantified, but establishment of the coastal refuges curtailed hunting and may have
contributed to a subsequent rise in Tufted Puffin numbers (Bent 1919). Currently, egg collection is
prohibited by state and federal law and although the patrolling of nesting sites is limited, this threat is
considered minimal.

Brown Pelican impacts. Since 2008, increasing numbers of Brown Pelicans (Pelecanus occidentalis) have

roosted on four islands (Willoughby Rock, Erin’s Bride, Grenville Arch, Grenville Pillar) currently or
formerly used by nesting Tufted Puffins in Washington, with hundreds of pelicans present on each of the
islands in 2014 (S. Pearson, pers. comm.). The birds have caused substantial vegetation change in their
roosting areas, which could potentially result in soil erosion and a loss of nesting habitat for puffins. This
impact should be monitored in the future to determine actual effects on puffins.

CONCLUSIONS AND RECOMMENDATION
Tufted Puffins are pelagic, pursuit diving seabirds that congregate annually on coastal islands and headlands
to breed and raise their chicks to fledging. In Washington, small nesting colonies in the San Juan archipelago
began declining as early as the 1950s, but the species was still considered a common bird along the outer
coast into the 1990s. However, Tufted Puffin sightings declined by 13.9% per year in pelagic surveys off
Westport, Washington, between 1983 and 2001 and 8.9% per year in offshore surveys along the outer
Washington coast from Cape Flattery to Point Grenville from 2001 to 2012. Colony visits from 2007 to 2010
noted widespread abandonment and an apparent order of magnitude population decline since the last
summary assessment from 1978-1984. Overall, 57% of the 44 historically documented puffin breeding sites
in Washington are no longer active. Maximum estimates once ranked nine colonies at 1,000 or more
individuals, with two reaching 10,000 or more birds, but none of the 19 remaining breeding sites now
contains more than a few hundred birds. In combination, of the 44 once-documented breeding sites, 39 have
either been abandoned or experienced an order of magnitude decline. Puffins in Washington’s inland marine
waters are at particular risk, with only two active breeding sites now remaining, both in the eastern Strait of
Juan de Fuca, that together contain a total population of a few dozen individuals.
Washington’s population trends fit a larger geographic pattern for Tufted Puffins, which appear to be
undergoing a range contraction. Though apparently stable in the northern portion of their range in Alaska
and Siberia, puffins have been declining throughout the southern extent of their range, including California,
Oregon, and Japan. Causes for the decline in Washington are not known, but may stem from a number of
current and historical threats, including reduced prey availability, changing ocean conditions, entrapment in
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 fishing nets, mortality from oil spills and chemical contaminants, human disturbance of breeding colonies,
impacts from introduced species, and increased Bald Eagle predation. Predicted shifts in ocean surface
temperature, ocean acidity, and weather patterns associated with climate change are expected to make
conditions challenging for piscivorous seabirds across the North Pacific, including Tufted Puffins. If the
current 8.9% annual rate of decline continues, the Washington population could become functionally
extirpated within about 40 years. Based on recent and continuing dramatic population declines and
widespread colony abandonment, and with conditions expected to worsen for this species in Washington and
throughout the North Pacific in the years ahead, it is recommended that the Tufted Puffin be listed as
endangered in Washington.

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45

Washington Department of Fish and Wildlife

 PERSONAL COMMUNICATIONS AND SOURCES OF UNPUBLISHED
DATA
Peter Arcese
University of British Columbia
Vancouver, British Columbia

Tony Scruton
The San Juan Preservation Trust
Waldron Island, Washington

Kurt Fresh
Northwest Fisheries Science Center
Seattle, Washington

Leslie Slater
Alaska Maritime National Wildlife Refuge
Homer, Alaska

Mark Hipfner
Pacific Wildlife Research Center
Environment Canada
Delta, British Columbia

Sue Thomas
Washington Maritime National Wildlife Refuge
Complex
Sequim, Washington

Peter Hodum
University of Puget Sound
Tacoma, Washington

Bill Tweit
Washington Department of Fish and Wildlife
Olympia, Washington

Scott Horton
Washington State Department of Natural Resources
Forks, Washington

Terence Wahl
Bellingham, Washington

Arthur Kettle
Alaska Maritime National Wildlife Refuge
Homer, Alaska
Deanna Lynch
U.S. Fish and Wildlife Service
Lacey, Washington

Lorna Wargo
Washington Department of Fish and Wildlife
Montesano, Washington
Ulrich Wilson
Washington Maritime National Wildlife Refuge
Complex (retired)
Sequim, Washington

Robert Paine
University of Washington (retired)
Seattle, Washington
Julia Parrish
University of Washington
Seattle, Washington
Scott Pearson
Washington Department of Fish and Wildlife
Olympia, Washington
Sara Perry
Seattle Aquarium
Seattle, Washington

January 2015

46

Washington Department of Fish and Wildlife

 Appendix A. Counts of Tufted Puffins at breeding sites and other locations in Washington,
1886-2013.
Colony Name

County

Year

Count

Observer

Reference

Bodelteh Island East

Clallam

1978

20

Pitman

Speich & Wahl 1989

Bodelteh Island East

Clallam

1982

6

Wilson

Speich & Wahl 1989

Bodelteh Island East
Bodelteh Island East
Bodelteh Island East

Bodelteh Island Middle

Bodelteh Island Middle
Bodelteh Island Middle
Bodelteh Island Middle
Bodelteh Island Middle
Bodelteh Island Middle
Bodelteh Island Middle
Bodelteh Island West
Bodelteh Island West
Bodelteh Island West
Bodelteh Island West
Bodelteh Island West

Bodelteh Island West
Bodelteh Island West
Bodelteh Island West

Bodelteh Islands (all)
Bodelteh Islands (all)
Bodelteh Islands (all)
Bodelteh Islands (all)

Bodelteh Islands (all)
Bodelteh Islands (all)
Bodelteh Islands (all)
Bodelteh Islands (all)
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cake Rock
Cakesosta

January 2015

Clallam
Clallam

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

1979
1979
2008
1978
1978
1979
1982
2008
2009
2010
1978
1978
1979
1979
1982
2008
2009
2010
1907
1959
1978
2007
2007
2008
2010

2010
1907
1959
1967

1968-1969
1978
1978
1978
1981
2007
2007
2008
2009

2010
2010
2010
1907

7

10
0

Breeding
Breeding
250
8

28
11
4

150-200

Breeding
130

Breeding
10
51
0

59

500-1,000
20

1,000-2,000
191
31

811,2
01

201

500
30

Breeding
Breeding
1,000
480+

500-1,000
70
17
0

982
60
74
50
47

500

47

Graybill
Pitman

Hodum et al.
Speich

Pitman
Pitman
Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Speich

Pitman
Pitman
Speich

Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Dawson

Kenyon & Scheffer
Pitman

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Dawson

Kenyon & Scheffer
Hancock
Cody

Speich
Speich

Pitman
Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Dawson

Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Dawson 1908b

Kenyon & Scheffer 1961
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Dawson 1908b

Kenyon & Scheffer 1961
Speich & Wahl 1989
Cody 1973

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Dawson 1908b

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Cakesosta

Clallam

1978

60-80

Speich

Speich & Wahl 1989

Cakesosta

Clallam

2008

822

Hodum et al.

Hodum et al., unpubl.

Cakesosta
Cakesosta
Cakesosta

Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island
Carroll Island

Carroll Island
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dhuoyautzachtahl
(Petrel Rock)
Dohodaaluh
Hand Rock

Hand Rock
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
January 2015

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

Clallam
Clallam
Clallam
Clallam
Clallam

1981
1982
2009
1907
1907
1907
1907

125+
6+
2

10,000

Breeding

Breeding3

1967

5,000
5,00010,000
100s

1978

1,000-2,000

1909
1968-1969
1978
1978
1978
1979
1979
1979
1981
1982
2008
2009
2010
2010
2010
1907
1919
1954
1979
1981
2007
2008
2009
2009
1907

1914-1915
1978
1907
1907
1907

Breeding
100s

4,000
8,000
2,000

Breeding
6,800
250

2,270
40

136
115
18

211
300

Specimen
Breeding
1,100
25
64
41
3
4

40

5,000
0

1,000

Breeding
Breeding

48

Wilson
Wilson

Hodum et al.
Dawson
Jones
Jones

Dawson
Dawson

Hancock
Cody

Speich

Pitman
Pitman
Speich

Pitman
Speich
Speich

Wilson
Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.

Hodum et al.
Dawson

Albrecht

Alcorn, Eddy
Speich

Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Dawson

Cantwell
Pitman

Dawson
Dawson
Jones

Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Dawson 1908a
Jones 1909

Speich & Wahl 1989
Dawson 1908b

Dawson and Bowles 1909
Speich & Wahl 1989
Cody 1973

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Dawson 1908b

Speich & Wahl 1989

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Dawson 1908b

Jewett et al. 1953

Speich & Wahl 1989
Dawson 1908b

Speich & Wahl 1989
Jones 1908a

Washington Department of Fish and Wildlife

 Colony Name
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
Jagged Island
(Wishalooth, Bald I.)
James Island
James Island
James Island
James Island

Reference

Clallam

1959

30

Kenyon & Scheffer

Kenyon & Scheffer 1961

Clallam

1978

10,000

Pitman

Speich & Wahl 1989

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

Clallam

Kochaauh

Clallam

Kochaauh

Clallam

Kochaauh

Clallam

Kochaauh

Clallam

Kochaauh

Clallam

Kochaauh

Clallam

Ozette Island
Ozette Island

Quillayute Needles Group4
Quillayute Needles Group4
Quillayute Needles Group4
Group4

Quillayute Needles Group4
Quillayute Needles Group4
Quillayute Needles Group4

January 2015

Observer

Clallam

Kochaauh

Seal Rock

Count

Clallam

Kochaauh

Quillayute Needles

Year

Clallam

James Island

Quillayute Needles

County

Group4

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

1967
1978
1979
1979
1979
1981
1982
2005
2005
2007
2008
2009
2010
2010
1975
2007
2007
2007
2007
1954
1967
1978
2007
2008
2009
2010
2010
1979
2007
1954
1959

1968-1969
2007
2007
2007
2010
2010
1976

Breeding
Breeding
1,000s

Breeding
12,000
950+

7,800
768
159
1

206
117
29
36

Breeding
13
30
12
0

Breeding
Breeding
200-400
30
24
1
0

33
0
0

Specimen
25

Breeding
39
63
0

35
5

Breeding

49

Hancock
Speich

Pitman
Speich
Speich

Wilson
Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Chappell

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Eddy

Hancock
Speich

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Speich

Hodum et al.
Anonymous

Kenyon & Scheffer
Cody

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Chappell

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989

Hodum et al., unpubl.
Speich & Wahl 1989

Kenyon & Scheffer 1961
Cody 1973

Hodum et al., unpubl.

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Seal Rock

Clallam

1978

25

Speich

Speich & Wahl 1989

Seal Rock

Clallam

1978

41

Wahl

Speich & Wahl 1989

Seal Rock
Seal Rock
Seal Rock
Seal Rock
Seal Rock

Silver Sides
Silver Sides
Silver Sides
Silver Sides
Table Rock
Table Rock

Tatoosh Island
Tatoosh Island
Tatoosh Island

Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island

Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island

Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island
Tatoosh Island

Unnamed Rock 1
Unnamed Rock 2
Unnamed Rock 2
White Rock
White Rock
White Rock
White Rock
White Rock
White Rock
White Rock
White Rock

Erin's Bride
Erin's Bride
January 2015

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam
Clallam

Grays Harbor
Grays Harbor

1978
1978
1978
1981
2008
1907
1978
1978
2008
1978
2009
1907
1914
1959
1968
1970
1971
1973
1973
1975
1978
1978
2005
2005
2005
2008
2008
2009
2009
2009
2010
2010
1978
1978
1982
1907
1959
1978

1979
2007
2007
2008
2009
1976
1978

36
24

Breeding
8
0

1,000

Breeding
200
812

Breeding
0

Breeding
Breeding
50

Breeding
200
20
60

50+

Breeding
100+
300
55

146
40
24
45
1
1
7
9

36
60

200
110

200-500
10

150

Breeding
3
0

65
1

Breeding
16+

50

Harrington-Tweit
Harrington-Tweit
Chappell
Wilson

Hodum et al.
Dawson
Pitman
Speich

Hodum et al.
Speich

Hodum et al.
Dawson

Cantwell

Kenyon & Scheffer
WA Dept. of Game
Paulson

Willapa NWR
Frazer

Leschner
Leschner
Boersma
Pitman

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Pitman
Pitman
Wilson

Dawson

Kenyon & Scheffer
Pitman
Speich

Hodum et al.
Hodum et al.
Hodum et al.

Hodum et al.
Hunn

Harrington-Tweit

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Dawson 1908b

Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Speich & Wahl 1989

Hodum et al., unpubl.
Dawson 1908b

Speich & Wahl 1989

Kenyon & Scheffer 1961
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Dawson 1908b

Kenyon & Scheffer 1961
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Erin's Bride

Grays Harbor

2007

32

Hodum et al.

Hodum et al., unpubl.

Grenville Arch

Grays Harbor

2007

0

Hodum et al.

Hodum et al., unpubl.

Erin's Bride

Grays Harbor

Grenville Arch

Grays Harbor

Grenville Arch

Grays Harbor

Grenville Arch

Grays Harbor

Grenville Pillar

Grays Harbor

Grenville Pillar
Grenville Pillar

Point Grenville Cliffs
Point Grenville Cliffs

Point Grenville Rocks5
Point Grenville Rocks5
Point Grenville Rocks5
Point Grenville

Rocks5

Point Grenville Rocks5
Point Grenville Rocks5
Point Grenville Rocks5
Point Grenville

Rocks5

Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Puffin Rock (Erin)
Split Rock
Split Rock

Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Willoughby Rock
Smith Island
Smith Island

January 2015

Grays Harbor

Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor

Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor

Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor
Grays Harbor

Grays Harbor
Grays Harbor
Island
Island

2008
1915
2009
2009
1980
2007
2009

pre-1921
1980
1973
1974
1974
1975
1976
1977
1979
1980
1907
1973
1976
1978
1978
1979
1980
1981
1982
2007
2010
2010
2010
2009
2009
1907
1977
1979
1981
2007
2007
2008
2009
2010
2010
2010
1914
1914

6

Breeding
0
0
3
0
0

Breeding
10-12
100
80

40-50
20
64

Breeding
Breeding
Breeding
2,000
98

Breeding
16

200
42
62
50
15
38
22
23
25
0
0

500

Breeding
28

120+
27
39
23
6
3
0
7

Breeding
500

51

Hodum et al.
Cantwell

Hodum et al.
Hodum et al.

Harrington-Tweit
Hodum et al.
Hodum et al.
Fletcher
Smith

Leschner

Hoge & Morris
Hoge & Hoge

Crowell & Nehls
Hoge & Hoge

Harrington-Tweit
Hoge & Hoge

Harrington-Tweit
Dawson
Frazer
Hunn

Harrington-Tweit
Speich
Speich

Harrington-Tweit
Wilson
Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Dawson

Harrington-Tweit
Speich

Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Cantwell
Cantwell

Hodum et al., unpubl.
Jewett et al. 1953

Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Crowell & Nehls 1975
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Dawson 1908b

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Dawson 1908b

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Smith Island

Island

1915

150

Cantwell

Speich & Wahl 1989

Smith Island

Island

1973

1

Manuwal

Speich & Wahl 1989

Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island
Smith Island

Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island
Alexander Island

Destruction Island
Destruction Island
Destruction Island

Destruction Island
Destruction Island
Destruction Island
Destruction Island
Destruction Island
Destruction Island
January 2015

Island
Island
Island
Island
Island
Island
Island
Island
Island
Island
Island
Island
Island
Island
Island
Island

Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson

Jefferson
Jefferson
Jefferson
Jefferson
Jefferson

Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson

1916
1917
1979
1982
1983
1983
1984
1985
1986
2007
2007
2007
2008
2008
2009
2010
1907
1907

1914-1915
1974
1974
1977
1978
1978
1978
1981
2005
2005
2005
2007
2007
2008
2009
2010
2010
2010

pre-1921
1963
1963
1963
1967

1968-1969
1973
1973
1973

150
150
44
8
1
6

10
3
6

21
11
15
25
18
5
4

5,000

Breeding
5,000

Breeding
300-400
80

600-1,000
400

4,000
45+
117
706
123
145
158
136
116
11
63
6

Present

Specimens
Specimen
Breeding
Breeding
Breeding
550

100s
400

52

Cantwell
Cantwell

Nisqually NWR

Nisqually NWR
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC

Hodum et al.
Hodum et al.

Hodum et al.
Hodum et al.
Hodum et al.
Thomas
Thomas

Dawson
Jones

Cantwell
Hoffman

Nysewander

Harrington-Tweit
Pitman
Speich
Speich

Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Lien

Hudson
LaFave
LaFave

Hancock
Cody

Leschner
Hoffman

Frazer, Reick

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.

S. Thomas, pers. comm.

S. Thomas, pers. comm.
Dawson 1908b
Jones 1908b

Speich & Wahl 1989

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Rathbun, unpubl.7

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Cody 1973

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Destruction Island

Jefferson

1975

650-700

Frazer

Speich & Wahl 1989

Destruction Island

Jefferson

1979

Breeding

Speich

Speich & Wahl 1989

Destruction Island
Destruction Island

Destruction Island
Destruction Island
Destruction Island
Destruction Island
Ghost Rock

Half Round Rock
Half Round Rock
Half Round Rock
Half Round Rock
Half Round Rock
Half Round Rock

Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island

Protection Island
Protection Island
Protection Island
Protection Island

Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island
Protection Island

Protection Island
Protection Island
Protection Island
Rounded Island
Rounded Island
January 2015

Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson

1976
1976
2007
2008
2010
2010
1981
1974
1978
1981
2007
2009
2009
1923
1955

1956-1959
1966
1967
1968
1971
1973

1975-1976
1976
1978
1979
1980
1980
1982
1984
1984
1986
1988
1989
1995
2007
2007
2007
2007
2008
2008
2011
2012
2013
1907
1978

Specimen
Specimen
922
52
30
19
25
1

20
2+
0
0
0

80-100
140

Breeding
Breeding
Breeding
Breeding
0

60-70
66
33

70-100
40
76
32
45
50
89
25
45
40
26
40
14
35
4

37
23
23
20
22

500

150

53

Anonymous
Welch

Hodum et al.
Hodum et al.
Hodum et al.

Hodum et al.
Wilson

Hoffman
Speich

Wilson

Hodum et al.
Hodum et al.
Hodum et al.
Pennington
Eddy

Richardson
Wahl
Wahl
Wahl
Wahl

Frazer

Wilson

Thompson
Pitman
Speich

Hirsch

Galusha et al.
Galusha et al.
Galusha et al.
WMNWRC
WMNWRC
WMNWRC
WMNWRC
Wilson

Hodum et al.
Hodum et al.

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
WMNWRC
WMNWRC
WMNWRC
Dawson
Speich

Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Richardson 1961

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Galusha et al. 1987

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Galusha et al. 1987
Galusha et al. 1987
Galusha et al. 1987

WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
Smith et al. 1997

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
Dawson 1908b

Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Rounded Island

Jefferson

1981

20

Wilson

Speich & Wahl 1989

Rounded Island

Jefferson

2008

26

Hodum et al.

Hodum et al., unpubl.

Rounded Island
Rounded Island
Rounded Island
Rounded Island
Rounded Island

Rounded Island
Rounded Island
Bare Island8
Bare Island8
Bare

Island8

Bare Island8
Bare Island8
Bare Island8
Bare

Island8

Bare Island8
Bare Island8
Bare Island8
Bare

Island8

Bare Island8
Bare Island8
Bare Island8
Bare

Island8

Bare Island8
Bare Island8
Bare Island8
Bare

Island8

Bare Island8
Bare Island8
Bare Island8
Bare

Island8

Bare Island8
Bird Rocks8
Bird Rocks8
Bird

Rocks8

Bird Rocks8
Bird Rocks8
Bird Rocks8
Castle

Island8

Castle Island8
Castle Island8
Colville

Island8

Colville Island8
Colville Island8
Colville Island8
Colville

Island8

January 2015

Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
Jefferson
San Juan
San Juan
San Juan

San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan

San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan

2007
2007
2009
2009
2010
2010
2010
1903
1905

1933-1935
1936
1937
1949

1956
1957
1959
1959
1962
1962
1963
1963
1967

1973-1975
1973
1974
1976
1978
2007
2007
2008
2008
1905
1937
1942
1984
2009
2010
1928
1961
2007
1957
1961
1962
1967
1968

115
65
2

45
50
14
12

Breeding3
40

Breeding
Breeding
100
16

Breeding
Present

Breeding
<40
20
14
2
1
5
4

18
3

Present
4
0
0
0
0
6
2

Breeding3
1
0
0
4
0
0

6+
2
6
0

Breeding

54

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Edson
Edson

Miller et al.
Miller

Jewett

Hudson

Richardson
Eddy

Bakus

Dickerman

Nisqually NWR
Eddy

Nisqually NWR

Hauser & Monson
Scruton

Manuwal
Manuwal
Eddy

Games

Speich & Wahl
Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
Edson

Jewett

Bushnell

WMNWRC
WMNWRC
WMNWRC
Rathbun
Eddy

Hodum et al.
Eddy
Eddy

Nisqually NWR
Wahl
Wahl

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Edson 1929b

Miller et al. 1935
Miller 1936

Speich & Wahl 1989
Speich & Wahl 1989

T. Scruton, pers. comm.
Speich & Wahl 1989
Speich & Wahl 1989
Dickerman 1960

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

T. Scruton, pers. comm.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.

Hodum et al., unpubl.
Hodum et al., unpubl.
Edson 1929a

Speich & Wahl 1989
Speich & Wahl 1989
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
Rathbun, unpubl.7

Speich & Wahl 1989

Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Island8

San Juan

1970

0

Wahl

Speich & Wahl 1989

Colville Island8

San Juan

1983

9

WMNWRC

WMNWRC, unpubl.6

Colville

Colville Island8
Colville Island8
Colville
Colville

Island8
Island8

Colville Island8
Colville Island8
Colville

Island8

Flattop

Island8

Flattop Island8
Flattop Island8
Flattop Island8
Flattop Island8
Flattop Island8
Goose Island
Goose Island
Hall Island8
Hall
Hall

Island8
Island8

Hall Island8

Matia Island8
Island8

Matia

Island8

Matia

Mummy Rocks
Mummy Rocks

North Pacific Rock
Puffin Island8
Puffin

Island8

Puffin Island8
Puffin Island8
Puffin Island8
Puffin
Puffin

Island8

Island8

Puffin Island8
Secar Rock
Secar Rock
Shark Reef

Skipjack Island8
Skipjack

Island8

Skipjack Island8
South Peapod8
Sucia

Islands8

The Sisters

The Sisters (Little Sister)
Turn Island
Turn Island

January 2015

San Juan
San Juan

San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
SanJuan

San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan
San Juan

1978
1979
1984
2007
2007
2008
2008
1904
1905
1905
1949
2008
2008
1937
2007
1978
1990
2009
2010
1984
2009
2010
1989
1990
1990
1938
1957
1957
1963
1963
1982
2008
2008
1982
1985
1978
1905

1933-1935
1936
1949
1886
1963
1976
1939
1979

20-30
5
5
0
0
0
0

Specimen
4-6
2

Specimen
0
0
0
0

10
1
0
0

10
0
0
1
2
1

Breeding3
4

8+
2
7
1
0
0
0
1
1

Present

Breeding
Breeding

Breeding

Breeding3
2
1

Specimen
0

55

Pitman
Wahl

WMNWRC

Hodum et al.
Hodum et al.
Hodum et al.
Hodum et al.
MacKay
Edson
Edson

Hudson

Hodum et al.
Hodum et al.
Jewett

Hodum et al.
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC
WMNWRC
Ray

Richardson
Eddy

Nisqually NWR

Hauser & Monson
WMNWRC

Hodum et al.
Hodum et al.
Wahl

WMNWRC
WMNWRC
Edson

Miller et al.
Miller

Hudson

Randolph, Johnson
Hauser & Monson
WMNWRC

Anonymous
Wahl

Speich & Wahl 1989
Speich & Wahl 1989
WMNWRC, unpubl.6

Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
Edson 1929b
Edson 1929b

Speich & Wahl 1989

Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989

Hodum et al., unpubl.
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6
WMNWRC, unpubl.6

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
WMNWRC, unpubl.6

Hodum et al., unpubl.
Hodum et al., unpubl.
Speich & Wahl 1989
WMNWRC, unpubl.6
WMNWRC, unpubl.6
Edson 1929b

Miller et al. 1935
Miller 1936

Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
WMNWRC, unpubl.6

Speich & Wahl 1989
Speich & Wahl 1989

Washington Department of Fish and Wildlife

 Colony Name

County

Year

Count

Observer

Reference

Unnamed (NWR Island 14)

San Juan

1980

6

WMNWRC

WMNWRC, unpubl.6

Waldron Island

San Juan

1975

0

Eddy

Speich & Wahl 1989

Waldron Island
Waldron Island
Whale Rocks
Viti Rocks
Viti Rocks
Viti Rocks

Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
Williamson Rocks
1

Williamson Rocks

San Juan
San Juan
San Juan
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit
Skagit

1895
1949
2007
1927
1949
1975
1905
1928
1928
1930
1930
1957

1957-1976
1963
1963
1967
1975
1977
1985
2008
2008

Specimen

Specimens
0

Specimen
4?

Edson

Hudson

Hodum et al.
Booth

Hudson

0, 4 old burrows
12

Breeding3
Several

Breeding3
Several
16

Breeding
8
1
5
1

6+
2
0
0

Eddy

Edson
Booth
Booth

Speich & Wahl 1989
Speich & Wahl 1989

Hodum et al., unpubl.
Speich & Wahl 1989
Speich & Wahl 1989
Speich & Wahl 1989
Edson 1929a

Speich & Wahl 1989
Speich & Wahl 1989

Booth
Booth

Thorsen & Booth
Thorsen

Thorsen & Galusha
Hauser & Monson
Nisqually NWR
Eddy

Thorsen

WMNWRC

Hodum et al.
Hodum et al.

Speich & Wahl 1989
Speich & Wahl 1989

Speich & Wahl 1989
Thorsen 1981

Thorsen & Galusha 1971
Speich & Wahl 1989

Speich & Wahl 1989
Speich & Wahl 1989

Thorsen 1980, 1981
WMNWRC, unpubl.6

Hodum et al., unpubl.
Hodum et al., unpubl.

No puffins were detected on Bodelteh East.
Colony size was estimated at ≥100 puffins (S. Pearson, pers. comm.).
3
Breeding was determined by the collection of one or more eggs, which are held as museum specimens (Speich and Wahl 1989).
4
The Quillayute Needles Group is comprised of various rocks and small islands, including Cakesota, Dhuoyautzachtahl, and Table
Rock (Speich and Wahl 1989), which are listed individually elsewhere in this appendix.
5
The Point Grenville Rocks are comprised of various rocks and small islands, including Erin’s Bride, Grenville Arch, Grenville Pillar,
and Puffin Island, as well as the Point Grenville Cliffs (Speich and Wahl 1989), which are listed individually elsewhere in this
appendix.
6
WMNWRC, Washington Marine National Wildlife Refuge Complex.
7
Unpublished field notes of S. F. Rathbun held at the Burke Museum, University of Washington, Seattle, Washington.
8
Classified here as a former breeding island. Speich and Wahl (1989) listed 11 San Juan Islands as current or former breeding
sites for Tufted Puffins, but we consider their supporting evidence as insufficient for three islands (The Sisters, Turn Island,
Waldron Island). We instead used the following criteria to establish an island as a breeding site in the San Juans: (1) records of
breeding, including collected egg specimens, are documented for the island in Speich and Wahl (1989), or (2) ≥3 puffins were
observed at least once at the island, and descriptions and photographs of the island (USFWS 2010) suggest that potentially
suitable habitat for breeding occurs there. Based on these criteria, we continue to classify 11 San Juan Islands as former breeding
sites. This includes two islands (Hall Island, Matia Island) where use was documented by the Washington Marine National Wildlife
Refuge Complex (unpubl. data) and one island (Castle Island) where breeding was documented by S. F. Rathbun (unpubl. notes);
use of these three islands was not reported in Speich and Wahl (1989).
2

January 2015

56

Washington Department of Fish and Wildlife

 Appendix B. Washington Administrative Codes 232-12-011, 232-12-014, and 232-12-297 and
Revised Codes of Washington 77.15.120 and 77.15.130.
WAC 232-12-011 Wildlife classified as protected shall not be hunted or fished.
Protected wildlife are designated into three subcategories: threatened, sensitive, and other.
(1) Threatened species are any wildlife species native to the state of Washington that are likely to become endangered within the
foreseeable future throughout a significant portion of their range within the state without cooperative management or removal of
threats. Protected wildlife designated as threatened include:
Common Name
Mazama pocket gopher
western gray squirrel
Steller (northern) sea lion
North American lynx
Aleutian Canada goose
bald eagle
ferruginous hawk
marbled murrelet
green sea turtle
loggerhead sea turtle
sage grouse
sharp-tailed grouse

Scientific Name
Thomomys mazama
Sciurus griseus
Eumetopias jubatus
Lynx canadensis
Branta canadensis leucopareia
Haliaeetus leucocephalus
Buteo regalis
Brachyramphus marmoratus
Chelonia mydas
Caretta caretta
Centrocercus urophasianus
Phasianus columbianus

(2) Sensitive species are any wildlife species native to the state of Washington that are vulnerable or declining and are likely to
become endangered or threatened in a significant portion of their range within the state without cooperative management or
removal of threats. Protected wildlife designated as sensitive include:
Common Name
gray whale
common Loon
peregrine falcon
Larch Mountain salamander
pygmy whitefish
margined sculpin
Olympic mudminnow

Scientific Name
Eschrichtius gibbosus
Gavia immer
Falco peregrinus
Plethodon larselli
Prosopium coulteri
Cottus marginatus
Novumbra hubbsi

(3) Other protected wildlife include:
Common Name
cony or pika
least chipmunk
yellow-pine chipmunk
Townsend's chipmunk
red-tailed chipmunk
hoary marmot
Olympic marmot
Cascade golden-mantled ground squirrel
golden-mantled ground squirrel
Washington ground squirrel
red squirrel
Douglas squirrel
northern flying squirrel
Wolverine
painted turtle
California mountain kingsnake

January 2015

Scientific Name
Ochotona princeps
Tamius minimus
Tamius amoenus
Tamius townsendii
Tamius ruficaudus
Marmota caligata
Marmota olympus
Spermophilus saturatus
Spermophilus lateralis
Spermophilus washingtoni
Tamiasciurus hudsonicus
Tamiasciurus douglasii
Glaucomys sabrinus
Gulo gulo
Chrysemys picta
Lampropeltis zonata

57

Washington Department of Fish and Wildlife

 All birds not classified as game birds, predatory birds or endangered species, or designated as threatened species or sensitive
species; all bats, except when found in or immediately adjacent to a dwelling or other occupied building; mammals of the order
Cetacea, including whales, porpoises, and mammals of the order Pinnipedia not otherwise classified as endangered species, or
designated as threatened species or sensitive species. This section shall not apply to hair seals and sea lions which are threatening
to damage or are damaging commercial fishing gear being utilized in a lawful manner or when said mammals are damaging or
threatening to damage commercial fish being lawfully taken with commercial gear.
[Statutory Authority: RCW 77.12.047, 77.12.655, 77.12.020. 06-04-066 (Order 06-09), § 232-12-011, filed 1/30/06, effective 3/2/06; 04-11-036
(Order 04-98), § 232-12-014, filed 5/12/04, effective 6/12/04. Statutory Authority: RCW 77.12.047, 77.12.655, 77.12.020. 02-11-069 (Order 0298), § 232-12-011, filed 5/10/02, effective 6/10/02. Statutory Authority: RCW 77.12.047. 02-08-048 (Order 02-53), § 232-12-011, filed 3/29/02,
effective 5/1/02; 00-17-106 (Order 00-149), § 232-12-011, filed 8/16/00, effective 9/16/00. Statutory Authority: RCW 77.12.040, 77.12.010,
77.12.020, 77.12.770. 00-10-001 (Order 00-47), § 232-12-011, filed 4/19/00, effective 5/20/00. Statutory Authority: RCW 77.12.040, 77.12.010,
77.12.020, 77.12.770, 77.12.780. 00-04-017 (Order 00-05), § 232-12-011, filed 1/24/00, effective 2/24/00. Statutory Authority: RCW 77.12.020.
98-23-013 (Order 98-232), § 232-12-011, filed 11/6/98, effective 12/7/98. Statutory Authority: RCW 77.12.040. 98-10-021 (Order 98-71), § 23212-011, filed 4/22/98, effective 5/23/98. Statutory Authority: RCW 77.12.040 and 75.08.080. 98-06-031, § 232-12-011, filed 2/26/98, effective
5/1/98. Statutory Authority: RCW 77.12.020. 97-18-019 (Order 97-167), § 232-12-011, filed 8/25/97, effective 9/25/97. Statutory Authority:
RCW 77.12.040, 77.12.020, 77.12.030 and 77.32.220. 97-12-048, § 232-12-011, filed 6/2/97, effective 7/3/97. Statutory Authority: RCW
77.12.020. 93-21-027 (Order 615), § 232-12-011, filed 10/14/93, effective 11/14/93; 90-11-065 (Order 441), § 232-12-011, filed 5/15/90,
effective 6/15/90. Statutory Authority: RCW 77.12.040. 89-11-061 (Order 392), § 232-12-011, filed 5/18/89; 82-19-026 (Order 192), § 232-12011, filed 9/9/82; 81-22-002 (Order 174), § 232-12-011, filed 10/22/81; 81-12-029 (Order 165), § 232-12-011, filed 6/1/81.]

WAC 232-12-014 Wildlife classified as endangered species. Endangered species include:
Common Name
pygmy rabbit
Fisher
gray wolf
grizzly bear
sea otter
sei whale
fin whale
blue whale
humpback whale
black right whale
sperm whale
killer whale
Columbian white-tailed deer
woodland caribou
American white pelican
brown pelican
sandhill crane
snowy plover
upland sandpiper
spotted owl
Streaked horned lark
western pond turtle
leatherback sea turtle
mardon skipper
Oregon silverspot butterfly
Taylor’s checkerspot
Oregon spotted frog
northern leopard frog

Scientific Name
Brachylagus idahoensis
Martes pennanti
Canis lupus
Ursus arctos
Enhydra lutris
Balaenoptera borealis
Balaenoptera physalus
Balaenoptera musculus
Megaptera novaeangliae
Balaena glacialis
Physeter macrocephalus
Orcinus orca
Odocoileus virginianus leucurus
Rangifer tarandus caribou
Pelecanus erythrorhynchos
Pelecanus occidentalis
Grus canadensis
Charadrius alexandrinus
Bartramia longicauda
Strix occidentalis
Eremophila alpestris strigata
Clemmys marmorata
Dermochelys coriacea
Polites mardon
Speyeria zerene hippolyta
Euphydryas editha taylori
Rana pretiosa
Rana pipiens

[Statutory Authority: RCW 77.12.047, 77.12.655, 77.12.020. 06-04-066 (Order 06-09), § 232-12-014, filed 1/30/06, effective 3/2/06. Statutory
Authority: RCW 77.12.047, 77.12.655, 77.12.020. 02-11-069 (Order 02-98), § 232-12-014, filed 5/10/02, effective 6/10/02. Statutory Authority:
RCW 77.12.040, 77.12.010, 77.12.020, 77.12.770, 77.12.780. 00-04-017 (Order 00-05), § 232-12-014, filed 1/24/00, effective 2/24/00. Statutory
Authority: RCW 77.12.020. 98-23-013 (Order 98-232), § 232-12-014, filed 11/6/98, effective 12/7/98; 97-18-019 (Order 97-167), § 232-12-014,
filed 8/25/97, effective 9/25/97; 93-21-026 (Order 616), § 232-12-014, filed 10/14/93, effective 11/14/93. Statutory Authority: RCW
77.12.020(6). 88-05-032 (Order 305), § 232-12-014, filed 2/12/88. Statutory Authority: RCW 77.12.040. 82-19-026 (Order 192), § 232-12-014,
filed 9/9/82; 81-22-002 (Order 174), § 232-12-014, filed 10/22/81; 81-12-029 (Order 165), § 232-12-014, filed 6/1/81.

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Washington Department of Fish and Wildlife

 WAC 232-12-297 Endangered, threatened, and
sensitive wildlife species classification.

2.9 "Significant portion of its range" means that portion of
a species' range likely to be essential to the long term
survival of the population in Washington.

PURPOSE

LISTING CRITERIA

1.1 The purpose of this rule is to identify and classify native
wildlife species that have need of protection and/or
management to ensure their survival as free-ranging
populations in Washington and to define the process by
which listing, management, recovery, and delisting of a
species can be achieved. These rules are established to
ensure that consistent procedures and criteria are followed
when classifying wildlife as endangered, or the protected
wildlife subcategories threatened or sensitive.
DEFINITIONS
For purposes of this rule, the following definitions apply:
2.1 "Classify" and all derivatives means to list or delist wildlife
species to or from endangered, or to or from the protected
wildlife subcategories threatened or sensitive.
2.2 "List" and all derivatives means to change the classification
status of a wildlife species to endangered, threatened, or
sensitive.
2.3 "Delist" and its derivatives means to change the
classification of endangered, threatened, or sensitive
species to a classification other than endangered,
threatened, or sensitive.

3.2 If a species is listed as endangered or threatened under
the federal Endangered Species Act, the agency will
recommend to the commission that it be listed as
endangered or threatened as specified in section 9.1.
If listed, the agency will proceed with development of
a recovery plan pursuant to section 11.1.
3.3 Species may be listed as endangered, threatened, or
sensitive only when populations are in danger of
failing, declining, or are vulnerable, due to factors
including but not restricted to limited numbers,
disease, predation, exploitation, or habitat loss or
change, pursuant to section 7.1.
3.4 Where a species of the class Insecta, based on
substantial evidence, is determined to present an
unreasonable risk to public health, the commission
may make the determination that the species need not
be listed as endangered, threatened, or sensitive.
DELISTING CRITERIA

2.4 "Endangered" means any wildlife species native to the state
of Washington that is seriously threatened with extinction
throughout all or a significant portion of its range within
the state.
2.5 "Threatened" means any wildlife species native to the state
of Washington that is likely to become an endangered
species within the forseeable future throughout a significant
portion of its range within the state without cooperative
management or removal of threats.
2.6 "Sensitive" means any wildlife species native to the state of
Washington that is vulnerable or declining and is likely to
become endangered or threatened in a significant portion of
its range within the state without cooperative management
or removal of threats.
2.7 "Species" means any group of animals classified as a
species or subspecies as commonly accepted by the
scientific community.
2.8 "Native" means any wildlife species naturally occurring in
Washington for purposes of breeding, resting, or foraging,
excluding introduced species not found historically in this
state.
January 2015

3.1 The commission shall list a wildlife species as
endangered, threatened, or sensitive solely on the basis
of the biological status of the species being
considered, based on the preponderance of scientific
data available, except as noted in section 3.4.

59

4.1 The commission shall delist a wildlife species from
endangered, threatened, or sensitive solely on the basis
of the biological status of the species being
considered, based on the preponderance of scientific
data available.
4.2 A species may be delisted from endangered,
threatened, or sensitive only when populations are no
longer in danger of failing, declining, are no longer
vulnerable, pursuant to section 3.3, or meet recovery
plan goals, and when it no longer meets the definitions
in sections 2.4, 2.5, or 2.6.
INITIATION OF LISTING PROCESS
5.1 Any one of the following events may initiate the
listing process.
5.1.1 The agency determines that a species
population may be in danger of failing,
declining, or vulnerable, pursuant to section
3.3.
5.1.2 A petition is received at the agency from an

Washington Department of Fish and Wildlife

 interested person. The petition should be addressed
to the director. It should set forth specific evidence
and scientific data which shows that the species
may be failing, declining, or vulnerable, pursuant to
section 3.3. Within 60 days, the agency shall either
deny the petition, stating the reasons, or initiate the
classification process.
5.1.3 An emergency, as defined by the Administrative
Procedure Act, chapter 34.05 RCW. The listing of
any species previously classified under emergency
rule shall be governed by the provisions of this
section.

the species' status in Washington and address factors
affecting its status, including those given under section
3.3. The status report shall be reviewed by the public
and scientific community. The status report will
include, but not be limited to an analysis of:
7.1.1 Historic, current, and future species population
trends.
7.1.2 Natural history, including ecological
relationships (e.g., food habits, home range,
habitat selection patterns).
7.1.3 Historic and current habitat trends.

5.1.4 The commission requests the agency review a
species of concern.
5.2 Upon initiation of the listing process the agency shall
publish a public notice in the Washington Register, and
notify those parties who have expressed their interest to the
department, announcing the initiation of the classification
process and calling for scientific information relevant to the
species status report under consideration pursuant to
section 7.1.
INITIATION OF DELISTING PROCESS
6.1 Any one of the following events may initiate the delisting
process:
6.1.1 The agency determines that a species population
may no longer be in danger of failing, declining, or
vulnerable, pursuant to section 3.3.
6.1.2 The agency receives a petition from an interested
person. The petition should be addressed to the
director. It should set forth specific evidence and
scientific data which shows that the species may no
longer be failing, declining, or vulnerable, pursuant
to section 3.3. Within 60 days, the agency shall
either deny the petition, stating the reasons, or
initiate the delisting process.

7.1.5 Historic and current species management
activities.
7.2 Except in an emergency under 5.1.3 above, the agency
shall prepare recommendations for species
classification, based upon scientific data contained in
the status report. Documents shall be prepared to
determine the environmental consequences of
adopting the recommendations pursuant to
requirements of the State Environmental Policy Act
(SEPA).
7.3 For the purpose of delisting, the status report will
include a review of recovery plan goals.
PUBLIC REVIEW
8.1 Except in an emergency under 5.1.3 above, prior to
making a recommendation to the commission, the
agency shall provide an opportunity for interested
parties to submit new scientific data relevant to the
status report, classification recommendation, and any
SEPA findings.
8.1.1 The agency shall allow at least 90 days for
public comment.

6.1.3 The commission requests the agency review a
species of concern.
6.2 Upon initiation of the delisting process the agency shall
publish a public notice in the Washington Register, and
notify those parties who have expressed their interest to the
department, announcing the initiation of the delisting
process and calling for scientific information relevant to the
species status report under consideration pursuant to
section 7.1.
SPECIES STATUS REVIEW AND AGENCY
RECOMMENDATIONS
7.1 Except in an emergency under 5.1.3 above, prior to making
a classification recommendation to the commission, the
agency shall prepare a preliminary species status report.
The report will include a review of information relevant to
January 2015

7.1.4 Population demographics (e.g., survival and
mortality rates, reproductive success) and their
relationship to long term sustainability.

60

FINAL RECOMMENDATIONS AND COMMISSION
ACTION
9.1 After the close of the public comment period, the
agency shall complete a final status report and
classification recommendation. SEPA documents will
be prepared, as necessary, for the final agency
recommendation for classification. The classification
recommendation will be presented to the commission
for action. The final species status report, agency
classification recommendation, and SEPA documents
will be made available to the public at least 30 days
prior to the commission meeting.
9.2 Notice of the proposed commission action will be
Washington Department of Fish and Wildlife

 published at least 30 days prior to the commission meeting.
PERIODIC SPECIES STATUS REVIEW
10.1 The agency shall conduct a review of each endangered,
threatened, or sensitive wildlife species at least every five
years after the date of its listing. This review shall include
an update of the species status report to determine whether
the status of the species warrants its current listing status or
deserves reclassification.
10.1.1 The agency shall notify any parties who have
expressed their interest to the department of the
periodic status review. This notice shall occur at
least one year prior to end of the five year period
required by section 10.1.

needed from and impacts to the department,
other agencies (including federal, state, and
local), tribes, landowners, and other interest
groups. The plan shall consider various
approaches to meeting recovery objectives
including, but not limited to regulation,
mitigation, acquisition, incentive, and
compensation mechanisms.
11.1.4 Public education needs.
11.1.5 A species monitoring plan, which requires
periodic review to allow the incorporation of
new information into the status report.

10.2 The status of all delisted species shall be reviewed at least
once, five years following the date of delisting.
10.3 The department shall evaluate the necessity of changing the
classification of the species being reviewed. The agency
shall report its findings to the commission at a commission
meeting. The agency shall notify the public of its findings
at least 30 days prior to presenting the findings to the
commission.
10.3.1 If the agency determines that new information
suggests that classification of a species should be
changed from its present state, the agency shall
initiate classification procedures provided for in
these rules starting with section 5.1.

11.2 Preparation of recovery and management plans will be
initiated by the agency within one year after the date
of listing.
11.2.1 Recovery and management plans for species
listed prior to 1990 or during the five years
following the adoption of these rules shall be
completed within five years after the date of
listing or adoption of these rules, whichever
comes later. Development of recovery plans
for endangered species will receive higher
priority than threatened or sensitive species.
11.2.2 Recovery and management plans for species
listed after five years following the adoption of
these rules shall be completed within three
years after the date of listing.

10.3.2 If the agency determines that conditions have not
changed significantly and that the classification of
the species should remain unchanged, the agency
shall recommend to the commission that the species
being reviewed shall retain its present classification
status.
10.4 Nothing in these rules shall be construed to automatically
delist a species without formal commission action.

RECOVERY AND MANAGEMENT OF LISTED SPECIES
11.1 The agency shall write a recovery plan for species listed as
endangered or threatened. The agency will write a
management plan for species listed as sensitive. Recovery
and management plans shall address the listing criteria
described in sections 3.1 and 3.3, and shall include, but are
not limited to:

11.2.3 The agency will publish a notice in the
Washington Register and notify any parties
who have expressed interest to the department
interested parties of the initiation of recovery
plan development.
11.2.4 If the deadlines defined in sections 11.2.1 and
11.2.2 are not met the department shall notify
the public and report the reasons for missing
the deadline and the strategy for completing
the plan at a commission meeting. The intent
of this section is to recognize current
department personnel resources are limiting
and that development of recovery plans for
some of the species may require significant
involvement by interests outside of the
department, and therefore take longer to
complete.
11.3 The agency shall provide an opportunity for interested
public to comment on the recovery plan and any
SEPA documents.

11.1.1 Target population objectives.
11.1.2 Criteria for reclassification.

CLASSIFICATION PROCEDURES REVIEW
11.1.3 An implementation plan for reaching population
objectives which will promote cooperative
management and be sensitive to landowner needs
and property rights. The plan will specify resources
January 2015

61

12.1 The agency and an ad hoc public group with members
representing a broad spectrum of interests, shall meet
as needed to accomplish the following:
Washington Department of Fish and Wildlife

 12.1.1 Monitor the progress of the development of
recovery and management plans and status reviews,
highlight problems, and make recommendations to
the department and other interested parties to
improve the effectiveness of these processes.
12.1.2 Review these classification procedures six years
after the adoption of these rules and report its
findings to the commission.
AUTHORITY

endangered under RCW 77.12.020. Species classified
as endangered are listed under WAC 232-12-014, as
amended.
13.2 Threatened and sensitive species shall be classified as
subcategories of protected wildlife. The commission
has the authority to classify wildlife as protected under
RCW 77.12.020. Species classified as protected are
listed under WAC 232-12-011, as amended.
[Statutory Authority: RCW 77.12.020. 90-11-066
(Order 442), § 232-12-297, filed 5/15/90, effective
6/15/90.]

13.1 The commission has the authority to classify wildlife as

RCW 77.15.120 Endangered fish or wildlife – Unlawful taking – Penalty.
(1) A person is guilty of unlawful taking of endangered fish or wildlife in the second degree if the person hunts, fishes,
possesses, maliciously harasses or kills fish or wildlife, or maliciously destroys the nests or eggs of fish or wildlife and the
fish or wildlife is designated by the commission as endangered, and the taking has not been authorized by rule of the
commission.
(2) A person is guilty of unlawful taking of endangered fish or wildlife in the first degree if the person has been:
(a) Convicted under subsection (1) of this section or convicted of any crime under this title involving the killing,
possessing, harassing, or harming of endangered fish or wildlife; and
(b) Within five years of the date of the prior conviction the person commits the act described by subsection (1) of this
section.
(3)(a) Unlawful taking of endangered fish or wildlife in the second degree is a gross misdemeanor.
(b) Unlawful taking of endangered fish or wildlife in the first degree is a class C felony. The department shall revoke
any licenses or tags used in connection with the crime and order the person's privileges to hunt, fish, trap, or obtain licenses
under this title to be suspended for two years.
[2000 c 107 § 236; 1998 c 190 § 13.]

RCW 77.15.130 Protected fish or wildlife — Unlawful taking — Penalty.
(1) A person is guilty of unlawful taking of protected fish or wildlife if:
(a) The person hunts, fishes, possesses, or maliciously kills protected fish or wildlife, or the person possesses or
maliciously destroys the eggs or nests of protected fish or wildlife, and the taking has not been authorized by rule of the
commission; or
(b) The person violates any rule of the commission regarding the taking, harming, harassment, possession, or transport
of protected fish or wildlife.
(2) Unlawful taking of protected fish or wildlife is a misdemeanor.
[1998 c 190 § 14.]

January 2015

62

Washington Department of Fish and Wildlife

 Appendix C. WDFW responses to public comments received during both the 90-day public
review period for the draft Washington State Status Report for the Tufted Puffin conducted from
September 12 to December 11, 2014, and the 14-day public review period for the final report
conducted under the State Environmental Policy Act (SEPA) from January 9-23, 2015. The
comments presented here are summaries of the remarks provided by one or more people.
Report Section

Comment and Response

General comments

1. I support the listing of Tufted Puffins as state endangered in Washington for the reasons
provided in the status report.
WDFW is recommending that Tufted Puffins be added to the state endangered species list
because of substantial scientific evidence indicating that the species has experienced a
major decline in abundance in Washington since about the 1970s. Tufted Puffin sightings
declined on average by 13.9% per year in pelagic surveys off Westport, Washington,
between 1983 and 2001, and 8.9% per year in offshore surveys along the outer Washington
coast from Cape Flattery to Point Grenville from 2001 to 2012. Colony visits from 2007 to
2010 noted widespread abandonment and an apparent order of magnitude population
decline since the last summary assessment from 1978-1984. Overall, 57% of the 44
historically documented puffin breeding sites in Washington are no longer active.
Maximum estimates once ranked nine colonies at 1,000 or more individuals, with two
reaching 10,000 or more birds, but none of the 19 remaining breeding sites now contains
more than a few hundred birds. In combination, of the 44 once-documented breeding sites,
39 have either been abandoned or experienced an order of magnitude decline. Puffins in
Washington’s inner marine waters are at particular risk, with only two active breeding
sites now remaining, both in the eastern Strait of Juan de Fuca, that together contain a
population of a few dozen individuals. None of the 11 historically known breeding colonies
in the San Juan Islands remain active.
2. I oppose the listing of Tufted Puffins as state endangered in Washington.
Comment noted. By law (WAC 232-12-297), species listings and delistings by the state
must be based solely on the biological status of the species and its continued existence in
the state.
3. There are concreate actions that can be taken in Washington to recover Tufted Puffins,
in particular doing more to restore prey fish. A plan should be formulated to stop the
decline of this species.
If the Washington Fish and Wildlife Commission classifies Tufted Puffins as a state
endangered species, WDFW will prepare a recovery plan for the species as required in
WAC 232-12-297, Section 11.1, which will describe different actions that can be taken to
recover puffins in Washington. These activities may include, for example, expanded
research on the species to determine the cause(s) of its decline; expanded monitoring of
prey fish trends; eradication of European rabbits from Destruction Island, where puffins
nest; management of invasive plants on nesting islands; and continued planning to avoid
oil spills in Washington’s marine waters.
4. The Washington State Department of Natural Resources hopes to collaborate with
WDFW in developing management actions for puffins on its aquatic reserves and other
aquatic lands in the state.

January 2015

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Washington Department of Fish and Wildlife

 Report Section

Comment and Response
WDFW appreciates this offer for collaboration and looks forward to working with the
Washington State Department of Natural Resources on puffin recovery planning and
conservation implementation in the future.
5. The puffin report’s conclusions and recommendation are completely unwarranted and
unsupported by the data available and presented. The report fails to adequately make a
case for listing Tufted Puffins as endangered in Washington. I want to see good,
documentable, applicable, verifiable, and real science that supports legitimate, not predetermined and ideological conclusions.
WDFW disagrees with this comment and believes the Washington State Status Report for
the Tufted Puffin was prepared using the best available scientific information on the
species. Draft versions of the report were reviewed by scientists and knowledgeable staff
within WDFW and numerous scientists and species experts outside of the agency. The final
status report reflects their suggested edits. None of these reviewers questioned the report’s
conclusions and recommendations, or its use of the science.
6. I used to see lots of sea ducks from Whidbey Island to Port Angeles in the 1970s and
1980s, and enjoyed hunting them. From what I read in your report, there is not enough
of a population to support harvesting puffins, therefore I support protecting them.
WDFW has never had a hunting season on Tufted Puffins. The status report does not
mention the hunting of puffins except during historical times, when tribal harvest of eggs
and birds was conducted.

Natural History

7. It might be useful to also cite the publication by Tyler et al. (1993), which states that
Tufted Puffins forage in fairly low densities.
This information was added to the status report (page 6).
8. Data confirming monogamy and long-term pair-bonding in this species are extremely
limited. We suggest using language indicating high mate fidelity, as opposed to
monogamy.
This correction was made in the status report (page 7).
9. The report notes that females lay only one egg, with a possible second egg being laid if
the first is lost. This naturally low reproductive rate should be discussed as a biological
factor causing Tufted Puffins to be particularly vulnerable to nest failure when
disturbances are present.
This information was not added to the status report because the authors were unable to
confirm in the scientific literature whether single-egg clutches do indeed make Tufted
Puffins more vulnerable to nest failure from disturbance.
10. We suggest including a reference by Peterson (1982) documenting Red Fox predation
on adults in Alaska.
This citation was not added to the status report because predation by Red Foxes was
already noted in the report.

January 2015

64

Washington Department of Fish and Wildlife

 Report Section

Comment and Response
11. We appreciate the inclusion of useful comparative data from related species on the
potential impact of gull kleptoparasitism. We have received anecdotal information that
these species also prey upon young Tufted Puffins. We suggest this as an area requiring
further research (to be added to the Research Needs section), particularly in light of
increasing abundance of gulls.
A statement about needing more information on gull kleptoparasitism was added to the
Research Needs section (page 20).

Population Status and
Trends

12. Table 2 in the status report shows that making comparisons between data sets is
problematic. For example, comparing Protection Island with Westport is difficult
because the former includes 10 years prior to the latter (with 1.5 times as many years
measured). We recommend finding a way to standardize years used for comparison
across sites. It would be helpful to see the data provided in Table 2 (separated out by
sites) plotted on one graph with time on the X-axis and percent population change on
the Y-axis.
This change was not made. Much of the information appearing in the table for locations
outside of Washington was taken from Piatt and Kitaysky (2002) and WDFW does not have
access to the data. Population trends for the four data sets in Washington are depicted in
greater detail in Figure 2 of the status report.

Conservation Status

13. The report mistakenly states that the U.S. Fish and Wildlife Service will decide whether
or not to federally list the Tufted Puffin in Washington, Oregon, and California in 2016
or 2017. In fact, this is when their listing evaluation will be initiated. A final listing
decision will likely not be made until 2018-2020.
This correction was made in the status report (pages v and 18).

Research, Monitoring, 14. The Restoration Potential section should also include information on habitat restoration
activities that could be done at current and former puffin breeding islands in
and Restoration
Washington.
This information was added to the status report (page 20).
Factors Affecting
Continuing Existence

15. The section on Adequacy of Existing Regulatory Mechanisms should clarify what
protections will be afforded to puffins through state listing as endangered.
This information was added to the status report (page 21).
16. I believe that the lack of sufficient small forage fish is very likely causing the decrease
in puffins and other marine waterbirds in Washington. WDFW and the Washington
Fish and Wildlife Commission have not done enough to reduce the harvest of forage
fish in the state.
The causes of the Tufted Puffin decline in Washington remain undetermined because of a
lack of scientific information. The status report includes a section on reduced prey
availability that discusses changes in forage fish populations in the state and how this
might affect Tufted Puffins. Harvest of several forage fish species in Washington have been
reduced in recent years in response to concerns about declining abundance of those
species. In addition, pelagic fisheries for sardines and anchovies are managed by the
Pacific Fishery Management Council (of which WDFW is a member) with a priority for
maintaining their ecosystem functions, primarily as a food source for seabirds, marine
mammals, salmon, and other predators.

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Washington Department of Fish and Wildlife

 Report Section

Comment and Response
17. In the Reduced Prey Availability section, we suggest citing a recent scientific paper by
Santora et al. (2014) that describes the relationships among prey species and seabirds in
California.
This reference was added to the status report (pages 21 and 23).
18. The section on oil spills should mention that the use of single-hull tanker vessels will be
completely phased out in the U.S. in January 2015.
This information was added to the status report (page 27).
19. In the Environmental Contaminants section, we suggest citing a recent scientific paper
by Desforges et al. (2014) that describes the occurrence of microplastics in coastal
British Columbia and adjacent waters of Washington.
Information from this reference was added to the status report (page 27).
20. Please prevent human disturbance at all remaining nesting sites for Tufted Puffins that
are under WDFW’s jurisdiction.
Nearly all current and former puffin breeding sites in Washington are managed by the U.S.
Fish and Wildlife Service (USFWS) as parts of their national wildlife refuges. These
islands are visited only occasionally by scientists and land managers, and are off-limits to
the general public. However, some former nesting sites in the San Juan Islands may be
subject to unlawful visitation by kayakers and boaters. If WDFW prepares a recovery plan
for puffins (see response to Comment 3), an action would likely be included to work with
the USFWS to minimize this type of visitation.
21. Is it possible that the growing cormorant population is crowding puffins out of their
traditional breeding sites?
There is no evidence of this occurring. The Double-crested Cormorant population has
greatly increased in Washington in recent years, but it is no longer known to breed on any
puffin nesting islands in the state. Pelagic and Brandt’s Cormorants nest on some of the
islands occupied by Tufted Puffins, but both species nest in areas separate from puffins and
thus do not compete for space with puffins. In addition, WDFW biologists have never
observed aggressive interactions between puffins and cormorants on these islands, which
can be an indication of competitive interactions.

January 2015

66

Washington Department of Fish and Wildlife

 Washington State Status Reports, Periodic Status Reviews, Recovery Plans,
and Conservation Plans
Status Reports

Periodic Status Reviews

2015
2007
2005

2015

2005
2004
2002
2000
1999
1999
1999
1999
1998
1998
1998
1998
1998
1997
1997
1997
1997
1993
1993
1993
1993
1993
1993
1993
1993

Tufted Puffin
Bald Eagle
Mazama Pocket Gopher,
Streaked Horned Lark, and
Taylor’s Checkerspot
Aleutian Canada Goose
Killer Whale
Peregrine Falcon
Common Loon
Northern Leopard Frog
Olympic Mudminnow
Mardon Skipper
Lynx Update
Fisher
Margined Sculpin
Pygmy Whitefish
Sharp-tailed Grouse
Sage-grouse
Aleutian Canada Goose
Gray Whale
Olive Ridley Sea Turtle
Oregon Spotted Frog
Larch Mountain Salamander
Lynx
Marbled Murrelet
Oregon Silverspot Butterfly
Pygmy Rabbit
Steller Sea Lion
Western Gray Squirrel
Western Pond Turtle

Steller Sea Lion

Recovery Plans
2012
2011
2007
2006
2004
2004
2003
2002
2001
2001
1999
1996
1995
1995
1995

Columbia Sharp-tailed Grouse
Gray Wolf
Western Gray Squirrel
Fisher
Sea Otter
Greater Sage-Grouse
Pygmy Rabbit: Addendum
Sandhill Crane
Pygmy Rabbit: Addendum
Lynx
Western Pond Turtle
Ferruginous Hawk
Pygmy Rabbit
Upland Sandpiper
Snowy Plover

Conservation Plans
2013

Bats

Reports, reviews, and plans are available on the Washington Department of Fish and Wildlife
website at: http://wdfw.wa.gov/publications/search.php

  

Tufted Pu??in near Smith Island, Washington (Photo Bill Hoglund)

References Reviewed for the Washington State Status Report for the Tufted Puffin
Table B presents the 305 references that are cited in the Washington State Status Report for the
Tufted Puffin. Each reference is categorized for its level of peer review pursuant to section
34.05.271 RCW, which is the codification of Substitute House Bill 2661 that passed the
Washington Legislature in 2014. A key to the review categories under section 34.05.271 RCW
is provided in Table A. References were categorized by Gary Wiles in January 2015.
Individual papers cited in the Washington State Status Report for the Tufted Puffin cover a
number of topics discussed in the report, including information on: 1) the species’ description,
taxonomy, distribution, and biology; 2) habitat requirements; 3) population status and trends; 4)
conservation status and protections; 5) research, monitoring, and restoration activities; and 6)
factors affecting the continued existence of the species.

Table A. Key to 34.05.271 RCW Categories:
Category
Code
34.05.271(1)(c) RCW
i
ii
iii
iv
v

vi

vii
viii

(i) Independent peer review: review is overseen by an independent third party.
(ii) Internal peer review: review by staff internal to the department of fish and wildlife.
(iii) External peer review: review by persons that are external to and selected by the department of
fish and wildlife.
(iv) Open review: documented open public review process that is not limited to invited
organizations or individuals.
(v) Legal and policy document: documents related to the legal framework for the significant agency
action including but not limited to: (A) federal and state statutes; (B) court and hearings board
decisions; (C) federal and state administrative rules and regulations; and (D) policy and regulatory
documents adopted by local governments.
(vi) Data from primary research, monitoring activities, or other sources, but that has not been
incorporated as part of documents reviewed under the processes described in (c)(i), (ii), (iii), and (iv)
of this subsection.
(vii) Records of the best professional judgment of department of fish and wildlife employees or
other individuals.
(viii) Other: Sources of information that do not fit into one of the categories identified in this
subsection (1)(c).

References for the Washington State
Status Report for the Tufted Puffin

1

Washington Department of Fish and Wildlife

 Table B

34.05.271
RCW
Review
Category

Reference
Abdul-Aziz, O. I., N. J. Mantua, and K. W. Myers. 2011. Potential climate change impacts on
thermal habitats of Pacific salmon (Oncorhynchus spp.) in the North Pacific Ocean and
adjacent seas. Canadian Journal of Fisheries and Aquatic Sciences 68:1660-1680.
Addison, B., R. C. Ydenberg, and B. D. Smith. 2007. Tufted puffins (Fratercula cirrhata) respond to
predation danger during colony approach flights. Auk 124:63-70.
Agler, B. A., S. J. Kendall, D. B. Irons, and S. P. Klosiewski. 1999. Declines in marine bird
populations in Prince William Sound, Alaska coincident with a climatic regime shift.
Waterbirds 22:98-103.
Agler, B. A., P. E. Seiser, S. J. Kendall and D. B. Irons. 1994. Marine bird and sea otter populations
of Prince William Sound, Alaska: population trends following the Exxon Valdez oil spill.
Restoration Project No. 93405, Final Report, U.S. Fish and Wildlife Service, Anchorage,
Alaska.
Ainley, D. G., and K. D. Hyrenbach. 2010. Top-down and bottom-up factors affecting seabird
population trends in the California current system (1985–2006). Progress in Oceanography
84:242–254.
Ainley, D. G., and T. J. Lewis. 1974. The history of Farallon Island marine bird populations, 18541972. Condor 76:432-446.
Ainley, D. G., W. J. Sydeman, S. A. Hatch, and U. Wilson. 1994. Seabird population trends along the
west coast of North America: causes and the extent of regional accordance. In J. R. Jehl, Jr.,
and N. K. Johnson, editors. A century of avifaunal change in western North America. Studies
in Avian Biology 15:119-133.
Albers, P. H. 1978. The effects of petroleum on different stages of incubation in bird eggs. Bulletin
of Environmental Contamination and Toxicology 19:624-630.
Alcorn, G. D. 1959. Puffins on the south Grays Harbor beaches. Murrelet 40:21.
Anderson, C. M., M. Mayes, and R. LaBelle. 2012. Update of occurrence rates for offshore oil
spills. OCS Report, BOEM 2012-069, BSEE 2012-069, Bureau of Ocean Energy Management
and Bureau of Safety and Environmental Enforcement, Department of the Interior, Herndon,
Virginia.
Anderson, D. M. 1997. Turning back the harmful red tide. Nature 388:513-514.
Anderson, P. J., and J. F. Piatt. 1999. Community reorganization in the Gulf of Alaska following
ocean climate regime shift. Marine Ecology Progress Series 189:117-123.
Anker-Nilssen, T., R. T. Barrett, and J. V. Krasnov. 1997. Long and short-term responses of seabirds
in the Norwegian and Barents Seas to changes in stocks of prey fish. Pages 683-698 in Forage
fishes in marine ecosystems: proceedings of the international symposium on the role of
forage fishes in marine ecosystems. Alaska Sea Grant College Program AK-SG-97-01,
University of Alaska, Fairbanks, Alaska.
Anthony, J. A., and D. D. Roby. 1997. Variation in lipid content of forage fishes and its effect on
energy provisioning rates to seabird nestlings. Pages 725-729 in Forage fishes in marine
ecosystems: proceedings of the international symposium on the role of forage fishes in
marine ecosystems. Alaska Sea Grant College Program AK-SG-97-01, University of Alaska,
Fairbanks, Alaska.
th
AOU (American Ornithologists’ Union). 1998. Check-list of North American birds. 7 edition.
American Ornithologists’ Union, Washington, D.C.
Aubry, K. B., and S. D. West. 1984. The status of native and introduced mammals on Destruction
Island, Washington. Murrelet 65:80-83.
Audubon Washington. 2004. State of the birds. Audubon Washington, Seattle, Washington.

References for the Washington State
Status Report for the Tufted Puffin

2

i
i
i

vi

i
i

i

i
i
vi
i
i

i

i

i
i
vi

Washington Department of Fish and Wildlife

 Table B

34.05.271
RCW
Review
Category

Reference
Audubon Washington. 2009. State of the birds. Audubon Washington, Seattle, Washington.
Avery-Gomm, S., J. F. Provencher, K. H. Morgan, and D. F. Bertram. 2013. Plastic ingestion in
marine-associated bird species from the eastern North Pacific. Marine Pollution Bulletin
72:257-259.
Bailey, E. P. 1976. Breeding bird distribution and abundance in the Barren Islands, Alaska.
Murrelet 57:2-12.
Bailey, E. P., and N. H. Faust. 1981. Summer distribution and abundance of marine birds and
mammals between Mitrofania and Sutwik Islands south of Alaska Peninsula. Murrelet 62:3442.
Baird, P. H. 1990. Influence of abiotic factors and prey distribution on diet and reproductive
success of three seabird species in Alaska. Ornis Scandinavica 21:224-235.
Baird, P. H. 1991. Optimal foraging and intraspecific competition in the Tufted Puffin. Condor
93:503-515.
Bargmann, G. 1998. Forage fish management plan. Washington Department of Fish and Wildlife,
Olympia, Washington. 77 pp.
Batten, S. D., and A. W. Walne. 2011. Variability in northwards extension of warm water copepods
in the NE Pacific. Journal of Plankton Research 33:1643-1653.
B.C. Conservation Data Centre. 2014. BC species and ecosystems explorer. British Columbia
Ministry of Environment, Victoria, British Columbia. http://a100.gov.bc.ca/pub/eswp/
[accessed December 18, 2014]
Bédard, J. 1969. Adaptive radiation in the Alcidae. Ibis 111:19-98.
Behrenfeld, M. J., R. T. O'Malley, D. A. Siegel, C. R. McClain, J. L. Sarmiento, et al. 2006. Climatedriven trends in contemporary ocean productivity. Nature 444:752-755.
Bent, A. C. 1919. Life history of North American diving birds. Dover Publications, New York, New
York (1963 reprint).
Bertram, D. F., D. L. Mackas, and S. M. McKinnell. 2001. The seasonal cycle revisited: interannual
variation and ecosystem consequences. Progress in Oceanography 49:283-307.
BirdLife International. 2012. Fratercula cirrhata. In IUCN Red List of Threatened Species. Version
2014.3. http://www.iucnredlist.org/ [accessed on December 19, 2014]
Blackburn, G. S., J. M. Hipfner, and R. C. Ydenberg. 2009. Evidence that Tufted Puffins Fratercula
cirrhata use colony overflights to reduce kleptoparasitism risk. Journal of Avian Biology
40:412-418.
Blight, L. K., and A. E. Burger. 1997. Occurrence of plastic particles in seabirds from the Eastern
North Pacific. Marine Pollution Bulletin 34:323-325.
Borstad, G., W. Crawford, J. M. Hipfner, R. Thomson, and K. Hyatt. 2011. Environmental control of
the breeding success of Rhinoceros Auklets at Triangle Island, British Columbia. Marine
Ecology Progress Series 424:285–302.
Bovy, K. 2007. Prehistoric human impacts on waterbirds at Watmough Bay, Washington, USA.
Journal of Island and Coastal Archaeology 2:210-230.
Bowles, J. H. 1922. Birds of Tacoma and vicinity. Murrelet 3:14-16.
Boyd, R. 1999. Indians, fire and the land in the Pacific Northwest. Oregon State University Press,
Corvallis, Oregon.
Branch, T. A., B. M. DeJoseph, L. J. Ray, and C. A. Wagner. 2013. Impacts of ocean acidification on
marine seafood. Trends in Ecology & Evolution 28:178-186.
Brazil, M. A. 1991. The birds of Japan. Smithsonian Institution Press, Washington, D.C. 448 pp.
Bridge, E. S. 2004. The effects of intense wing molt on diving in alcids and potential influences on
the evolution of molt patterns. Journal of Experimental Biology 207:3003-3014.
References for the Washington State
Status Report for the Tufted Puffin

3

vi
i
i
i
i
i
Ii,iii,iv
i
vi
i
i
i
i
i
i
i
i
i
i
i
i
i
i

Washington Department of Fish and Wildlife

 Table B

34.05.271
RCW
Review
Category

Reference
Briggs, K. T., D. H. Varaoujen, W. W. Williams, R. G. Ford, M. L. Bonnell, and J. N. Casey. 1992.
Seabirds of the Oregon and Washington OCS, 1989-1990. Pages 3.142-3.145 in J. J.
Brueggeman, editor. Oregon and Washington marine mammal and seabird surveys: final
report. Pacific OCS Region, Minerals Management Service, U.S. Department of the Interior,
Los Angeles, California.
Brodeur, R. D., J. P. Fisher, R. L. Emmett, C. A. Morgan, and E. Casillas. 2005. Species composition
and community structure of pelagic nekton off Oregon and Washington under variable
oceanographic conditions. Marine Ecology Progress Series 298:41-57.
Brodier, S., B. Pisanu, A. Villers, E. Pettex, M. Lioret, J.-L. Chapuis, and V. Bretagnolle. 2011.
Responses of seabirds to the rabbit eradication on Ile Verte, sub-Antarctic Kerguelen
Archipelago. Animal Conservation 14:459-465.
Burger, A. E., and D. M. Fry. 1993. Effects of oil pollution on seabirds in the Northeast Pacific.
Pages 254-263 in K. Vermeer, K. T. Briggs, K. H. Morgan, and D. Siegel-Causey, editors. The
status, ecology, and conservation of marine birds of the North Pacific. Canadian Wildlife
Service Special Publication, Ottawa, Ontario.
Burger, J., and M. Gochfeld. 2007. Metals and radionuclides in birds and eggs from Amchitka and
Kiska Islands in the Bering Sea/Pacific Ocean ecosystem. Environmental Monitoring and
Assessment 127:105-117.
Burger, J., and M. Gochfeld. 2009a. Comparison of arsenic, cadmium, chromium, lead,
manganese, mercury and selenium in feathers in Bald Eagle (Haliaeetus leucocephalus), and
comparison with Common Eider (Somateria mollissima), Glaucous-winged Gull (Larus
glaucescens), Pigeon Guillemot (Cepphus columba), and Tufted Puffin (Fratercula cirrhata)
from the Aleutian chain of Alaska. Environmental Monitoring and Assessment 152:357-367.
Burger, J., and M. Gochfeld. 2009b. Mercury and other metals in feathers of Common Eider
(Somateria mollissima) and Tufted Puffin (Fratercula cirrhata) from the Aleutian chain of
Alaska. Archives of Environmental Contamination and Toxicology 56:596-606.
Burger, J., M. Gochfeld, D. Kosson, C. W. Powers, et al. 2007. Radionuclides in marine fishes and
birds from Amchitka and Kiska Islands in the Aleutians: establishing a baseline. Health
Physics 92:265-279.
Burrell, G. C. 1980. Some observations on nesting Tufted Puffins, Destruction Island, Washington.
Murrelet 61:92-94.
Byrd, G. V., E. C. Murphy, G. W. Kaiser, A. Y. Kondratyev, and Y. V. Shibaev. 1993. Status and
ecology of offshore fish-feeding alcids (murres and puffins) in the North Pacific. Pages 176186 in K. Vermeer, K. T. Briggs, K. H. Morgan, and D. Siegel-Causey, editors. The status,
ecology, and conservation of marine birds of the North Pacific. Canadian Wildlife Service
Special Publication, Ottawa, Ontario.
Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. Kaiser, and M. C. E. McNall.
1990. The birds of British Columbia. Volume 2: diurnal birds of prey through woodpeckers.
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Carter, H. R., and S. G. Sealy. 2011. Historical breeding records of four species of alcid in British
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Clapp R. B. 1986. A summary of alcid records from Hawaii. Colonial Waterbirds 9:104-107.
Clapp, R. B. and J. B. Giezentanner. 1979. Tufted Puffin from Laysan – first occurrence in the
tropical Pacific. ‘Elepaio 40:120-121.
Clausen, K. K., and P. Clausen. 2014. Forecasting future drowning of coastal waterbird habitats
reveals a major conservation concern. Biological Conservation 171:177-185.

References for the Washington State
Status Report for the Tufted Puffin

4

i

i

i

i

i

i

i

i
i

i

i
i
i
i
i

Washington Department of Fish and Wildlife

 Table B

34.05.271
RCW
Review
Category

Reference
Cody, M. L. 1973. Coexistence, coevolution and convergent evolution in seabird communities.
Ecology 54:31-44.
Couch, L. K. 1929. Introduced European Rabbits in the San Juan Islands, Washington. Journal of
Mammalogy 10:334-336.
Courchamp, F., M. Langlais, and G. Sugihara. 2000. Rabbits killing birds: modeling the
hyperpredation process. Journal of Animal Ecology 69:154-164.
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seabird response to forage fish depletion – one-third for the birds. Science 334:1703-1706.
Davies, W. E., J. M. Hipfner, K. A. Hobson, and R. C. Ydenberg. 2009. Seabird seasonal
trophodynamics: isotopic patterns in a community of Pacific alcids. Marine Ecology Progress
Series 382:211-219.
Dawson, D. A., F. M. Hunter, J. Pandhal, R. Buckland, A. Parham, et al. 2005. Assessment of 17
new Whiskered Auklet (Aethia pygmaea) microsatellite loci in 42 seabirds identifies 5–15
polymorphic markers for each of nine Alcinae species. Molecular Ecology Notes 5:289–297.
Dawson, W. L. 1908a. The new reserves on the Washington Coast. Condor 10:45-49.
Dawson, W. L. 1908b. The bird colonies of the Olympiades. Auk 25:153-166.
Dawson, W. L. 1911. Another fortnight on the Farallones. Condor 13:171-83.
Dawson, W. L. and J. H. Bowles. 1909. The birds of Washington: a complete scientific and popular
account of the 372 species of birds found in the state. Occidental Publishing Company,
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Pacific Ocean, 1976–1985. Marine Pollution Bulletin 18:311-316.
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nets in the North Pacific. Pages 204-211 in K. Vermeer, K. T. Briggs, K. H. Morgan, and D.
Siegel-Causey, editors. The status, ecology, and conservation of marine birds of the North
Pacific. Canadian Wildlife Service Special Publication, Ottawa, Ontario.
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References for the Washington State
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vi
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vi
i

i

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i
i

Washington Department of Fish and Wildlife

 Table B

34.05.271
RCW
Review
Category

Reference
Di Lorenzo, E., N. Schneider, K. M. Cobb, P. J. S. Franks, K. Chhaket, et al. 2008. North Pacific Gyre
Oscillation links ocean climate and ecosystem change. Geophysical Research Letters 35,
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Donald, P. 2010. Facing extinction: the world’s rarest birds and the race to save them. T. & A. D.
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Doney, S. C., M. Ruckelshaus, J. E. Duffy, J. P. Barry, F. Chan, C. A. English, H. M. Galindo, J. M.
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Dragoo, D. E., G. V. Byrd, and D. B. Irons. 2010. Breeding status, population trends and diets of
seabirds in Alaska, 2007. U.S. Fish and Wildlife Service Report AMNWR 2010/08, U.S. Fish
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temperate North Pacific. Pages 241-253 in K. Vermeer, K. T. Briggs, K. H. Morgan, and D.
Siegel-Causey, editors. The status, ecology, and conservation of marine birds of the North
Pacific. Canadian Wildlife Service Special Publication, Ottawa, Ontario.
Emmett, R. L., R. D. Brodeur, T. W. Miller, S. S. Pool, P. J. Bentley, G. K. Krutzikowsky, and J.
McCrae. 2005. Pacific sardine (Sardinops sagax) abundance, distribution, and ecological
relationships in the Pacific Northwest. California Cooperative Oceanic Fisheries
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Etkin, D. S., and J. Neel. 2001. Investing in spill prevention-has it reduced vessel spills and
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fauna and ecosystem processes. ICES Journal of Marine Science 65:414–432.
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Bloxton, B. Galleher, B. Hogoboom, M. Lance, and R. Young. 2011. Marbled Murrelet
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climate change on marine faunas. Trends in Ecology and Evolution 8:361–366.
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variability on the oceanic ecosystems of the NE Pacific. Fisheries Oceanography 7:1-21.

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 Table B

34.05.271
RCW
Review
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Reference
Frazer, D. A. 1974. Breeding biology of the Tufted Puffin on Destruction Island: a brief summary of
the 1974 field season. University of Washington, Seattle, Washington.
Frazer, D. A. 1975. Breeding biology of the Tufted Puffin (Lunda cirrhata): a review. M.S. thesis,
University of Washington, Seattle, Washington.
Frederiksen, M., T. Anker-Nilssen, G. Beaugrand, and S. Wanless. 2013. Climate, copepods and
seabirds in the boreal Northeast Atlantic – current state and future outlook. Global Change
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Friesen, V. L., A. J. Baker, and J. F. Piatt. 1996. Phylogenetic relationships within the Alcidae
(Charadriiformes: Aves) inferred from total molecular evidence. Molecular Biology and
Evolution 13:359-367.
Fry, D. M. 1995. Reproductive effects in birds exposed to pesticides and industrial chemicals.
Environmental Health Perspectives 103:165-171.
Galusha, J. G., B. Vorvick, M. R. Opp, and P. T. Vorvick. 1987. Nesting season censuses of seabirds
on Protection Island, Washington. Murrelet 68:103-107.
Gill, F. 2007. Ornithology. 3rd edition. W. H. Freeman and Co., New York, New York.
Gill, R. E., Jr., and G. A. Sanger. 1979. Tufted Puffins nesting in estuarine habitat. Auk 96:792-794.
Gjerdrum, C. 2004. Parental provisioning and nestling departure decisions: a supplementary
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