Endangered and Threatened Wildlife; Determination on Whether To List the Harbor Seals in Iliamna Lake, Alaska as a Threatened or Endangered Species, 81074-81086 [2016-27690]
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Endangered and Threatened Wildlife;
Determination on Whether To List the
Harbor Seals in Iliamna Lake, Alaska
as a Threatened or Endangered
Species
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
ACTION: Notice of a listing
determination.
AGENCY:
We, NMFS, have completed
our review of the status of eastern North
Pacific harbor seals (Phoca vitulina
richardii) in Iliamna Lake, Alaska. Our
review was in response to a petition to
list these seals as threatened or
endangered under the Endangered
Species Act (ESA). Based on the best
scientific and commercial information
available, we conclude that the seals in
Iliamna Lake do not constitute a species,
subspecies, or distinct population
segment (DPS) under the ESA. As a
result, we conclude that listing the
harbor seals in Iliamna Lake, Alaska is
not warranted.
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SUMMARY:
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This listing determination is
made as of November 17, 2016.
ADDRESSES: This finding and supporting
information are available on our Web
page at: https://
alaskafisheries.noaa.gov/pr/harborseals. Supporting documentation used
in preparing this listing determination is
available for public inspection, by
appointment, during normal business
hours at the office of NMFS Alaska
Region, Protected Resources Division,
709 West 9th Street, Room 461, Juneau,
AK 99801. This documentation includes
the petition, the Biological Review
Team’s DPS report, information
provided by the public and interested
parties, and scientific and commercial
data gathered for the review.
FOR FURTHER INFORMATION CONTACT:
Mandy Migura, NMFS Alaska Region,
(907) 271–1332; Jon Kurland, NMFS
Alaska Region, (907) 586–7638; or Lisa
Manning, NMFS Office of Protected
Resources, (301) 427–8466.
SUPPLEMENTARY INFORMATION:
DATES:
Background
On November 19, 2012, we received
a petition submitted by the Center for
Biological Diversity (CBD) to list the
harbor seals in Iliamna Lake, Alaska as
a threatened or endangered species
under the ESA, and to designate critical
habitat concurrent with listing. CBD
asserted that the harbor seals found in
Iliamna Lake constitute a DPS of Pacific
harbor seals and contended that the
seals in Iliamna Lake face threats
warranting protection as a listed species
under the ESA. Iliamna Lake is the
largest freshwater lake in Alaska and is
connected to the Bristol Bay region of
the Bering Sea by the Kvichak River.
On May 17, 2013 (78 FR 29098), we
found that the petition presented
substantial information indicating that
listing the seals in Iliamna Lake under
the ESA may be warranted, and we
requested comments from the public to
inform our status review, and to help us
determine whether these seals should be
listed as threatened or endangered. To
assist with our status review, we
convened a Biological Review Team
(BRT), composed of federal scientists
with expertise in marine mammal
biology and marine mammal genetics, to
review the available information about
the status of the species, and provide an
assessment regarding the seals in
Iliamna Lake. The BRT compiled
information about the harbor seals in
Iliamna Lake in a DPS Report (Boveng
et al., 2016).
In this notice, we announce our
finding that the petitioned action to list
harbor seals in Iliamna Lake under the
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ESA as either threatened or endangered
is not warranted because the seals do
not constitute a distinct population
segment (DPS) and thus are not a
separate ‘‘species,’’ as the ESA defines
that term. Speficically, while we
conclude that the seals are a discrete
population, the best scientific and
commercial data available suggest that
they are not significant to the greater
taxon to which they belong, i.e., the
eastern North Pacific harbor seal
subspecies (Phoca vitulina richardii).
ESA Statutory, Regulatory, and Policy
Considerations
Section 3 of the ESA defines a
‘‘species’’ as ‘‘any subspecies of fish or
wildlife or plants, and any distinct
population segment of any species of
vertebrate fish or wildlife which
interbreeds when mature.’’ Section 3 of
the ESA further defines an endangered
species as ‘‘any species which is in
danger of extinction throughout all or a
significant portion of its range’’ and a
threatened species as one ‘‘which is
likely to become an endangered species
within the foreseeable future throughout
all or a significant portion of its range.’’
Thus, we interpret an ‘‘endangered
species’’ to be one that is presently in
danger of extinction. A ‘‘threatened
species,’’ on the other hand, is not
presently in danger of extinction, but is
likely to become so in the foreseeable
future. In other words, the primary
statutory difference between a
threatened and endangered species is
the timing of when a species may be in
danger of extinction, either presently
(endangered) or in the foreseeable future
(threatened).
Under section 4(a)(1) of the ESA, we
must determine whether a species is
threatened or endangered because of
any one or a combination of the
following factors: (A) The present or
threatened destruction, modification, or
curtailment of its habitat or range; (B)
overutilization for commercial,
recreational, scientific, or educational
purposes; (C) disease or predation; (D)
inadequacy of existing regulatory
mechanisms; or (E) other natural or
human-made factors affecting its
continued existence. We must make this
determination based solely on the best
scientific and commercial data available
after conducting a review of the status
of the species and taking into account
those efforts being made by states or
foreign governments to protect the
species.
The first step in determining whether
the harbor seals in Iliamna Lake warrant
listing under the ESA is to assess if they
meet the ESA’s definition of ‘‘species.’’
Although there has been speculation
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regarding the taxonomy of the seals in
Iliamna Lake (i.e., whether they are
harbor seals, spotted seals, or hybrids),
recent genetic analyses (O’Corry-Crowe
2013) provide a high degree of
confidence these seals are harbor seals
(Phoca vitulina). The data available are
insufficient to suggest the seals in
Iliamna Lake, Alaska are a separate
subspecies of harbor seal apart from the
subspecies P. v. richardii (Boveng et al.,
2016), which ranges from Mexico to
Alaska. Therefore, we assessed whether
the harbor seals in Iliamna Lake
constitute a distinct population segment
of P. v. richardii.
The U.S. Fish and Wildlife Service
(USFWS) and NMFS (the ‘‘Services’’)
adopted the Policy Regarding the
Recognition of Distinct Vertebrate
Population Segments under the ESA
(the DPS Policy, 61 FR 4722; February
7, 1996) to clarify the Services’
interpretation of the term ‘‘distinct
population segment’’ for the purposes of
listing, delisting, and reclassifying
vertebrates under the ESA. The DPS
Policy establishes two criteria that must
be met for a population or group of
populations to be considered a DPS: (1)
The population segment must be
discrete in relation to the remainder of
the species (or subspecies) to which it
belongs; and (2) the population segment
must be significant to the remainder of
the species (or subspecies) to which it
belongs. In this case, harbor seals in
Iliamna Lake would need to be both
discrete from and significant to the
eastern North Pacific subspecies of
harbor seals (P. v. richardii), to be
designated as a DPS.
If the seals in Iliamna Lake were
found to meet the DPS criteria, we
would then conduct a status review and
determine whether they are threatened
or endangered because of any one or a
combination of the factors from section
4(a)(1) of the ESA. Such a determination
would be based solely on the best
scientific and commercial data
available. Here, because we concluded
that the seal population in Iliamna Lake
is not a DPS, we did not conduct a
status review of the population under
section 4(a)(1) of the ESA.
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Harbor Seal Biology and Life History
Physical Description
Harbor seals (Phoca vitulina) range in
length and size from 1.5–1.9 meters (m)
and 75–180 kilograms (kg) for males,
and 1.4–1.7 m and 60–145 kg for
females, with weights varying
seasonally (Sease 1992). At birth, harbor
seal pups are approximately 0.75–1.0 m
in length and weigh 10–20 kg (Sease
1992). There is a large amount of natural
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variation in harbor seal coats with
coloration ranging from tan/brown to
light gray/black with patterns of spots,
rings, and blotches that vary between
individuals (Shaughnessy and Fay 1977;
Kelly 1981). Variable patterns in seal
coats have been well documented and
may be a result of the age or sex of the
animal, season, location, or the
environment they inhabit (Shaughnessy
and Fay 1977; Kelly 1981; Moss 1992;
Caro et al., 2012). The stage of molting
also has an impact on the appearance of
their coats.
Life History
On average, harbor seals reach sexual
maturity at the age of five for both
females and males; however, females
exhibit a larger range of age at maturity
(Calkins and Pitcher 1979). The
variation depends on population size
and trend, body condition, and prey
resources (Pitcher and Calkin 1979;
Mclaren and Smith 1985; Atkinson
1997). Harbor seals in the eastern North
Pacific subspecies also exhibit natural
variation in the timing of pupping,
ranging from March to September (Bigg
1969; Temte et al., 1991; Searse 1992),
depending in part on general geographic
location. Aerial surveys of harbor seals
in Iliamna Lake since 2010 have
documented that pupping occurs in the
lake, with pups observed during aerial
surveys in June, July, and August (Burns
et al., 2012; Burns et al., 2013; Boveng
et al., 2016; NMML unpubl. data).
Harbor seals molt annually following
pupping (Pitcher and Calkins 1979).
Molting usually lasts 1–2 months,
during which time seals spend a large
amount of time hauled-out (Pitcher and
Calkins 1979; Daniel et al., 2003).
Molting occurs in stages across the
body, affecting coloration and pattern of
the coat throughout the molt.
Harbor seals are considered
opportunistic foragers and feed on a
wide variety of prey found in marine,
estuarine, and fresh waters (Carretta et
al., 2015). Since they inhabit coastal
waters, harbor seal dives are often less
than 50 m and last 2–5 minutes (Bowen
et al., 1999; Frost et al., 2001, 2006)
which influences the prey species
available for foraging. Alaskan harbor
seals have been documented to forage
on pollock, Pacific cod, Pacific sand
lance, sculpins, Pacific salmon, trout,
char, graylings, flatfishes, capelin,
eulachon, smelt, and Pacific herring
(Hobson et al., 1997; Iverson et al., 1997;
Houser et al., 2008; Geiger et al., 2013).
Power and Gregoire (1978) report harbor
seal diet in Lower Seal Lake, Quebec
being dominated by lake and brook
trout. Harbor seals have also been
documented to follow salmon and other
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anadromous fish up rivers and into
freshwater lakes where they may remain
for extended periods (e.g. Bigg 1969a,
1981, and Hoover 1988 as cited in Sease
1992; Middlemas et al., 2006). One of
the largest sockeye salmon populations
in the world run up the Kvichak River
into Iliamna Lake annually in June and
July. Harbor seals have been observed to
follow these fish runs seasonally from
Bristol Bay, although whether those
seals enter Iliamna Lake has not been
documented.
Distribution and Abundance
Harbor seals are one of the most
widespread pinniped species and are
found throughout the northern
hemisphere, ranging from temperate to
polar regions. As of 2008, the
worldwide harbor seal population was
estimated between 350,000 and 500,000
mature individuals (Thompson and
¨ ¨
Harkonen 2008). Currently, there are
five recognized subspecies of harbor
seals: P. v. vitulina in the eastern
Atlantic; P. v. concolor in the western
Atlantic; P. v. mellonae in some lakes
and rivers draining into eastern Hudson
Bay; P. v. richardii in the eastern North
Pacific; and P. v. stejnegeri (also known
as P. v. kurilensis) in the western North
Pacific (Rice 1998; Berta and Churchill
2012).
The harbor seals found in Iliamna
Lake are classified as part of the
subspecies P. v. richardii, also
commonly referred to as eastern North
Pacific harbor seals. Eastern North
Pacific harbor seals range from Mexico
to Alaska (Carretta et al., 2015), with an
estimated abundance of 360,000
individuals (DFO 2010). More than
205,000 harbor seals occur in Alaska
(Muto and Angliss 2015).
Eastern North Pacific harbor seals in
Alaska are divided into 12 separate
stocks under the Marine Mammal
Protection Act; however, these stocks do
not represent taxonomic delineations,
and all 12 stocks are part of the
subspecies P. v. richardii. Harbor seals
in Iliamna Lake are part of the Bristol
Bay stock, which was estimated at
approximately 32,350 individuals based
on a 2011 survey (Muto and Angliss
2015), an increase from the estimated
18,577 seals in 2005 (Allen and Angliss
2014).
Aerial surveys of harbor seals in
Iliamna Lake have primarily been
conducted in the summer and have
consistently documented fewer than 350
animals (Mathisen and Kline 1992;
Small 2001; Withrow and Yano 2009;
Burns et al., 2012; Burns et al., 2013;
NMML unpubl. data). The standard
protocol for harbor seal aerial surveys is
that only seals on land are counted and
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seals in the water are not counted
(Burns et al., 2011; Burns et al., 2013).
It is likely that not all seals haul-out at
the same time and some seals present in
the water were not counted during the
surveys of Iliamna Lake. Thus, the
actual number of seals in Iliamna Lake
at the time of these surveys may have
been greater than the number of seals
reported during the aerial surveys. To
estimate abundance and trends in seal
numbers in Iliamna Lake, a simple
demographic model was developed
(Boveng et al., in prep as reported in
Boveng et al., 2016). That model
indicates that the number of seals in the
lake, about 400, has been relatively
stable from 1984–2013 with little to no
evidence of a trend over the past 5,10,
and 15-year horizons. In 2011,
household surveys of local residents
from six communities in the Iliamna
Lake region were conducted. Based
upon a synthesis of the information
provided by this local traditional
knowledge (LTK) of Iliamna Lake
residents, the population size of seals in
the lake was believed to be
approximately 329 individuals, with a
general belief that the population was
increasing (Burns et al., 2013).
Habitat Use and Movements
Harbor seals typically inhabit nearshore coastal waters, but are well known
for their use of estuaries and rivers, and
have been recorded over 200 kilometers
(km) upstream (see review in COSEWIC
2007). Harbor seals are known to haulout on a variety of natural and manmade
substrates which include beaches,
sandbars, rocks, islands, ice, docks,
piers, and boats. Their varied haul-out
substrates are an example of the
behavioral plasticity of harbor seals to
adapt to a range of environmental
settings and conditions (Komers 1997;
Vincent et al., 2010).
Harbor seals are often described as a
sedentary, non-migratory species, with
considerable site fidelity to one or a few
haul-outs, with large scale movements
being rare. Traditional thinking is that
harbor seals generally stay within 50 km
of a primary haul-out site (e.g., see
Peterson et al., 2012). However, Burns
(2002) states this is a ‘‘gross
oversimplification’’ and instead states
that harbor seals move quite extensively
in some cases, including movements
characterized as ‘‘migrations, juvenile
dispersal, seasonal shifts, shifts related
to breeding activity, responses to seals
habitat exclusion, responses to acute or
chronic disturbance, and immigration/
emigration, occasionally on a relatively
large scale.’’ Satellite tagging studies
document that harbor seals have large
home ranges with haul-out sites that
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vary seasonally and by individual, with
some seals migrating hundreds of km
between breeding and post-breeding
habitats (e.g., Lowry et al., 2001; Lesage
et al., 2004; Peterson et al., 2012;
Womble and Gende 2013). These
studies also report strong evidence of
site fidelity by harbor seals to their
breeding or locations where they were
tagged during summer. In the St.
Lawrence estuary in Canada, over half
of the satellite tagged harbor seals left
their summer haul-out areas once solid
ice formed within the bays of the
estuary, and migrated between 65 km
and 520 km to over-wintering sites
(Lesage et al., 2004). In the Pacific
Northwest region of the United States,
Hardee (2008) reported that harbor seal
movements up to 100 km from the
tagging site occurred most frequently
outside of the breeding season, and that
some adult males made trips in excess
of 200 km roundtrip that lasted 1–8
weeks between April and August.
Hardee (2008) observed long-distance
and long-duration movements by harbor
seals throughout the study period, with
males making multiple roundtrip
movements greater than 200 km that
were not associated with a migratory
over-wintering behavior. Hardee’s
(2008) study, as well as a study of
harbor seals from the Wadden Sea,
Denmark (Tougaard et al., 2003 as cited
in Hardee 2008), contradict the
traditional view that harbor seals reside
in a limited geographic area and do not
leave that home area for extended
periods of time. Peterson et al. (2012)
documented adult male harbor seals in
the Pacific Northwest moving rapidly
between haul-outs, at times traveling
over 100 km in about two days. That
study also concluded that some adult
male harbor seals had secondary haulout sites greater than 100 km from the
primary haul-out site; that the locations
of, and distances between, primary and
secondary haul-outs varied by seal; and
that seasonal migrations over 100 km by
adult male seals were more common
than previously believed. In Alaska,
Lowry et al. (2001) reported juvenile
harbor seal movements of 300–500 km,
and Womble and Gende (2013)
documented extensive migrations of
harbor seals from Glacier Bay during the
post-breeding season, with some
females traveling to Prince William
Sound, a distance up to 900 km one
way. A harbor seal tagged in the Egegik
and Ugashik region of eastern Bristol
Bay traveled in excess of 470 km, and
8 of 14 tagged harbor seals traveled in
excess of 100 km from a major haul-out
site (ADF&G unpubl. data).
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There is also variation in individual
movements of harbor seals within a
population, with some seals traveling
great distances seasonally while others
stay within a smaller area year-round.
Womble and Gende (2013) noted that
some harbor seals in Glacier Bay,
Alaska, were residents year-round
whereas others were migratory. For the
migrating harbor seals, there was a high
degree of site fidelity back to Glacier
Bay the following pupping/breeding
season despite the extensive migration
away from the breeding area during the
post-breeding season (Womble and
Gende 2013). Lesage et al. (2004)
documented that half of the tagged
harbor seals in the St. Lawrence estuary
in Canada left their summer haul-out
areas and migrated up to 520 km to
over-wintering sites, whereas the other
half stayed year-round. Peterson et al.
(2012) concluded that some harbor seals
in the Pacific Northwest had spatially
separated primary and secondary haulouts, while other seals stayed relatively
close to a primary haul-out year-round.
Sharples et al. (2012) documented
highly variable individual harbor seal
movements for seals tagged in the
British Isles. This study also concluded
that region and season better explained
the variation in foraging movements
than the individual seal’s sex, size, and
body condition (Sharples et al. 2012),
suggesting the local habitat conditions
and distance to profitable feeding
grounds may influence the foraging
movements of the seals.
No harbor seals in Iliamna Lake have
been satellite tagged, thus there are no
data available about harbor seals
movements in Iliamna Lake comparable
to those discussed in the preceding
paragraphs. Data on habitat use and
movements of harbor seals in Iliamna
Lake are from aerial surveys
documenting locations where harbor
seals were hauled-out (e.g., Mathisen
and Kline 1992; Small 2001; Withrow
and Yano 2009; Burns et al., 2012;
Burns et al., 2013), and the LTK of
residents, including Alaska Native
subsistence hunters around Iliamna
Lake (e.g., Burns et al., 2013; Van Lanen
et al., 2013). In Iliamna Lake, hauled-out
harbor seals are observed primarily in
the northeastern portion of the lake, but
some local residents report seeing seals
in the southwestern portion of the lake,
especially near the Kvichak River and
Igiugig (Burns et al., 2013). The majority
of aerial surveys of Iliamna Lake were
conducted during the summer/ice-free
season, with a small number of recent
(2010–2013) surveys also flown during
the winter/ice-present season. The
recent aerial surveys documented
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seasonal variations in seal presence and
abundance in the lake, with
significantly greater numbers of seals
observed hauled-out during the summer
pupping and molting periods (e.g., 237
seals observed August 4, 2013) than
during the winter (e.g., 9 seals observed
April 4, 2013) (Burns et al., 2011;
Withrow et al., 2012; Burns et al., 2012;
Burns et al., 2013; NMML unpubl. data).
While harbor seals are known to haulout on ice, recent aerial surveys have
documented few seals hauled-out
during winter surveys in Iliamna Lake.
For example, an aerial survey flown in
April 2010, when the lake was almost
completely frozen-over, documented
only 11 seals; observers reported they
‘‘did not see any areas that could
support the several hundred seals that
have been documented in the summer’’
(Withrow et al., 2011). Another aerial
survey in April 2013 observed only nine
hauled-out seals (NMML unpubl. data).
Although fewer seals are documented
during winter months, there has been
some speculation, primarily by some
local residents (Burns et al., 2013; Van
Lanen et al., 2013), that all the seals
remain in the lake year-round and are
undetectable during winter aerial
surveys. It is possible seals present in
the lake in winter are not observed
because they are either in the water or
they are under the ice in areas with air
pockets, which may become accessible
along shorelines when the lake’s water
level drops after a heavy layer of ice has
formed at the surface. The particular
environmental condition of under-ice
air pockets has been scientifically
documented in the Lacs des Loups
Marins in Canada (Twomey 1939 as
cited in Smith and Horonowitsch 1987;
Smith and Horonowitsch 1987). The
Lacs des Loups Marins are home to
harbor seals in subspecies P. v.
mellonae, who reside in freshwater
lakes year-round and are believed to use
under ice haul-outs when the lakes are
iced-over (Smith and Horonowitsch
1987; Smith 1997; DFO 2016). While
neither this environmental condition
nor the use of under-ice air pockets by
harbor seals have been scientifically
assessed in Iliamna Lake, the use of
under ice air pockets or chambers could
explain why fewer seals are observed in
Iliamna Lake when it is frozen
compared to when it is not. However,
this theory does not explain why only
eight seals were counted in November
2010 (Burns et al., 2011) when the lake
was not iced-over. There currently is no
scientific evidence available to
determine whether air chambers or
haul-outs are used by seals under the ice
in Iliamna Lake during the winter;
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however, local residents have reported
hearing seals under the ice in such
spaces (Burns et al., 2013). Regardless of
the number of seals present in winter,
the aerial surveys provide scientific
evidence of some level of year-round
presence of harbor seals in Iliamna
Lake.
Conclusions drawn from recent aerial
surveys suggest that some harbor seals
may be year-round residents of Iliamna
Lake whereas other harbor seals may
seasonally migrate to and from the lake
(Burns et al., 2011; Withrow et al., 2011;
Burns et al., 2012; Burns et al., 2013).
Some of the LTK regarding the
migration patterns of seals in Iliamna
Lake are inconsistent, and collectively
they do not provide clarity (see Burns et
al., 2013). Some LTK reports indicate
harbor seals migrate between Iliamna
Lake and Bristol Bay and are frequently
seen traversing the Kvichak River (e.g.,
Alvarez 2013; Burns et al., 2013; Igiugig
Tribal Village Council 2013; Mohr 2013;
Wilson 2013), while other reports
indicate that the seals do not migrate
and are present in the lake year-round
(e.g., Burns et al., 2013; Jacko 2013;
Mohr 2013). Local residents around
Iliamna Lake indicate that observations
of harbor seals in the Kvichak River are
typically made beginning in spring,
peak during mid-summer, and decline
to zero in the winter months; however,
some residents of Levelock on the
Kvichak River have observed seals in
the river in the winter (Burns et al.,
2013). This suggests that the Kvichak
River may be used seasonally as a
migration route between Iliamna Lake
and Bristol Bay.
No scientific data are available to
determine whether enough fish remain
in Iliamna Lake to support hundreds of
seals during winter. Some LTK indicates
that the lake may not have sufficient
food available to support the number of
seals observed in summer months on a
year-round basis. A local seal hunter
recently noted that two seals harvested
during two consecutive winters in the
lake had not ‘‘one drop of food in the
stomach or intestines’’ (Burns et al.,
2013). Another seal hunter recollected
shooting a seal in March one year that
was very skinny and had no fat on it,
and speculated that during cold winters
there was inadequate food for the seals
(Burns et al., 2013). However, the
hunter also mentioned that it was very
rare to find a skinny seal in Iliamna
Lake. During our public comment
period we received a comment that
provided calculations of the abundance
of non-salmonid freshwater fish
available during the overwinter period
and indicated that a population of
approximately 300 seals could not be
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81077
sustained on the levels of freshwater
fish available in the winter. We have no
information to support or refute the
calculations provided by the
commenter.
Alternatively, there may be adequate
abundance of prey available in the lake
year-round, but some seals could leave
the lake in winter for other reasons. In
the St. Lawrence estuary, a study of
satellite-tagged harbor seals found that
seals left summer haul-out areas when
solid ice formed within the bays of the
estuary despite ‘‘evidence of high
abundance of potential prey for harbor
seals in the estuary during winter’’
(Lesage et al., 2004). This study
concluded that availability of prey in
winter ‘‘is not the primary factor which
influences the movement and
distribution patterns of harbor seals’’
(Lesage et al., 2004). As discussed
earlier, harbor seals have been
documented to have spatially separated
home ranges which vary seasonally (e.g.
Lowry et al., 2001; Lesage et al., 2004;
Peterson et al., 2012; Womble and
Gende 2013), but also high site fidelity
to breeding locations. Thus, it is
plausible that some harbor seals from
Bristol Bay seasonally follow the salmon
to Iliamna Lake and return to Bristol
Bay for winter, but there are no data
available either to support or refute this
scenario.
Whether seals migrate seasonally
between Iliamna Lake and Bristol Bay
has not been scientifically investigated,
with the exception of a few recent aerial
surveys of Iliamna Lake and the Kvichak
River. Aerial surveys of the Kvichak
River (five complete or partial river
surveys conducted from 2008–2013)
have failed to document harbor seal
presence in the river (Burns et al., 2013),
but it is possible that seals in the river
may have been missed during the
surveys or that the surveys were
conducted when seals were not using
the river. For example, during an aerial
survey in 2011, the survey crew
received a report of seals in a tributary
of the Kvichak River near Kastinak
Flats, but the survey crew was unable to
locate the seals when they flew over the
area approximately 30 minutes later
(Burns et al., 2013; D. Withrow, NMML,
pers. comm.). Additionally, Burns et al.,
(2013) postulated that seals present in
the Kvichak River may not be accounted
for as a result of the survey
methodology, which only counts seals
hauled-out, not those in the water.
Other reports suggest harbor seals are
regularly seen throughout the Kvichak
River (Burns et al., 2013; Van Lanen et
al., 2013; ADF&G unpubl. data). Of 14
harbor seals satellite tagged in Egegik
and Ugashik Bays within eastern Bristol
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‘‘discreteness’’ and ‘‘significance’’
criteria of our DPS policy (61 FR 4722;
February 7, 1996). If a population
segment is found to be discrete and
significant, it is a DPS and is considered
a ‘‘species’’ under the ESA. If the
population is not both discrete and
significant, it does not meet the criteria
for designation as a DPS and does not
qualify as a ‘‘species’’ as defined by the
ESA; thus, we need not evaluate its
status relative to the factors in section
4(a)(1) of the ESA because it cannot be
listed as a threatened or endangered
species. Our assessment first addresses
the discreteness of the harbor seals
found in Iliamna Lake, and then
addresses whether these seals are
significant to P. v. richardii, as these
Subsistence Harvest
terms are defined in our DPS policy (61
Harbor seals are an important
FR 4722; February 7, 1996).
resource for Alaska Native communities
As discussed above, we know from
surrounding Iliamna Lake. Harbor seals
formal scientific studies and LTK that at
are not only a food source, but also
least some harbor seals are present in
provide materials that can be used for
the lake year-round; i.e. are residents of
clothing, handicrafts, and cultural
Iliamna Lake. What is not clear from the
traditions. Reports of harvesting harbor
science or LTK is whether harbor seals
seals by indigenous people around
from Bristol Bay migrate to Iliamna
Iliamna Lake date back to the early
Lake. The BRT considered four
1800s and LTK suggests that seals have
scenarios: (1) The population of seals in
inhabited the lake for many centuries
Iliamna Lake is self-sustaining with
(Fall et al., 2006; Van Lanen 2012; Burns seals being year-round residents of the
et al., 2013). The majority of hunting
lake, and no migration of seals from
occurs during February and March;
Bristol Bay into the lake occurs; (2)
however, some animals have been
there are resident seals in the lake, and
harvested in summer and occasionally
some seals from Bristol Bay migrate to
in winter (Burns et al., 2013). Seven
the lake during the summer, but there is
communities around Iliamna Lake and
no interbreeding of seals from the two
along the Kvichak River were surveyed
regions and the Bristol Bay seals do not
regarding their harvest of marine
stay in the lake during winter; (3)
mammals: Pedro Bay, Pope-Vannoy
Iliamna Lake contains a mix of seals
Landing, Kokhanok, Newhalen, Igiugig,
born in the lake and those born in the
Iliamna, and Levelock (Burns et al.,
marine environment but who migrated
2013). Between 1982 and 2011,
to the lake (either temporarily or
approximately 150 seals were harvested permanently), and these seals are
in Iliamna Lake; however, there is a
interbreeding; or (4) there is no selfmarked difference in the number of
sustaining population of seals in the
seals harvested each of those years
lake and migration is necessary to
(Burns et al., 2013). For instance, there
sustain the population of seals in the
were no reported harvests of seals in
lake. The BRT found three of the four
1982 and 1996, yet 33 were harvested in scenarios to be plausible, favoring
1991. The most recent survey in 2011
explanations 1 and 2, but not ruling out
reported that 44 percent of households
3. None of the BRT members considered
surveyed from these seven communities the forth scenario likely (Boveng et al.,
2016). For our DPS analyses, we
used ‘‘freshwater’’ harbor seal products
recognize that questions remain
and 13 percent used ‘‘saltwater’’ harbor
regarding whether there is migration,
seal products in some capacity,
and references below to seals in or from
resulting from an estimated harvest of
Iliamna Lake are not meant to imply
29 seals (five ‘‘saltwater’’ and 24
that their birth location (either in
‘‘freshwater’’) (Burns et al., 2013).
Iliamna Lake or the marine
Distinct Population Segment (DPS)
environment) is known, but rather are
Assessment
an indication of the seals’ location in
As described above, only species,
Iliamna Lake at time of observation or
subspecies, and DPSs are eligible for
sampling.
listing as a threatened or endangered
Discreteness
species under the ESA. A DPS is a
We first sought to determine whether
population or group of populations of a
the harbor seal population in Iliamna
vertebrate species that meet both the
asabaliauskas on DSK3SPTVN1PROD with NOTICES
Bay in 2000 and 2001, none were
documented in the Kvichak River or
Iliamna Lake (ADF&G unpubl. data).
However, the sample size is too small to
conclude that migration between Bristol
Bay and Iliamna Lake does not occur.
We did not find any scientific evidence
to conclude the harbor seals in Iliamna
Lake constitute a closed population
with no migration between the lake and
marine waters, and the documented
LTK on this question was inconsistent.
In the absence of persuasive evidence of
a closed population, for purposes of our
DPS assessment, we assumed that
harbor seal migration between Iliamna
Lake and Bristol Bay (or beyond) is
possible.
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Lake is discrete in relation to the
remainder of the taxon to which it
belongs (i.e., the eastern North Pacific
harbor seal subspecies, P. v. richardii).
A population segment of a vertebrate
species may be considered discrete if it
satisfies either one of the following
conditions specified in our DPS policy:
‘‘(1) it is markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors.
Quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation; or
(2) it is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the ESA.’’ Because Iliamna
Lake is entirely within the United
States, the second discreteness criterion
identified above is not relevant. Thus,
we focused our assessment of
discreteness on whether the harbor seals
in Iliamna Lake are markedly separated
from other harbor seals in the
subspecies P. v. richardii, with
emphasis on the nearest harbor seal
stock in adjacent Bristol Bay. In
addition to examining four categories of
factors (i.e., physical, physiological,
ecological, or behavioral factors) as
mechanisms with the potential for
providing marked separation by limiting
the dispersal of breeders between
populations, the BRT recognized that
dispersal rates often cannot be directly
measured in natural populations. As
such, the BRT also decided to separately
review the available genetic information
for evidence of separation.
Physical Factors: Iliamna Lake is
located at the base of the Alaska
Peninsula, where it drains through the
Kvichak River into Bristol Bay. Thus,
harbor seal habitat in Iliamna Lake is
separated from the nearest habitat
commonly used by harbor seals in
Bristol Bay by the Kvichak River.
Reports regarding the length of the
Kvichak River vary, with some older
documents reporting the river is
approximately 80 km (50 mi) in length
(e.g., Orth 1971; BLM 2004), whereas
more recent reports suggest it is closer
to 115–120 km (71–75 mi) (e.g.,
Withrow and Yano 2009; Boveng et al.,
2016; validated by a measurement of the
river path between Kogging and Iliamna
Lake using a high resolution
topographic map). The discrepancy in
reported distances of the river could be
explained by changes in the river itself
over time, variances in the starting and
ending measurement points, or by using
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straight-line measurements on a map
versus tracing the path of the river.
Although seals are found
predominantly in the northeast region of
Iliamna Lake, the most recent studies
indicate harbor seals are found
throughout Iliamna Lake, in rivers
draining into the lake (Iliamna,
Newhalen, and Gilbralter rivers), and
throughout the Kvichak River (Alvarez
2013; Burns et al., 2011; Burns et al.,
2012; Burns et al., 2013; Igiugig Tribal
Village Council 2013; Mohr 2013; Van
Lanen et al., 2013; Wilson 2013). The
distance that seals would have to travel
from the lake to Bristol Bay is well
within the known distances that harbor
seals travel (see previous discussion in
‘‘Habitat Use and Movements’’). Thus,
the evidence available does not indicate
that the length of the Kvichak River nor
the distance to the northeast region of
Iliamna Lake (approximately 180 km
from Bristol Bay) would be a physical
barrier separating seals in Iliamna Lake
from those in Bristol Bay.
Physical factors that could impede
harbor seal passage in the Kvichak River
include shallow braided sandbars and
ice cover during winter. Although
poorly adapted for travel on land,
harbor seals in other areas have been
suspected to cross land up to 0.15 km
long and on inclines as steep as 25
degrees to get from one body of water
to another (COSEWIC 2007), so it is
reasonable to assume harbor seals have
the capability to cross shallow braided
sandbars in the Kvichak River.
Millions of sockeye salmon enter
Iliamna Lake from marine waters
annually via the Kvichak River along
with other species of anadromous
salmon. Also, another marine mammal
species has been reported to travel to
Iliamna Lake via the Kvichak River.
Beluga whales, which are less agile and
much larger than harbor seals, have
been documented in the Kvichak River
(Frost et al., 1983; Quakenbush 2002) in
the spring, summer, and fall (Chythlook
and Coiley 1994) and have been
observed near Igiugig (Burns et al.,
2013; Wilson 2013) and in Iliamna Lake
(Mohr 2013). Thus, the available
evidence suggests the Kvichak River is
passable for harbor seals, at least part of
the year when the river is not frozen
over.
Individual BRT members were not in
agreement regarding the scientific
support for discreteness due to physical
factors, but concluded ‘‘no strong
evidence was found either for or against
marked separation by physical barriers
between harbor seals in Iliamna Lake
and those in Bristol Bay’’ (Boveng et al.,
2016). When we considered the best
available information indicating that
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there is access between Iliamna Lake
and Bristol Bay via the Kvichak River,
which is passable at least part of the
year, and that the distance between the
two locations is within documented
migration distances of harbor seals,
along with with the opinion of the BRT,
we concluded that the best available
information does not support a
conclusion that there is separation due
to physical factors. As such, we find
that harbor seals in Iliamna Lake are not
markedly separated from other harbor
seals of the subspecies P. v. richardii as
a consequence of physical factors.
Physiological Factors: Unlike the Lacs
des Loups Marins harbor seals in
Canada, a landlocked population that
lives exclusively in freshwater lakes and
rivers and has documented
physiological differences from the
adjacent harbor seal population in
marine waters (Smith et al., 1994), no
studies exist suggesting there are
statistically significant morphological or
physiological differences between
harbor seals in Iliamna Lake and other
members of the subspecies P. v.
richardii. Consequently, our
discreteness analysis considered other
types of evidence which may suggest
physiological differences. Specifically,
we considered observations obtained
primarily from those with LTK of seals
in Iliamna Lake having a different size,
taste, pelage, and timing of pupping as
compared to seals in Bristol Bay.
The concentration and availability of
salmon to seals in Iliamna Lake in the
summer may account for perceived
differences reported by LTK in size and
taste of seals in Iliamna Lake compared
to seals in Bristol Bay. For example,
several respondents of a recent LTK
survey indicated that the ‘‘physical size
of the seals grows every year following
the salmon runs’’ (Burns et al., 2013),
suggesting high availability and
consumption of energy-rich salmon
results in growth of seals during the
summer. While the well-fed seals may
have experienced salmon-fueled growth,
the flavor of the harvested seals has
been reported to become less desirable
after the salmon runs, which is
reportedly why seals in Iliamna Lake are
not normally hunted in fall (Burns et al.,
2013). The LTK perception of
differences in pelage pattern and
coloration is conflicting (see Burns et
al., 2013), and no formal studies have
been conducted to determine if there are
significant differences in pelage patterns
for harbor seals in Iliamna Lake versus
elsewhere. Burns et al., (2013) speculate
that the timing of the harvest of harbor
seals in relation to the timing of the
annual molt may play a role in the
perceptions of difference in pelage
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81079
texture or coloration. The observed
variances in taste, body size, and pelage
traits are more likely a reflection of
seasonal diet, normal phenotypic
plasticity, and individual variation
rather than an indication that the seals
in Iliamna Lake are physiologically
distinct from those in the adjacent
marine environment.
The timing of pupping for eastern
North Pacific harbor seals ranges from
March to September (Bigg 1969; Temte
et al., 1991; Sease 1992). In Iliamna
Lake, LTK reports about the timing of
pupping are variable, with some reports
of seal pups born on the lake ice during
March and April, and other reports
indicating pups are born during the first
half of June (Burns et al., 2013). LTK
observations of seal pup sightings in
Iliamna Lake ranged from February to
September, with the majority of pup
sightings between April and August
(Burns et al., 2013). Between 2009 and
2013, aerial surveys of Iliamna Lake
documented newborn pups in June,
July, and August (Burns et al., 2013).
Both aerial survey observations and
local resident observations of newborn
seal pups in Iliamna Lake are within the
normal range of pupping dates for the
eastern North Pacific harbor seal
subspecies.
Jemison and Kelly (2001) and
Reijnders et al. (2010) showed that the
timing of harbor seal pupping in the
same location can shift by as much as
several weeks over the course of a few
decades. A review of data from 1975–
2006 for harbor seals in Nanvek Bay,
Alaska, (which is the main location
within Bristol Bay for which harbor seal
pupping data are available) indicates
that the average peak pupping date can
vary by a couple of weeks over just a
few years (e.g., June 18 in 2002 vs. July
3 in 2006; see Table 1 in Boveng et al.,
2016). This observed natural variation
in timing of harbor seal pupping, along
with scarcity of available data, may
account for seemingly conflicting
information in the scientific literature
about the timing of pupping in Iliamna
Lake relative to other harbor seals in
Alaska (e.g., Burns et al., 2013 states
‘‘when compared to Bristol Bay seals
only, the timing of pupping in Iliamna
does not appear to be substantially
delayed’’ versus Withrow et al. (2011)
which states ‘‘Elsewhere in Alaska we
observe harbor seals pupping much
earlier, in May and June’’). According to
the BRT report (Boveng et al., 2016), the
latest peak pupping date estimated for
the Nanvek Bay region of Bristol Bay
was July 5 (1990). Iliamna Lake aerial
surveys flown in 2010, 2011, and 2013
indicate that the earliest peak pupping
date was July 9 (2010). Sparse data
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about pupping dates in both Bristol Bay
and Iliamna Lake lead us to conclude
that while we do not know the precise
timing of peak pupping of harbor seals
in either region, we do know that timing
of peak pupping can vary by a couple
of weeks among years within a given
location. Therefore, an overlap of the
timing of pupping between seals in
Bristol Bay and Iliamna Lake is possible,
even though there may be a 15-day
delay in the average peak pupping date
in Iliamna Lake (July 12) versus the
average peak pupping date in Nanvek
Bay (June 27) (see Boveng et al., 2016).
Burns et al. (2013) also concluded that
compared to Bristol Bay, the timing of
pupping in Iliamna Lake does not
appear to be substantially delayed. A
model developed to estimate the
abundance and trend of harbor seals in
Iliamna Lake (Boveng et al., in prep as
cited in Boveng et al., 2016) predicted
a peak pupping date of July 20 (versus
the July 12 peak pupping date suggested
by a simple average of the dates of
maximum pup counts presented in
Table 1 of the BRT Report); however,
there was substantial imprecision in the
model’s estimate for the peak of pup
counts in the lake.
Individual BRT members were not all
in agreement regarding the degree of
scientific support for discreteness based
upon marked separation due to
physiological factors. Regarding
differences in physiological traits such
as pelage coloration or texture and seal
size and taste, the BRT report stated
‘‘whether any of these differences truly
reflect physiological differences or
separation is not clear, and the BRT was
unaware of any documentation that
these traits are heritable and would
indicate separation or novel genetic
diversity’’ (Boveng et al., 2016).
Regarding physiological separation
based on the notion that pupping in
Iliamna Lake is potentially delayed by
two to six weeks when compared to
nearby populations, the BRT stated,
‘‘The sparsity of information currently
available for Iliamna Lake, imprecision
in determining the timing for any of the
comparison populations, and the length
of the harbor seal pupping period
(approximately 6–10 weeks), reduce the
confidence that can be placed on the
apparent difference’’ (Boveng et al.,
2016).
When we considered all the evidence
currently available to us, including the
lack of direct measures of physiological
factors, the possibility that perceived
differences in seals’ appearance may be
the result of natural individual
variation, the imprecision of estimating
pupping dates due to limited data, the
potential overlap of pupping seasons
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between Iliamna Lake and Bristol Bay,
and the large timeframe (March to
September) for typical pupping times
across the eastern North Pacific harbor
seal taxon, we concluded that the
available information is too weak for us
to make a determination that there is
separation based on physiological
factors. As such, based on the available
evidence, we find that harbor seals in
Iliamna Lake are not markedly separated
from other harbor seals of the
subspecies P. v. richardii as a
consequence of physiological factors.
Ecological Factors: Harbor seals are
known to pursue and aggregate around
concentrations of anadromous prey,
particularly salmon (e.g., London et al.,
2001, Orr et al., 2004, and Wright et al.,
2007, as cited in Peterson et al., 2012;
Middlemas et al., 2006; Hauser et al.,
2008). Changes in distribution of
seasonally abundant prey in the Pacific
Northwest have been suggested as a
possible explanation for seasonal
movements of harbor seals in that area
(Peterson et al., 2012), as harbor seals
may move deliberately to exploit
regions of higher prey availability
(Hardee 2008). In Alaska, movements of
125 km by adult female harbor seals
have coincided with seasonal eulachon
runs in the Copper River Delta (Lowry
et al., 2001). Savarese and Burns (2010)
documented peak harbor seal numbers
coincident with peaks in regional
salmon abundance in the Bering Glacier
region, and contended the salmon
attracted large numbers of harbor seals
to the region. Peterson et al. (2012)
speculated that the observations of
harbor seals using spatially separated
haul-out sites on a seasonal basis may
be related to seasonal changes in prey
distribution and foraging opportunities.
Hauser et al. (2008) examined
foraging by harbor seals in Iliamna Lake
during July and August, when salmon
are very abundant in the lake, and
reported that the seals predominately
fed on large salmonids (salmon, trout,
char, and graylings) during the summer
months. In addition to salmonids,
Hauser et al. (2008) documented
lampreys, smelts, sculpins, whitefishes,
sticklebacks, and other unidentified
prey items in the scat samples of harbor
seals in Iliamna Lake. Thus, harbor seals
in Iliamna Lake appear to be
opportunistic feeders, consistent with
the general pattern of harbor seals
foraging on a wide variety of fish and
invertebrate prey across their range,
with regional differences in diet
diversity (Jemison 2001; COSEWIC
2007). The prey items and seasonal
concentration of salmon in the diet of
seals in Iliamna Lake are consistent with
those documented for harbor seals in
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other freshwater systems. For example,
Middlemas et al. (2006) documented a
summer peak in the contribution of
salmonid prey to the diet of harbor seals
observed in a Scottish river system;
Beck et al. (1970) documented a seal in
Edehon Lake, Canada with both trout
and whitefish in its stomach; and Power
and Gregoire (1978) reported that harbor
seals in lakes ate various freshwater fish
present in the lakes, including trout.
Smith et al. (1996) examined stomachs
of four harbor seals from the Lacs des
Loups Marins which contained in large
part lake whitefish, lake trout, and
brook trout. Scat collected in the
Nanvak Bay region of Bristol Bay also
showed that harbor seals have a diverse
diet, including some of the same types
of prey species consumed in Iliamna
Lake (e.g., salmon, smelts, sculpins) as
well as other prey species (e.g.,
codfishes, herring, squid/octopus)
(Jemison 2001).
Stable isotope analyses of whiskers
and muscle tissue can provide some
insights about harbor seal diets from
several months prior to the date the
samples were collected. Samples
collected from a small number of
subsistence harvested harbor seals from
Iliamna Lake provide preliminary
evidence that those specific seals
consumed freshwater fish during the
previous winter (Burns et al., 2013).
These preliminary data and the typical
timing of ice melt in the Kvichak River
and Iliamna Lake (May–June) suggest
that these samples were most likely
collected from seals which had
overwintered in the lake. However,
these preliminary stable isotope data are
not especially revealing due to the lack
of data on whisker growth rates, tissue
turnover times, and direct measures of
the isotopic signature of potential prey
resources (Burns et al., 2013).
If ecological factors prevented harbor
seals in Iliamna Lake from mixing with
other harbors seals during mating
season, then there could be marked
separation as a result of lack of
opportunities for interbreeding.
However, when considering the timing
of the annual ice melt in the Kvichak
River and Iliamna Lake, the sockeye
salmon runs into Iliamna Lake, and the
presumed mating seasons of seals in
Bristol Bay and in Iliamna Lake, the
BRT concluded that the timing of these
events would not preclude
opportunities for interbreeding by seals
migrating from Bristol Bay to Iliamna
Lake (Boveng et al., 2016).
The BRT members were in general
agreement regarding the degree of
scientific support for discreteness based
upon marked separation due to
ecological factors, and concluded there
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was ‘‘no strong evidence for separation’’
as a result of any of the ecological
factors considered. Based on the
available evidence, we find that harbor
seals in Iliamna Lake are not markedly
separated from other harbor seals of the
subspecies P. v. richardii as a result of
ecological factors.
Behavioral Factors: There are no
scientific or LTK data available to assess
whether mating behaviors (e.g.,
vocalizations or mate attraction
displays) differ for seals in Iliamna Lake
relative to those in Bristol Bay or other
areas of the eastern North Pacific harbor
seal range. Absent data available
regarding mating behaviors of harbor
seals in Iliamna Lake, the BRT
construed the selection of relatively
remote pupping sites in the northeastern
region of Iliamna Lake (nearly 200 km
from pupping sites in Bristol Bay) to be
a behavior, and suggested the selection
of the unusual location was evidence of
some degree of separation, especially
given harbor seals’ site fidelity to
breeding locations. The selection of
distant pupping sites could be
interpreted to mean that harbor seals in
Iliamna Lake are not freely breeding
with harbor seals in Bristol Bay, and
lead to a conclusion there is marked
separation. However, even a small
amount of breeding dispersal from
marine populations of harbor seals into
Iliamna Lake could render the degree of
genetic differentiation insignificant
(Boveng et al., 2016), suggesting there
may not be marked separation. The
available LTK does not resolve this
question, as opinions vary regarding
whether seals in the lake are residents,
migrants, or a mix of both (see Burns et
al., 2013).
Previously we mentioned that harbor
seals commonly follow anadromous
prey into freshwater environments, such
as rivers and lakes. Thus, we do not
consider the mere presence of harbor
seals in Iliamna Lake to be a behavioral
adaptation suggestive of marked
separation from harbor seals in the
marine environment. However, some
Alaska Natives in the Iliamna Lake
region, including subsistence hunters,
have postulated that the seals
overwinter in the lake by using underice air gaps and haul-outs (Burns et al.,
2013), although such winter habitats
have not been documented in Iliamna
Lake. Lack of data complicates a
determination of whether use of underice shelters would be a special, learned
behavioral adaptation that is unique to
harbor seals over-wintering in
freshwater environments, or if this
behavior would be one that any harbor
seal in a similar environment may
adopt. Similar under-ice habitats in the
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Lacs des Loups Marins in Canada have
been suggested as potential harbor seal
lairs or breathing chambers (e.g., Smith
and Horonowitsch 1987; COSEWIC
2007). This, in turn, suggests that use of
such under-ice habitats may be an
example of the behavioral plasticity that
results in harbor seals using a range of
behaviors and habitats in response to
environmental conditions (Komers
1997; Vincent et al., 2010).
The Lacs des Loups Marins harbor
seal population has shown evidence of
modifying typical harbor seal behavior
and adapting to its environment. It is
postulated that, because no pups have
been observed being born on the ice
during that species’ pupping time
period (April, when the lakes are
frozen), the Lacs des Loups Marins
harbor seals have learned and adapted
to their situation by whelping in underice shelters similar to subnivean birth
lairs (snow caves) used by ringed seals
(Consortium Gilles Shooner & Associes
et al., 1991 as cited in Smith 1997). On
the contrary, Burns et al. (2013) include
information from local residents near
Iliamna Lake who suggest some harbor
seal pups may be born in Iliamna Lake
in March and April, when the lake is
still frozen, but pup on the ice, not
under it. Due to this reported on-ice
pupping, even if the harbor seals in
Iliamna Lake utilize under-ice habitats
as shelters or breathing chambers, such
behavior would not be an adaptation
necessary for successful pupping by
seals that use the lake. Thus, unlike the
Lacs des Loups Marins harbor seals, the
evidence suggests that harbor seals in
Iliamna Lake have not developed novel
behaviors to facilitate pupping in a lake
environment.
The BRT members were in general
agreement regarding the degree of
scientific support for discreteness based
upon marked separation due to
behavioral factors, as determined by
selection of pupping locations far from
those in Bristol Bay, and the ambiguity
regarding the degree of migration and
breeding dispersal (if any). Their
judgment suggests behavioral separation
is possible, but the available evidence is
not strong, or is contradicted by other
evidence. Our review of behavioral
factors indicates that the observed
harbor seal behaviors in Iliamna Lake
are not uncommon; harbor seals in
Iliamna Lake have not been documented
to display behaviors outside the range of
normal harbor seal behaviors (e.g., no
unique mating, pupping, or foraging
behaviors reported), although there are
unresolved questions about migration
and use of under ice shelters. There is
no information available to suggest that
harbor seals living in ice conditions
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81081
year-round in a freshwater system
would require different behavioral
adaptations from harbor seals living in
ice conditions in a saltwater or estuarine
system. Despite the lack of these
obvious indications of potential
behavioral separation, we recognize the
possibility that the selection of pupping
locations distant from other known
pupping locations could be construed as
a behavior and indicate marked
separation as a result of the selection of
pupping sites limiting the potential for
interbreeding. Therefore, we find that
the best available evidence is not
conclusive but indicates that harbor
seals in Iliamna Lake may be markedly
separated from other harbor seals of the
subspecies P. v. richardii as a
consequence of behavioral factors.
Genetics: To further consider whether
harbor seals in Iliamna Lake are
markedly separated from other
populations of eastern North Pacific
harbor seals as a consequence of
physical, physiological, ecological, or
behavioral factors, we examined
available genetic evidence which may
be indicative of separation. Genetic
samples available from harbor seals in
Iliamna Lake were compared to genetic
samples available from harbor seals in
the Egegik and Ugashik regions of
eastern Bristol Bay. Bristol Bay has the
nearest concentration of seals to Iliamna
Lake, and the BRT determined ‘‘the
seals in eastern Bristol Bay would be
expected to be the most similar to the
Iliamna Lake seals if there is breeding
dispersal between the two areas, and
therefore would be expected to pose the
most stringent test for demonstrating
discreteness’’ (Boveng et al., 2016).
Genetic samples have been collected
and analyzed from 13 harbor seals in
Iliamna Lake collected in six years from
1996 through 2012. The mitochondrial
DNA (mtDNA) analysis revealed that 11
of 13 seals sampled from Iliamna Lake
exhibited the same mtDNA haplotype
(O’Corry-Crowe 2013), meaning all 11
seals had the same group of genes
inherited from their female parent. The
remaining two DNA samples did not
yield results for this test. This specific
mtDNA haplotype (Pvit-Hap#7) is the
most common haplotype found in
harbor seals sampled from Bristol Bay
and is observed in roughly 21 percent of
harbor seals from the Egegik and
Ugashik regions of Bristol Bay (Burns et
al., 2013; O’Corry-Crowe 2013). Thus,
this haplotype is not unique to harbor
seals in Iliamna Lake.
The identification of only one mtDNA
haplotype in harbor seals from Iliamna
Lake appears to suggest unusually low
genetic diversity. For comparison, 76
harbor seals sampled from the Egegik
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and Ugashik regions of eastern Bristol
Bay exhibited 33 different mtDNA
haplotypes (O’Corry-Crowe 2013; Burns
et al., 2013). If seals from the Egegik and
Ugashik regions were immigrating into
the lake and staying year-round, there
would be almost an 80 percent
likelihood that one of the other mtDNA
haplotypes, not Pvit-Hap#7, would be
seen in samples collected from Iliamna
Lake (O’Corry-Crowe 2013). However,
because mtDNA is inherited from the
mother, mtDNA diversity analysis
cannot determine if male seals are
migrating to and from the lake and
breeding with resident female seals.
Hardee (2008) recognized similar
limitations of mtDNA given
observations of male harbor seals in the
Pacific Northwest traveling larger
distances than previously believed,
possibly to mate in a separate
geographic region before returning to
their home site. Therefore, conclusive
results about the level of genetic
diversity require analyses using nuclear
DNA (nDNA; which also provides
information from the male parent), and
more formal analyses of mtDNA with
statistical comparisons to harbor seals
sampled from other regions within the
range of the taxon (O’Corry-Crowe
2013). These more stringent data
regarding genetic diversity do not exist.
In addition to examining the existing
genetic diversity of the samples,
analyses were conducted to examine the
extent of genetic differentiation between
harbor seals sampled in Iliamna Lake
from those sampled in the Egegik and
Ugashik regions of eastern Bristol Bay.
The results of analyses examining
genetic differentiation using both
mtDNA and nDNA suggest that the
harbor seals sampled in Iliamna Lake
were genetically differentiated from
harbor seals sampled in the Egegik and
Ugashik regions of eastern Bristol Bay
(Burns et al., 2013; O’Corry-Crowe
2013). The results of these analyses also
suggest that male and female-mediated
dispersal between the Egegik and
Ugashik regions of eastern Bristol Bay
and Iliamna Lake was restricted (Burns
et al., 2013; O’Corry-Crowe 2013).
Although no directed comparisons were
conducted between Iliamna Lake
samples and genetic samples collected
from harbor seals in other areas of
Bristol Bay or other portions of the
range of the taxon, the measure of
mtDNA genetic differentiation between
seals in Iliamna Lake and those in
eastern Bristol Bay yielded results
showing substantially greater genetic
differentiation than all previous
pairwise comparisons between the other
major centers of harbor seal abundance
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in Alaska (O’Corry-Crowe 2012; Boveng
et al., 2016). These genetic
differentiation results are suggestive of
the presence of a small, isolated
population of harbor seals in Iliamna
Lake.
O’Corry-Crowe (2013) identifies
several limitations of the findings for
the Iliamna Lake samples. He cautions
that the sample size is extremely small
and that questions regarding the
patterns of kinship among the collected
samples remain unresolved (i.e., if some
of the samples were from related
individuals, then the data could be
skewed and not representative of a
random sampling of the population),
and indicates that genetic differentiation
may be enhanced in small populations
when there is a rapid rate of genetic
drift, even when there is continued gene
flow. Although the 13 genetic samples
from seals in Iliamna Lake were
collected between 1996 and 2012, most
samples were collected during months
when seasonal migrants would not be
expected to be in the lake, thus the
power to detect seasonal migrants may
be low. Conversely, the timing of the
samples may be benficial for
considering if the resident seals in the
lake are discrete from their marine
counterparts because for most samples
seasonal migrants would not be
expected to be present in the lake.
O’Corry-Crowe (2013) also provides
recommendations for future genetic
research to resolve lingering issues,
including analyzing 20 microsatellite
loci (only 9–11 loci were analyzed) and
updating the techniques used for the
analyses to newer technologies, which
would increase the power to resolve
genetic questions. We also note that the
tests for genetic differentiation
compared the Iliamna Lake samples
solely against samples collected from
the Egegik and Ugashik regions of
eastern Bristol Bay. Thus, the samples
used for the comparison group may not
be representative of all the seals that
could migrate to Iliamna Lake.
The genetic data available suggest the
harbor seals sampled in Iliamna Lake
have low mtDNA diversity, possess the
most common mtDNA haplotype found
in Bristol Bay harbor seals, and are
genetically differentiated from harbor
seals sampled in the Egegik and Ugashik
regions of eastern Bristol Bay. Given the
concerns about the limited nature of the
available genetic information previously
discussed here and by O’Corry-Crowe
(2013), ambiguity remains regarding the
degree of separation, and hence
discreteness, of harbor seals in Iliamna
Lake. However, in the absence of more
samples collected from a greater number
of seals in Iliamna Lake and the Kvichak
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River, to include the potential migration
season, and/or completion of additional
tests such as those recommended by
O’Corry-Crowe (2013), we consider the
existing genetic results to be the best
available data upon which to base our
determination. These genetic results
support a decision that harbor seals in
Iliamna Lake are markedly separated
from harbor seals in eastern Bristol Bay,
and by assumption, from the remainder
of the taxon.
Discreteness Conclusion
We find the available evidence for
discreteness based on physical,
physiological, or ecological factors to be
unconvincing. The available evidence
based on behavioral factors is not
conclusive, but the selection of pupping
locations distant from other known
pupping locations could be construed as
a behavior and indicate marked
separation as a result of the selection of
pupping sites limiting the potential for
interbreeding. The strongest evidence
for discreteness derives from 13 genetic
samples collected from seals in Iliamna
Lake. Analyses of these samples
strongly indicate the seals from Iliamna
Lake are genetically differentiated from
seals sampled in two locations within
Bristol Bay (Ugashik and Egegik), the
nearest concentration of seals to Iliamna
Lake with genetic data available.
Genetic comparisons of samples for the
entire taxon do not exist, but this region
within Bristol Bay was expected to
provide the most stringent comparison
for discreteness if there is breeding
dispersal between the two regions. The
BRT was in strong agreement that the
genetic data reflect marked separation,
although the BRT acknowledged that
the mechanism of such separation is
unknown and the data are limited. It is
possible that the limited available
genetic data may accurately represent
the situation in both Iliamna Lake and
all of Bristol Bay, or that additional
genetic analysis from P. v. richardii
animals sampled from elsewhere in
their range or from additional seals in
Iliamna Lake, could result in a different
conclusion. Nonetheless, the best
available genetic information leads us to
conclude that some portion, and
perhaps all, of the harbor seals in
Iliamna Lake likely constitute a resident
population that is genetically
differentiated from harbor seals in
eastern Bristol Bay, and thus meet the
criteria for consideration as a discrete
entity per our DPS policy (61 FR 4722;
February 7, 1996).
Significance
Having determined that resident seals
from Iliamna Lake are likely discrete, at
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least from harbor seals in the Egegik and
Ugashik regions of nearby Bristol Bay,
we next sought to determine whether
they are significant to the P. v. richardii
subspecies.
In carrying out the significance
examination per our DPS policy (61 FR
4722; February 7, 1996), we are to
consider available scientific evidence of
the population’s importance to the
taxon to which it belongs. This
consideration may include, but is not
limited to, the following: (1) Persistence
of the discrete population segment in an
ecological setting unusual or unique for
the taxon; (2) evidence that loss of the
discrete population segment would
result in a significant gap in the range
of the taxon; (3) evidence that the
discrete population segment represents
the only surviving natural occurrence of
a taxon that may be more abundant
elsewhere as an introduced population
outside its historic range; or (4)
evidence that the discrete population
segment differs markedly from other
populations of the species in its genetic
characteristics.
This determination, however, is
highly fact specific and may consider
factors besides those enumerated above.
Further, significance of the discrete
population segment is not necessarily
determined by existence of one of these
classes of information standing alone.
Information analyzed under these and
any other applicable considerations is
evaluated relative to the biological and
ecological importance of the discrete
population to the taxon as a whole.
Accordingly, all relevant and available
biological and ecological information is
analyzed. As we explained in the DPS
policy, ‘‘the principal significance to be
considered in a potential DPS will be
the significance to the taxon to which it
belongs’’ (61 FR 4722, 4724; February 7,
1996). Finally,we assessed the biological
and ecological significance of the seals
in Iliamna Lake to the P. v. richardii (the
eastern North Pacific harbor seal) taxon
in light of Congressional guidance that
the authority to list DPSs be used
‘‘sparingly’’ while conserving the
genetic diversity of the species (see
Senate Report 151, 96th Congress, 1st
Session).
Persistence in an Unusual or Unique
Ecological Setting: In assessing the
‘‘persistence of the discrete population
segment in an ecological setting unusual
or unique for the taxon,’’ we considered
whether specific characteristics of the
Iliamna Lake environment are unusual
or unique; whether persistence in the
Iliamna Lake environment is unusual or
unique; and whether there are
adaptations as a result of persistence in
an unusual or unique environment
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which would result in the discrete
population being biologically or
ecologically significant to the taxon P. v.
richardii.
The diet of harbor seals in Iliamna
Lake is consistent with what we would
expect for the species occupying a
freshwater system dominated by
anadromous salmon. Hauser et al.
(2008) indicate that harbor seals in
Iliamna Lake consumed large amounts
of sockeye salmon when they were
seasonally abundant, and also fed on
trout, char, graylings, lampreys, smelts,
sculpins, whitefishes, sticklebacks, and
other unidentified prey items. Burns et
al. (2013) examined eight harbor seal
stomachs collected from seals harvested
from Iliamna Lake in 2011 and 2012;
only three had identifiable prey items
and the remaining five stomach were
either empty, only had worms, or had
unidentifiable contents. An examination
of the identifiable prey items found that
these seals had consumed small or
young salmonids (salmon and/or trout),
threespine stickleback, and Arctic
grayling or lake whitefish (Burns et al.,
2013). The variety and types of prey
items in the diet of these sampled seals
in Iliamna Lake reflects harbor seals
being opportunistic feeders (Carretta et
al., 2015), and the available data suggest
no unusual or unique prey for the
habitat occupied.
We also considered whether the
habitat available for use by seals in
Iliamna Lake is unusual or unique.
Harbor seals commonly use reefs, sand
and gravel beaches, sand and mud bars,
island beaches, and ice (glacial ice, pan
ice, sea ice, or icebergs) as haul-out
sites. Harbor seals in Iliamna Lake are
known to haul-out on rocky and sandy
substrates, sand bars, small islands, and
ice near pressure cracks or polynas
(Burns et al., 2011; Burns et al., 2012).
None of these haul-out substrates are
unique or unusual for harbor seals.
Harbor seals in Iliamna Lake are
reported to pup both on ice (Burns et al.
2013) and other haul-outs in the absence
of ice. There is no evidence of seals in
Iliamna Lake pupping in air pockets
beneath the ice, which would be
unusual. Such use has been
hypothesized for the harbor seals in the
Lacs des Loups Marins (Consortium
Gilles Shooner & Associes et al. 1991 as
cited in Smith 1997; DFO 2016).
According to LTK, pupping in Iliamna
Lake likely occurs at island beaches or
sandbars in the northeastern portion of
the lake, which is consistent with the
types of substrates upon which aerial
surveys documented pups (i.e., on lowlying islands and sand spits; Burns et
al., 2013). Nothing suggests that harbor
seals in Iliamna Lake display unusual or
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81083
unique pupping behaviors (including
habitat usage).
Smith and Horonowitsch (1987)
studied the ice at one location within
the Lacs des Loups Marins and
documented what they refer to as
‘‘shoreline ice-steps’’ which they
speculated could be used as breathing
chambers for over-wintering seals in the
lake. LTK suggests the presence and use
of similar under-ice haul-outs in Iliamna
Lake (Burns et al., 2013). While this
would represent unusual habitat use for
harbor seals in general, and unique
habitat for harbor seals of P. v. richardii,
it would be consistent with the general
observation that harbor seals exhibit
wide variation in habitat use, rather
than being indicative of an adaptation
by seals in Iliamna Lake that would be
significant to the P. v. richardii taxon as
a whole (see further discussion of
habitat adaptation below).
Harbor seals have the broadest
distribution and occur in more different
habitats than any other pinniped species
(Burns 2002; COSEWIC 2007), and are
frequently and commonly observed in
freshwater systems (Burns 2002).
Mansfield (1967) provides information
about sightings of harbor seals in rivers
and lakes in Arctic Canada (referencing
Doutt 1942 and Harper 1961 for detailed
summaries of Arctic harbor seals’
freshwater distribution), indicating that
harbor seals have ‘‘a strong liking for
fresh water’’ and are often found in
estuaries and freshwater habitats ‘‘far
from the sea.’’ Beck et al., (1970) report
harbor seals in the Thlewiaza River
system and associated lakes west of
Hudson Bay. Smith et al. (1994) and
Smith (1997) provide an extensive list of
reports of harbor seals documented in
freshwater systems. Smith et al. (1996)
conducted analyses involving both the
Lacs des Loups Marins harbor seals as
well as a second group of ‘‘lacustrine’’
harbor seals from Kasegalik Lake in
Canada’s Northwest Territory.
Middlemas et al. (2006) provide
documentation of harbor seals in a
Scottish river system. The Committee on
the Status of Endangered Wildlife in
Canada (COSEWIC) reports that harbor
seals occasionally ascend the St.
Lawrence River to the Great Lakes
(COSEWIC 2007). In the Bristol Bay
region, harbor seals have been observed
in other lakes in addition to Iliamna
Lake, such as Lake Becharof and Naknek
Lake (Mathisen and Kline 1992). Thus,
the presence of harbor seals in
freshwater systems or lakes, including
Iliamna Lake, is not unusual or unique
for the species.
Year-round persistence of harbor seals
in a lake is less common. Besides the
unknown number of harbor seals
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occupying Iliamna Lake through the
winter, the Lacs des Loups Marins
harbor seals are the only other
documented instance of harbor seals
persisting in freshwater systems yearround. However, a review of available
literature suggests the possibility this
scenario may be more prevalent than
just these two groups of harbor seals.
For example, Mansfield (1967) states
that the population of freshwater harbor
seals in the Upper and Lower Seal Lakes
east of Hudson Bay (a.k.a. the Lacs des
Loups Marins) is not unique given
reports of harbor seals found in other
freshwater systems of Canada. Beck et
al. (1970) postulated that harbor seals
may live in the Thlewiaza River and
associated lakes year-round, and
documented a pup in the Edehon Lake,
leading them to conclude that harbor
seal reproduction is successful in that
freshwater habitat. Beck et al. (1970)
also concluded that individual seals in
those lakes may be born and spend most
or all of their lives in freshwater, but
there was no reason to believe they were
an isolated population. In Alaska,
winter aerial surveys led Savarese and
Burns (2010) to suggest that harbor seals
are present year-round in Vitus Lake, a
tidally-influenced lake near the Bering
Glacier. No pups were documented
during that study and diet and genetic
data indicated seals from various stocks
moved into Vitus Lake to take advantage
of local salmon runs (Savarese and
Burns 2010). These reports of potential
year-round presence of harbor seals in
various freshwater systems are sporadic,
and do not confirm self-sustaining
populations exist in those other
freshwater systems. Therefore, it is
reasonable to conclude that the yearround persistence of a discrete
population of harbor seals in the
freshwater environment of Iliamna Lake
is at least unusual, if not unique, to the
P. v. richardii harbor seal taxon.
The BRT considered whether the
persistence of the population of harbor
seals in this setting is important to the
taxon as a whole (see discussion in
Boveng et al., 2016). Specifically, the
BRT considered whether harbor seals in
Iliamna Lake exhibit any adaptations to
the environment which would be
biologically or ecologically significant to
the P. v. richardii harbor seal taxon. The
evidence of such adaptations is not
necessarily required to demonstrate
significance; however, the BRT
examined such evidence here in light of
harbor seals’ widespread and diverse
habitat and diet. The BRT considered
the physiology of the seals in Iliamna
Lake as well as their over-wintering
strategy as possible indicators of
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adaptations of potential importance for
the taxon.
As previously discussed, some local
residents of the Iliamna Lake region
have suggested they think the harbor
seals harvested from Iliamna Lake taste,
look, or feel different (e.g., seals are
fatter; pelage is softer) from those
harvested in the marine environment
(Burns et al., 2013). There was,
however, a lack of consensus regarding
the perceived differences (e.g., some say
seals from Iliamna Lake are darker than
marine counterparts, others say the seals
are lighter) among the local residents
interviewed. Moreover, attributes such
as fatness and softness of the coat, or the
way the seals taste when consumed, are
not necessarily inherited traits and
could be acquired during time spent in
the lake. Unlike other lake seal species,
there are no data available to document
whether morphological (e.g.,
craniometric) differences exist; if such
morphological differences are present,
they are not distinct enough to be
generally recognized in traditional
knowledge of Alaska Native residents in
the area (see discussion in Boveng et al.,
2016). There is no evidence to suggest
these reported physical differences in
fatness, softness, or taste are adaptations
that would convey significance of these
seals to the taxon.
The use of air gaps under the ice in
winter is a potential adaptation to
freshwater life in sub-Arctic regions,
and is only documented among harbor
seals in one location (P. v. mellonae of
Lacs des Loups Marins). Whether the
use of under-ice shelters would be a true
adaptation to a freshwater environment
which freezes over, or would simply be
a response to habitat conditions that
may be used by any harbor seal exposed
to those conditions, remains uncertain.
On the importance of this particular
behavior relative to significance of seals
in Iliamna Lake to the P. v. richardii
subspecies, the BRT concluded any
assessment would ‘‘be in the realm of
judgment or even speculation’’ (Boveng
et al.,2016). Even though harbor seals in
Iliamna Lake cope with the extensive
ice cover in winter, there is no
indication they have adapted or
modified their breeding, whelping, or
pup-rearing behaviors in a manner
unusual for, or of significance to, the
taxon.
The BRT members were in strong
agreement that harbor seals persisting
year-round and breeding in a freshwater
lake that freezes over almost completely
nearly every year is unique for the
subspecies P. v. richardii, and unusual
for the harbor seal species. However,
there was a lack of consensus amongst
BRT members whether the available
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evidence reflects physical, life-history,
or other adaptations as a result of
persisting in an unusual or unique
ecological setting which would make
the harbor seal population in Iliamna
Lake biologically or ecologically
significant to the broader taxon. The
discrepancies in opinion stemmed from
‘‘differences in assessing the weights of
several lines of qualitative and indirect
evidence’’ (Boveng et al., 2016). The
BRT also concluded (1) seals from the
marine population would be able to
persist in the Iliamna Lake setting, and
(2) even if seals from the marine
population were unable to persist in
Iliamna Lake, the ‘‘lack of ‘ecological
exchangeability’ is not important to the
persistence of the taxon as a whole’’
(Boveng et al., 2016). Ultimately, the
BRT’s assessment favored ‘‘a conclusion
that the evidence does not support
significance’’ (Boveng et al., 2016). We
agree that persistence of a population of
harbor seals in the unusual or unique
ecological setting of Iliamna Lake in and
of itself does not confer significance of
that population to the taxon. The
absence of evidence suggesting the
harbor seals in Iliamna Lake have
adaptations to their environment which
would benefit the taxon to which they
belong leads us to determine that the
persistence of a population of harbor
seals in Iliamna Lake is not significant
to the subspecies P. v. richardii.
Evidence That Loss Would Result in
Significant Gap in Range: Eastern North
Pacific harbor seals range from Mexico
northward along the coastlines of the
continental U.S. and Canada and much
of Alaska. In Alaska, harbor seals of this
subspecies are distributed almost
continuously throughout the southern
coastal waters in the region surrounding
Iliamna Lake. In assessing whether the
loss of harbor seals in Iliamna Lake
would result in a significant gap in the
range, we considered a scenario
whereby all the seals in the lake were
extirpated and there was no migration
into the lake, either because there is no
migration currently occurring or
because a future physical barrier
prevents migration. Given the extensive
and continuous range of the eastern
North Pacific harbor seals, the loss of
the small proportion of habitat in
Iliamna Lake would not result in a
significant gap in the range.
Furthermore, the evidence indicating
possible seasonal movement of some
harbor seals from Bristol Bay to Iliamna
Lake suggests that the habitat in this
portion of the range could be
reoccupied.
The loss of harbor seals in Iliamna
Lake would not have a detrimental
impact to other harbor seal populations
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that comprise the subspecies P. v.
richardii, as this is not an interstitial
population of harbor seals whose loss
would isolate another population from
the main group. Additionally, there are
only an estimated 400 harbor seals in
Iliamna Lake (Boveng et al., 2016), so
this population represents a minute
fraction of the total population ofeastern
North Pacific harbor seals, estimated at
360,000 (DFO 2010).
The BRT was in strong agreement that
the evidence is clear that the loss of the
Iliamna Lake segment would not result
in a significant gap in the range of the
taxon, and we agree.
Evidence of Only Surviving Natural
Occurrence: Harbor seals in taxon P. v.
richardii are currently found throughout
their historic range along the coasts
from Baja California, Mexico, northward
to Alaska, and west through the Gulf of
Alaska and Aleutian Islands, and in the
Bering Sea north to Cape Newenham
and the Pribilof Islands. There are no
known introductions of this species to
any place outside its historic range, thus
it is naturally occurring wherever it
occurs. The BRT was unanimous in its
assessment that harbor seals in Iliamna
Lake are not the only surviving natural
occurrence of the taxon. We concur in
that determination.
Evidence of Marked Difference in
Genetic Characteristics: As discussed
above, the limited genetic data available
from seals in Iliamna Lake indicate 11
of 13 (2 samples did not yield results)
sampled seals had the same mtDNA
haplotype, an indication of possible low
genetic diversity (O’Corry-Crowe 2013).
Unlike the Lacs des Loups Marins
harbor seals, which exhibit mtDNA
haplotypes that are only found in seals
from the Lacs des Loups Marins (Smith
1997), the single mtDNA haplotype
exhibited in the harbor seals in Iliamna
Lake is not unique to Iliamna Lake.
Rather, it is the most common mtDNA
haplotype found in samples from harbor
seals in Bristol Bay (O’Corry-Crowe
2013; Van Lanen et al., 2013). One
plausible explanation for the single
haplotype found in all the harbor seal
samples from Iliamna Lake is that these
seals are simply a genetic subset of seals
from Bristol Bay, and have lost rather
than gained substantial amounts of
genetic diversity since isolation. An
alternative explanation is the seals in
Iliamna Lake have been isolated a long
time, during which they may have
accumulated genetic differences at other
loci (not currently examined) via
mutation, especially for loci under
selective pressure (i.e., adaptation).
However, as previously discussed, only
a small number of genetic loci were
tested and the sample size was small, so
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the reason for a single mtDNA
haplotype is undeterminable at this
time. We conclude that the best
scientific and commercial data
available, a single mtDNA haplotype
which is commonly found in other
populations of the taxon and the data
used to assess discreteness of the
population, do not indicate that harbor
seals in Iliamna Lake have novel genes
which could be significant to the taxon
as a whole.
There is no strong evidence to
indicate the existence of phenotypic
differences between harbor seals in
Iliamna Lake and those in other portions
of the taxon’s range. Although there
have been some LTK reports that the
seals in Iliamna Lake may taste different
or have pelage of varying appearance
from seals in Bristol Bay, there have
been no studies assessing whether these
perceived differences are the result of
significant differences in genetics. The
BRT members did not reach consensus
regarding this issue, with a slight
preponderance of opinion favoring the
conclusion that the genetic
characteristics of seals in Iliamna Lake
did not convey significance to these
seals in regards to P. v. richardii. Some
members considered the data available
as mostly insufficient for drawing a
conclusion regarding significance, and
some considered the evidence against
significance slightly more persuasive
than the evidence for significance.
Accordingly, we find that the genetic
characteristics (i.e., mtDNA haplotype)
found in seals from Iliamna Lake do not
differ markedly from those found in
Bristol Bay and therefore determine that
the best available genetic data, albeit
limited, supports a conclusion that
harbor seals in Iliamna Lake do not have
genetic characteristics that are
significant to the taxon as a whole.
Overall Significance to the Taxon: We
considered several factors that could
indicate whether harbor seals in Iliamna
Lake may be biologically and
ecologically significant to the taxon as a
whole. Of the four factors delinated in
the 1996 DPS policy, we conclude that
there is evidence of only one: The
population persists in an unusual or
unique setting for the taxon. As we
explained in our policy, ‘‘occurrence in
an unusual ecological setting is
potentially an indication that a
population segment represents a
significant resource of the kind sought
to be conserved by the’’ ESA and in
‘‘any actual case of a DPS recognized in
part on this basis, the Services will
describe in detail the nature of this
significance when accepting a petition
or proposing a rule’’ (61 FR at 4724).
While year-round persistence in the
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81085
freshwater environment of Iliamna Lake
is unique to the taxon P. v. richardii and
unusual for the entire species, we agree
with the BRT (Boveng et al., 2016) that
the best scientific and commercial data
available are limited and suggest that
the persistence of the seals in Iliamna
Lake is not significant to the taxon as a
whole. The loss of the Iliamna Lake
segment would not result in a gap in the
range of the taxon, and the harbor seals
in Iliamna Lake are not the only
surviving natural occurrence of the
taxon; thus harbor seals in Iliamna Lake
do not demonstrate significance to the
taxon based on these factors. Further,
available genetic data suggest that
harbor seals in Iliamna Lake are not
significant to the larger taxon. Although
the best available genetic data indicate
that at least some of the seals in Iliamna
Lake are distinct from harbor seals in
the eastern regions of nearby Bristol
Bay, the genetic characteristics (e.g., the
single mtDNA observed in samples from
seals in Iliamna Lake is the most
common haplotype found in seals frim
Bristol Bay) do not appear to differ in
ways that would convey significance to
the P. v. richardii subspecies.
Individual BRT members were not in
agreement regarding the degree of
scientific support overall for or against
the significance of seals in Iliamna Lake
to the P. v. richardii subspecies, but
stated ‘‘the slight majority judgment
against significance of the population
segment . . . summarized a diversity of
views about how much weight to place
on the various lines of mostly weak and
qualitative evidence’’ and that ‘‘the
evidence itself must be characterized as
mostly indirect, qualitative rather than
quantitative, and equivocal for the
purpose of demonstrating biological or
ecological importance to the broader
taxon’’ (Boveng et al., 2016). Taking into
consideration the totality of all the
information discussed above regarding
the possible significance of harbor seals
in Iliamna Lake to the P. v. richardii
taxon, including the qualitative and
equivocal nature of the available
information, along with the guidance
from legislative history to identify DPSs
‘‘sparingly,’’ we find that the available
evidence supports a conclusion that the
harbor seals in Iliamna Lake are not
significant to the remainder of the
taxon.
DPS Conclusion
Based on the best scientific and
commercial data available, we find the
evidence for marked separation of
harbor seals in Iliamna Lake from the
remainder of the taxon based on
physical, physiological, ecological or
behavioral factors to be unconvincing or
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81086
Federal Register / Vol. 81, No. 222 / Thursday, November 17, 2016 / Notices
asabaliauskas on DSK3SPTVN1PROD with NOTICES
weak. The strongest support for marked
separation comes from the best available
genetic data which, although limited
and preliminary, support a conclusion
that at least some of the harbor seals in
Iliamna Lake are likely isolated from
harbor seals in the Egegik and Ugashik
regions of eastern Bristol Bay. Thus, we
conclude that the harbor seal population
in Iliamna Lake is separated from other
populations of the taxon and meet the
discreteness criterion of our DPS policy
(61 FR 4722; February 7, 1996).
Per the second component of our DPS
Policy, we are to consider available
scientific evidence of the discrete
population’s importance to the taxon to
which it belongs (61 FR 4722; February
7, 1996). Our review of the best
available information suggests the only
characteristic which may make this
population of harbor seals unique
within its taxon is the fact that they
persist year-round in a freshwater
system which freezes over to some
degree in most winters. While that
characteristic is unique within the
subspecies P. v. richardii, we
determined such persistence is not
biologically or ecologically important to
the taxon as a whole. Furthermore, the
information available supports a
conclusion that loss of this population
would not be detrimental to the
persistence of the taxon or constitute a
gap in the range of the taxon; this
population is not the only natural
surviving population; and there are no
unique genetic characteristics conveying
significance of this population to the
taxon. After reviewing the best available
data as they apply to the significance
criterion, we conclude that the harbor
seals in Iliamna Lake are not significant
to the taxon P. v. richardii.
Under our DPS Policy, both the
discreteness and significance elements
must be met to qualify as a DPS. Our
review has determined that the seals
persisting year-round in Iliamna Lake
are discrete but not significant;
therefore, the harbor seals in Iliamna
Lake do not qualify as a DPS and are not
a listable entity under the ESA.
Finding
In assessing whether the actions in
the petition are warranted, we reviewed
the best available scientific and
commercial information available,
including the BRT report, the petition
and literature cited in the petition,
published and grey literature relevant to
the topic, correspondence with experts
in academic and government
institutions, documentation of LTK, and
public comments. On the basis of this
review, we have determined that harbor
seals in Iliamna Lake meet the criteria
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for discreteness but do not meet the
criteria for significance. As such, the
harbor seals in Iliamna Lake do not meet
all the criteria necessary to constitute a
DPS, and thus are not a listable entity
under the ESA. Therefore, we find that
the petitioned actions to list the harbor
seals in Iliamna Lake as a threatened or
endangered species under the ESA, and
to designate critical habitat, are not
warranted.
In our 90-day finding (78 FR 29098;
May 17, 2013), we indicated we were
commencing a status review of the
harbor seals in Iliamna Lake. To assist
our evaluation of whether the seals in
Iliamna Lake constitute a DPS, the BRT
prepared a report which compiled
background information about the
harbor seals in Iliamna Lake and
evaluated the scientific information
relevant to the DPS criteria (Boveng et
al., 2016). Upon our determination that
the DPS criteria were not met and the
seals in Iliamna Lake are not a ‘‘species’’
under the ESA, there is no need to
complete the status review by
conducting a threats assessment or
extinction risk assessment in light of the
factors in section 4(a)(1) of the ESA.
In some instances, where we find a
petitioned action is not warranted
because the petitioned population does
not constitute a ‘‘species’’ under the
ESA, we have initiated a status review
of a related or larger population (e.g.,
the 12-month determination that the
petitioned action to list Lynn Canal
Pacific herring was not warranted,
followed by a status review of the
Southeast Alaska population of Pacific
herring; 73 FR 19824; April 11, 2008).
Here, the scope of the petition was
limited to the seals in Iliamna Lake, and
since the most recent abundance data
for the Bristol Bay harbor seal stock (the
stock that includes seals in Iliamna
Lake) indicates this stock increased
from an estimated 18,577 seals in 2005
to an estimated 32,350 seals in 2011
(Allen and Angliss 2014; Muto and
Angliss 2015), we are not initiating a
status review of the Bristol Bay harbor
seal stock at this time.
References
A complete list of all references cited
herein is available upon request (see
ADDRESSES).
Authority
The authority for this action is the
Endangered Species act of 1973, as
amended (16 U.S.C. 1531 et seq.).
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Dated: November 10, 2016.
Samuel D. Rauch, III,
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2016–27690 Filed 11–16–16; 8:45 am]
BILLING CODE 3510–22–P
CONSUMER PRODUCT SAFETY
COMMISSION
[CPSC Docket No. 17–C0001]
PetSmart, Inc., Provisional Acceptance
of a Settlement Agreement and Order
Consumer Product Safety
Commission
ACTION: Notice.
AGENCY:
It is the policy of the
Commission to publish settlements
which it provisionally accepts under the
Consumer Product Safety Act in the
Federal Register in accordance with the
terms of the Consumer Product Safety
Commission’s regulations. Published
below is a provisionally-accepted
Settlement Agreement with PetSmart,
Inc., containing a civil penalty in the
amount of four million, two hundred
fifty thousand dollars ($4,250,000)
within thirty (30) days of service of the
Commission’s final Order accepting the
Settlement Agreement.
DATES: Any interested person may ask
the Commission not to accept this
agreement or otherwise comment on its
contents by filing a written request with
the Office of the Secretary by December
2, 2016.
ADDRESSES: Persons wishing to
comment on this Settlement Agreement
should send written comments to the
Comment 17–C0001, Office of the
Secretary, Consumer Product Safety
Commission, 4330 East-West Highway,
Room 820, Bethesda, Maryland 20814–
4408.
FOR FURTHER INFORMATION CONTACT:
Philip Z. Brown, Trial Attorney,
Division of Compliance, Office of the
General Counsel, Consumer Product
Safety Commission, 4330 East-West
Highway, Bethesda, Maryland 20814–
4408; telephone (301) 504–7645.
SUPPLEMENTARY INFORMATION: The text of
the Agreement and Order appears
below.1
SUMMARY:
1 The Commission voted (4–1) to provisionally
accept the Settlement Agreement and Order
regarding PetSmart, Inc. Chairman Kaye,
Commissioner Adler, Commissioner Robinson and
Commissioner Mohorovic voted to provisionally
accept the Settlement Agreement and Order.
Commissioner Buerkle voted to reject the
Settlement Agreement and Order.
E:\FR\FM\17NON1.SGM
17NON1
Agencies
[Federal Register Volume 81, Number 222 (Thursday, November 17, 2016)]
[Notices]
[Pages 81074-81086]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2016-27690]
-----------------------------------------------------------------------
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
[Docket No. 121120640-6943-02]
RIN 0648-XC365
Endangered and Threatened Wildlife; Determination on Whether To
List the Harbor Seals in Iliamna Lake, Alaska as a Threatened or
Endangered Species
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Commerce.
ACTION: Notice of a listing determination.
-----------------------------------------------------------------------
SUMMARY: We, NMFS, have completed our review of the status of eastern
North Pacific harbor seals (Phoca vitulina richardii) in Iliamna Lake,
Alaska. Our review was in response to a petition to list these seals as
threatened or endangered under the Endangered Species Act (ESA). Based
on the best scientific and commercial information available, we
conclude that the seals in Iliamna Lake do not constitute a species,
subspecies, or distinct population segment (DPS) under the ESA. As a
result, we conclude that listing the harbor seals in Iliamna Lake,
Alaska is not warranted.
DATES: This listing determination is made as of November 17, 2016.
ADDRESSES: This finding and supporting information are available on our
Web page at: https://alaskafisheries.noaa.gov/pr/harbor-seals.
Supporting documentation used in preparing this listing determination
is available for public inspection, by appointment, during normal
business hours at the office of NMFS Alaska Region, Protected Resources
Division, 709 West 9th Street, Room 461, Juneau, AK 99801. This
documentation includes the petition, the Biological Review Team's DPS
report, information provided by the public and interested parties, and
scientific and commercial data gathered for the review.
FOR FURTHER INFORMATION CONTACT: Mandy Migura, NMFS Alaska Region,
(907) 271-1332; Jon Kurland, NMFS Alaska Region, (907) 586-7638; or
Lisa Manning, NMFS Office of Protected Resources, (301) 427-8466.
SUPPLEMENTARY INFORMATION:
Background
On November 19, 2012, we received a petition submitted by the
Center for Biological Diversity (CBD) to list the harbor seals in
Iliamna Lake, Alaska as a threatened or endangered species under the
ESA, and to designate critical habitat concurrent with listing. CBD
asserted that the harbor seals found in Iliamna Lake constitute a DPS
of Pacific harbor seals and contended that the seals in Iliamna Lake
face threats warranting protection as a listed species under the ESA.
Iliamna Lake is the largest freshwater lake in Alaska and is connected
to the Bristol Bay region of the Bering Sea by the Kvichak River.
On May 17, 2013 (78 FR 29098), we found that the petition presented
substantial information indicating that listing the seals in Iliamna
Lake under the ESA may be warranted, and we requested comments from the
public to inform our status review, and to help us determine whether
these seals should be listed as threatened or endangered. To assist
with our status review, we convened a Biological Review Team (BRT),
composed of federal scientists with expertise in marine mammal biology
and marine mammal genetics, to review the available information about
the status of the species, and provide an assessment regarding the
seals in Iliamna Lake. The BRT compiled information about the harbor
seals in Iliamna Lake in a DPS Report (Boveng et al., 2016).
In this notice, we announce our finding that the petitioned action
to list harbor seals in Iliamna Lake under the ESA as either threatened
or endangered is not warranted because the seals do not constitute a
distinct population segment (DPS) and thus are not a separate
``species,'' as the ESA defines that term. Speficically, while we
conclude that the seals are a discrete population, the best scientific
and commercial data available suggest that they are not significant to
the greater taxon to which they belong, i.e., the eastern North Pacific
harbor seal subspecies (Phoca vitulina richardii).
ESA Statutory, Regulatory, and Policy Considerations
Section 3 of the ESA defines a ``species'' as ``any subspecies of
fish or wildlife or plants, and any distinct population segment of any
species of vertebrate fish or wildlife which interbreeds when mature.''
Section 3 of the ESA further defines an endangered species as ``any
species which is in danger of extinction throughout all or a
significant portion of its range'' and a threatened species as one
``which is likely to become an endangered species within the
foreseeable future throughout all or a significant portion of its
range.'' Thus, we interpret an ``endangered species'' to be one that is
presently in danger of extinction. A ``threatened species,'' on the
other hand, is not presently in danger of extinction, but is likely to
become so in the foreseeable future. In other words, the primary
statutory difference between a threatened and endangered species is the
timing of when a species may be in danger of extinction, either
presently (endangered) or in the foreseeable future (threatened).
Under section 4(a)(1) of the ESA, we must determine whether a
species is threatened or endangered because of any one or a combination
of the following factors: (A) The present or threatened destruction,
modification, or curtailment of its habitat or range; (B)
overutilization for commercial, recreational, scientific, or
educational purposes; (C) disease or predation; (D) inadequacy of
existing regulatory mechanisms; or (E) other natural or human-made
factors affecting its continued existence. We must make this
determination based solely on the best scientific and commercial data
available after conducting a review of the status of the species and
taking into account those efforts being made by states or foreign
governments to protect the species.
The first step in determining whether the harbor seals in Iliamna
Lake warrant listing under the ESA is to assess if they meet the ESA's
definition of ``species.'' Although there has been speculation
[[Page 81075]]
regarding the taxonomy of the seals in Iliamna Lake (i.e., whether they
are harbor seals, spotted seals, or hybrids), recent genetic analyses
(O'Corry-Crowe 2013) provide a high degree of confidence these seals
are harbor seals (Phoca vitulina). The data available are insufficient
to suggest the seals in Iliamna Lake, Alaska are a separate subspecies
of harbor seal apart from the subspecies P. v. richardii (Boveng et
al., 2016), which ranges from Mexico to Alaska. Therefore, we assessed
whether the harbor seals in Iliamna Lake constitute a distinct
population segment of P. v. richardii.
The U.S. Fish and Wildlife Service (USFWS) and NMFS (the
``Services'') adopted the Policy Regarding the Recognition of Distinct
Vertebrate Population Segments under the ESA (the DPS Policy, 61 FR
4722; February 7, 1996) to clarify the Services' interpretation of the
term ``distinct population segment'' for the purposes of listing,
delisting, and reclassifying vertebrates under the ESA. The DPS Policy
establishes two criteria that must be met for a population or group of
populations to be considered a DPS: (1) The population segment must be
discrete in relation to the remainder of the species (or subspecies) to
which it belongs; and (2) the population segment must be significant to
the remainder of the species (or subspecies) to which it belongs. In
this case, harbor seals in Iliamna Lake would need to be both discrete
from and significant to the eastern North Pacific subspecies of harbor
seals (P. v. richardii), to be designated as a DPS.
If the seals in Iliamna Lake were found to meet the DPS criteria,
we would then conduct a status review and determine whether they are
threatened or endangered because of any one or a combination of the
factors from section 4(a)(1) of the ESA. Such a determination would be
based solely on the best scientific and commercial data available.
Here, because we concluded that the seal population in Iliamna Lake is
not a DPS, we did not conduct a status review of the population under
section 4(a)(1) of the ESA.
Harbor Seal Biology and Life History
Physical Description
Harbor seals (Phoca vitulina) range in length and size from 1.5-1.9
meters (m) and 75-180 kilograms (kg) for males, and 1.4-1.7 m and 60-
145 kg for females, with weights varying seasonally (Sease 1992). At
birth, harbor seal pups are approximately 0.75-1.0 m in length and
weigh 10-20 kg (Sease 1992). There is a large amount of natural
variation in harbor seal coats with coloration ranging from tan/brown
to light gray/black with patterns of spots, rings, and blotches that
vary between individuals (Shaughnessy and Fay 1977; Kelly 1981).
Variable patterns in seal coats have been well documented and may be a
result of the age or sex of the animal, season, location, or the
environment they inhabit (Shaughnessy and Fay 1977; Kelly 1981; Moss
1992; Caro et al., 2012). The stage of molting also has an impact on
the appearance of their coats.
Life History
On average, harbor seals reach sexual maturity at the age of five
for both females and males; however, females exhibit a larger range of
age at maturity (Calkins and Pitcher 1979). The variation depends on
population size and trend, body condition, and prey resources (Pitcher
and Calkin 1979; Mclaren and Smith 1985; Atkinson 1997). Harbor seals
in the eastern North Pacific subspecies also exhibit natural variation
in the timing of pupping, ranging from March to September (Bigg 1969;
Temte et al., 1991; Searse 1992), depending in part on general
geographic location. Aerial surveys of harbor seals in Iliamna Lake
since 2010 have documented that pupping occurs in the lake, with pups
observed during aerial surveys in June, July, and August (Burns et al.,
2012; Burns et al., 2013; Boveng et al., 2016; NMML unpubl. data).
Harbor seals molt annually following pupping (Pitcher and Calkins
1979). Molting usually lasts 1-2 months, during which time seals spend
a large amount of time hauled-out (Pitcher and Calkins 1979; Daniel et
al., 2003). Molting occurs in stages across the body, affecting
coloration and pattern of the coat throughout the molt.
Harbor seals are considered opportunistic foragers and feed on a
wide variety of prey found in marine, estuarine, and fresh waters
(Carretta et al., 2015). Since they inhabit coastal waters, harbor seal
dives are often less than 50 m and last 2-5 minutes (Bowen et al.,
1999; Frost et al., 2001, 2006) which influences the prey species
available for foraging. Alaskan harbor seals have been documented to
forage on pollock, Pacific cod, Pacific sand lance, sculpins, Pacific
salmon, trout, char, graylings, flatfishes, capelin, eulachon, smelt,
and Pacific herring (Hobson et al., 1997; Iverson et al., 1997; Houser
et al., 2008; Geiger et al., 2013). Power and Gregoire (1978) report
harbor seal diet in Lower Seal Lake, Quebec being dominated by lake and
brook trout. Harbor seals have also been documented to follow salmon
and other anadromous fish up rivers and into freshwater lakes where
they may remain for extended periods (e.g. Bigg 1969a, 1981, and Hoover
1988 as cited in Sease 1992; Middlemas et al., 2006). One of the
largest sockeye salmon populations in the world run up the Kvichak
River into Iliamna Lake annually in June and July. Harbor seals have
been observed to follow these fish runs seasonally from Bristol Bay,
although whether those seals enter Iliamna Lake has not been
documented.
Distribution and Abundance
Harbor seals are one of the most widespread pinniped species and
are found throughout the northern hemisphere, ranging from temperate to
polar regions. As of 2008, the worldwide harbor seal population was
estimated between 350,000 and 500,000 mature individuals (Thompson and
H[auml]rk[ouml]nen 2008). Currently, there are five recognized
subspecies of harbor seals: P. v. vitulina in the eastern Atlantic; P.
v. concolor in the western Atlantic; P. v. mellonae in some lakes and
rivers draining into eastern Hudson Bay; P. v. richardii in the eastern
North Pacific; and P. v. stejnegeri (also known as P. v. kurilensis) in
the western North Pacific (Rice 1998; Berta and Churchill 2012).
The harbor seals found in Iliamna Lake are classified as part of
the subspecies P. v. richardii, also commonly referred to as eastern
North Pacific harbor seals. Eastern North Pacific harbor seals range
from Mexico to Alaska (Carretta et al., 2015), with an estimated
abundance of 360,000 individuals (DFO 2010). More than 205,000 harbor
seals occur in Alaska (Muto and Angliss 2015).
Eastern North Pacific harbor seals in Alaska are divided into 12
separate stocks under the Marine Mammal Protection Act; however, these
stocks do not represent taxonomic delineations, and all 12 stocks are
part of the subspecies P. v. richardii. Harbor seals in Iliamna Lake
are part of the Bristol Bay stock, which was estimated at approximately
32,350 individuals based on a 2011 survey (Muto and Angliss 2015), an
increase from the estimated 18,577 seals in 2005 (Allen and Angliss
2014).
Aerial surveys of harbor seals in Iliamna Lake have primarily been
conducted in the summer and have consistently documented fewer than 350
animals (Mathisen and Kline 1992; Small 2001; Withrow and Yano 2009;
Burns et al., 2012; Burns et al., 2013; NMML unpubl. data). The
standard protocol for harbor seal aerial surveys is that only seals on
land are counted and
[[Page 81076]]
seals in the water are not counted (Burns et al., 2011; Burns et al.,
2013). It is likely that not all seals haul-out at the same time and
some seals present in the water were not counted during the surveys of
Iliamna Lake. Thus, the actual number of seals in Iliamna Lake at the
time of these surveys may have been greater than the number of seals
reported during the aerial surveys. To estimate abundance and trends in
seal numbers in Iliamna Lake, a simple demographic model was developed
(Boveng et al., in prep as reported in Boveng et al., 2016). That model
indicates that the number of seals in the lake, about 400, has been
relatively stable from 1984-2013 with little to no evidence of a trend
over the past 5,10, and 15-year horizons. In 2011, household surveys of
local residents from six communities in the Iliamna Lake region were
conducted. Based upon a synthesis of the information provided by this
local traditional knowledge (LTK) of Iliamna Lake residents, the
population size of seals in the lake was believed to be approximately
329 individuals, with a general belief that the population was
increasing (Burns et al., 2013).
Habitat Use and Movements
Harbor seals typically inhabit near-shore coastal waters, but are
well known for their use of estuaries and rivers, and have been
recorded over 200 kilometers (km) upstream (see review in COSEWIC
2007). Harbor seals are known to haul-out on a variety of natural and
manmade substrates which include beaches, sandbars, rocks, islands,
ice, docks, piers, and boats. Their varied haul-out substrates are an
example of the behavioral plasticity of harbor seals to adapt to a
range of environmental settings and conditions (Komers 1997; Vincent et
al., 2010).
Harbor seals are often described as a sedentary, non-migratory
species, with considerable site fidelity to one or a few haul-outs,
with large scale movements being rare. Traditional thinking is that
harbor seals generally stay within 50 km of a primary haul-out site
(e.g., see Peterson et al., 2012). However, Burns (2002) states this is
a ``gross oversimplification'' and instead states that harbor seals
move quite extensively in some cases, including movements characterized
as ``migrations, juvenile dispersal, seasonal shifts, shifts related to
breeding activity, responses to seals habitat exclusion, responses to
acute or chronic disturbance, and immigration/emigration, occasionally
on a relatively large scale.'' Satellite tagging studies document that
harbor seals have large home ranges with haul-out sites that vary
seasonally and by individual, with some seals migrating hundreds of km
between breeding and post-breeding habitats (e.g., Lowry et al., 2001;
Lesage et al., 2004; Peterson et al., 2012; Womble and Gende 2013).
These studies also report strong evidence of site fidelity by harbor
seals to their breeding or locations where they were tagged during
summer. In the St. Lawrence estuary in Canada, over half of the
satellite tagged harbor seals left their summer haul-out areas once
solid ice formed within the bays of the estuary, and migrated between
65 km and 520 km to over-wintering sites (Lesage et al., 2004). In the
Pacific Northwest region of the United States, Hardee (2008) reported
that harbor seal movements up to 100 km from the tagging site occurred
most frequently outside of the breeding season, and that some adult
males made trips in excess of 200 km roundtrip that lasted 1-8 weeks
between April and August. Hardee (2008) observed long-distance and
long-duration movements by harbor seals throughout the study period,
with males making multiple roundtrip movements greater than 200 km that
were not associated with a migratory over-wintering behavior. Hardee's
(2008) study, as well as a study of harbor seals from the Wadden Sea,
Denmark (Tougaard et al., 2003 as cited in Hardee 2008), contradict the
traditional view that harbor seals reside in a limited geographic area
and do not leave that home area for extended periods of time. Peterson
et al. (2012) documented adult male harbor seals in the Pacific
Northwest moving rapidly between haul-outs, at times traveling over 100
km in about two days. That study also concluded that some adult male
harbor seals had secondary haul-out sites greater than 100 km from the
primary haul-out site; that the locations of, and distances between,
primary and secondary haul-outs varied by seal; and that seasonal
migrations over 100 km by adult male seals were more common than
previously believed. In Alaska, Lowry et al. (2001) reported juvenile
harbor seal movements of 300-500 km, and Womble and Gende (2013)
documented extensive migrations of harbor seals from Glacier Bay during
the post-breeding season, with some females traveling to Prince William
Sound, a distance up to 900 km one way. A harbor seal tagged in the
Egegik and Ugashik region of eastern Bristol Bay traveled in excess of
470 km, and 8 of 14 tagged harbor seals traveled in excess of 100 km
from a major haul-out site (ADF&G unpubl. data).
There is also variation in individual movements of harbor seals
within a population, with some seals traveling great distances
seasonally while others stay within a smaller area year-round. Womble
and Gende (2013) noted that some harbor seals in Glacier Bay, Alaska,
were residents year-round whereas others were migratory. For the
migrating harbor seals, there was a high degree of site fidelity back
to Glacier Bay the following pupping/breeding season despite the
extensive migration away from the breeding area during the post-
breeding season (Womble and Gende 2013). Lesage et al. (2004)
documented that half of the tagged harbor seals in the St. Lawrence
estuary in Canada left their summer haul-out areas and migrated up to
520 km to over-wintering sites, whereas the other half stayed year-
round. Peterson et al. (2012) concluded that some harbor seals in the
Pacific Northwest had spatially separated primary and secondary haul-
outs, while other seals stayed relatively close to a primary haul-out
year-round. Sharples et al. (2012) documented highly variable
individual harbor seal movements for seals tagged in the British Isles.
This study also concluded that region and season better explained the
variation in foraging movements than the individual seal's sex, size,
and body condition (Sharples et al. 2012), suggesting the local habitat
conditions and distance to profitable feeding grounds may influence the
foraging movements of the seals.
No harbor seals in Iliamna Lake have been satellite tagged, thus
there are no data available about harbor seals movements in Iliamna
Lake comparable to those discussed in the preceding paragraphs. Data on
habitat use and movements of harbor seals in Iliamna Lake are from
aerial surveys documenting locations where harbor seals were hauled-out
(e.g., Mathisen and Kline 1992; Small 2001; Withrow and Yano 2009;
Burns et al., 2012; Burns et al., 2013), and the LTK of residents,
including Alaska Native subsistence hunters around Iliamna Lake (e.g.,
Burns et al., 2013; Van Lanen et al., 2013). In Iliamna Lake, hauled-
out harbor seals are observed primarily in the northeastern portion of
the lake, but some local residents report seeing seals in the
southwestern portion of the lake, especially near the Kvichak River and
Igiugig (Burns et al., 2013). The majority of aerial surveys of Iliamna
Lake were conducted during the summer/ice-free season, with a small
number of recent (2010-2013) surveys also flown during the winter/ice-
present season. The recent aerial surveys documented
[[Page 81077]]
seasonal variations in seal presence and abundance in the lake, with
significantly greater numbers of seals observed hauled-out during the
summer pupping and molting periods (e.g., 237 seals observed August 4,
2013) than during the winter (e.g., 9 seals observed April 4, 2013)
(Burns et al., 2011; Withrow et al., 2012; Burns et al., 2012; Burns et
al., 2013; NMML unpubl. data).
While harbor seals are known to haul-out on ice, recent aerial
surveys have documented few seals hauled-out during winter surveys in
Iliamna Lake. For example, an aerial survey flown in April 2010, when
the lake was almost completely frozen-over, documented only 11 seals;
observers reported they ``did not see any areas that could support the
several hundred seals that have been documented in the summer''
(Withrow et al., 2011). Another aerial survey in April 2013 observed
only nine hauled-out seals (NMML unpubl. data). Although fewer seals
are documented during winter months, there has been some speculation,
primarily by some local residents (Burns et al., 2013; Van Lanen et
al., 2013), that all the seals remain in the lake year-round and are
undetectable during winter aerial surveys. It is possible seals present
in the lake in winter are not observed because they are either in the
water or they are under the ice in areas with air pockets, which may
become accessible along shorelines when the lake's water level drops
after a heavy layer of ice has formed at the surface. The particular
environmental condition of under-ice air pockets has been
scientifically documented in the Lacs des Loups Marins in Canada
(Twomey 1939 as cited in Smith and Horonowitsch 1987; Smith and
Horonowitsch 1987). The Lacs des Loups Marins are home to harbor seals
in subspecies P. v. mellonae, who reside in freshwater lakes year-round
and are believed to use under ice haul-outs when the lakes are iced-
over (Smith and Horonowitsch 1987; Smith 1997; DFO 2016). While neither
this environmental condition nor the use of under-ice air pockets by
harbor seals have been scientifically assessed in Iliamna Lake, the use
of under ice air pockets or chambers could explain why fewer seals are
observed in Iliamna Lake when it is frozen compared to when it is not.
However, this theory does not explain why only eight seals were counted
in November 2010 (Burns et al., 2011) when the lake was not iced-over.
There currently is no scientific evidence available to determine
whether air chambers or haul-outs are used by seals under the ice in
Iliamna Lake during the winter; however, local residents have reported
hearing seals under the ice in such spaces (Burns et al., 2013).
Regardless of the number of seals present in winter, the aerial surveys
provide scientific evidence of some level of year-round presence of
harbor seals in Iliamna Lake.
Conclusions drawn from recent aerial surveys suggest that some
harbor seals may be year-round residents of Iliamna Lake whereas other
harbor seals may seasonally migrate to and from the lake (Burns et al.,
2011; Withrow et al., 2011; Burns et al., 2012; Burns et al., 2013).
Some of the LTK regarding the migration patterns of seals in Iliamna
Lake are inconsistent, and collectively they do not provide clarity
(see Burns et al., 2013). Some LTK reports indicate harbor seals
migrate between Iliamna Lake and Bristol Bay and are frequently seen
traversing the Kvichak River (e.g., Alvarez 2013; Burns et al., 2013;
Igiugig Tribal Village Council 2013; Mohr 2013; Wilson 2013), while
other reports indicate that the seals do not migrate and are present in
the lake year-round (e.g., Burns et al., 2013; Jacko 2013; Mohr 2013).
Local residents around Iliamna Lake indicate that observations of
harbor seals in the Kvichak River are typically made beginning in
spring, peak during mid-summer, and decline to zero in the winter
months; however, some residents of Levelock on the Kvichak River have
observed seals in the river in the winter (Burns et al., 2013). This
suggests that the Kvichak River may be used seasonally as a migration
route between Iliamna Lake and Bristol Bay.
No scientific data are available to determine whether enough fish
remain in Iliamna Lake to support hundreds of seals during winter. Some
LTK indicates that the lake may not have sufficient food available to
support the number of seals observed in summer months on a year-round
basis. A local seal hunter recently noted that two seals harvested
during two consecutive winters in the lake had not ``one drop of food
in the stomach or intestines'' (Burns et al., 2013). Another seal
hunter recollected shooting a seal in March one year that was very
skinny and had no fat on it, and speculated that during cold winters
there was inadequate food for the seals (Burns et al., 2013). However,
the hunter also mentioned that it was very rare to find a skinny seal
in Iliamna Lake. During our public comment period we received a comment
that provided calculations of the abundance of non-salmonid freshwater
fish available during the overwinter period and indicated that a
population of approximately 300 seals could not be sustained on the
levels of freshwater fish available in the winter. We have no
information to support or refute the calculations provided by the
commenter.
Alternatively, there may be adequate abundance of prey available in
the lake year-round, but some seals could leave the lake in winter for
other reasons. In the St. Lawrence estuary, a study of satellite-tagged
harbor seals found that seals left summer haul-out areas when solid ice
formed within the bays of the estuary despite ``evidence of high
abundance of potential prey for harbor seals in the estuary during
winter'' (Lesage et al., 2004). This study concluded that availability
of prey in winter ``is not the primary factor which influences the
movement and distribution patterns of harbor seals'' (Lesage et al.,
2004). As discussed earlier, harbor seals have been documented to have
spatially separated home ranges which vary seasonally (e.g. Lowry et
al., 2001; Lesage et al., 2004; Peterson et al., 2012; Womble and Gende
2013), but also high site fidelity to breeding locations. Thus, it is
plausible that some harbor seals from Bristol Bay seasonally follow the
salmon to Iliamna Lake and return to Bristol Bay for winter, but there
are no data available either to support or refute this scenario.
Whether seals migrate seasonally between Iliamna Lake and Bristol
Bay has not been scientifically investigated, with the exception of a
few recent aerial surveys of Iliamna Lake and the Kvichak River. Aerial
surveys of the Kvichak River (five complete or partial river surveys
conducted from 2008-2013) have failed to document harbor seal presence
in the river (Burns et al., 2013), but it is possible that seals in the
river may have been missed during the surveys or that the surveys were
conducted when seals were not using the river. For example, during an
aerial survey in 2011, the survey crew received a report of seals in a
tributary of the Kvichak River near Kastinak Flats, but the survey crew
was unable to locate the seals when they flew over the area
approximately 30 minutes later (Burns et al., 2013; D. Withrow, NMML,
pers. comm.). Additionally, Burns et al., (2013) postulated that seals
present in the Kvichak River may not be accounted for as a result of
the survey methodology, which only counts seals hauled-out, not those
in the water. Other reports suggest harbor seals are regularly seen
throughout the Kvichak River (Burns et al., 2013; Van Lanen et al.,
2013; ADF&G unpubl. data). Of 14 harbor seals satellite tagged in
Egegik and Ugashik Bays within eastern Bristol
[[Page 81078]]
Bay in 2000 and 2001, none were documented in the Kvichak River or
Iliamna Lake (ADF&G unpubl. data). However, the sample size is too
small to conclude that migration between Bristol Bay and Iliamna Lake
does not occur. We did not find any scientific evidence to conclude the
harbor seals in Iliamna Lake constitute a closed population with no
migration between the lake and marine waters, and the documented LTK on
this question was inconsistent. In the absence of persuasive evidence
of a closed population, for purposes of our DPS assessment, we assumed
that harbor seal migration between Iliamna Lake and Bristol Bay (or
beyond) is possible.
Subsistence Harvest
Harbor seals are an important resource for Alaska Native
communities surrounding Iliamna Lake. Harbor seals are not only a food
source, but also provide materials that can be used for clothing,
handicrafts, and cultural traditions. Reports of harvesting harbor
seals by indigenous people around Iliamna Lake date back to the early
1800s and LTK suggests that seals have inhabited the lake for many
centuries (Fall et al., 2006; Van Lanen 2012; Burns et al., 2013). The
majority of hunting occurs during February and March; however, some
animals have been harvested in summer and occasionally in winter (Burns
et al., 2013). Seven communities around Iliamna Lake and along the
Kvichak River were surveyed regarding their harvest of marine mammals:
Pedro Bay, Pope-Vannoy Landing, Kokhanok, Newhalen, Igiugig, Iliamna,
and Levelock (Burns et al., 2013). Between 1982 and 2011, approximately
150 seals were harvested in Iliamna Lake; however, there is a marked
difference in the number of seals harvested each of those years (Burns
et al., 2013). For instance, there were no reported harvests of seals
in 1982 and 1996, yet 33 were harvested in 1991. The most recent survey
in 2011 reported that 44 percent of households surveyed from these
seven communities used ``freshwater'' harbor seal products and 13
percent used ``saltwater'' harbor seal products in some capacity,
resulting from an estimated harvest of 29 seals (five ``saltwater'' and
24 ``freshwater'') (Burns et al., 2013).
Distinct Population Segment (DPS) Assessment
As described above, only species, subspecies, and DPSs are eligible
for listing as a threatened or endangered species under the ESA. A DPS
is a population or group of populations of a vertebrate species that
meet both the ``discreteness'' and ``significance'' criteria of our DPS
policy (61 FR 4722; February 7, 1996). If a population segment is found
to be discrete and significant, it is a DPS and is considered a
``species'' under the ESA. If the population is not both discrete and
significant, it does not meet the criteria for designation as a DPS and
does not qualify as a ``species'' as defined by the ESA; thus, we need
not evaluate its status relative to the factors in section 4(a)(1) of
the ESA because it cannot be listed as a threatened or endangered
species. Our assessment first addresses the discreteness of the harbor
seals found in Iliamna Lake, and then addresses whether these seals are
significant to P. v. richardii, as these terms are defined in our DPS
policy (61 FR 4722; February 7, 1996).
As discussed above, we know from formal scientific studies and LTK
that at least some harbor seals are present in the lake year-round;
i.e. are residents of Iliamna Lake. What is not clear from the science
or LTK is whether harbor seals from Bristol Bay migrate to Iliamna
Lake. The BRT considered four scenarios: (1) The population of seals in
Iliamna Lake is self-sustaining with seals being year-round residents
of the lake, and no migration of seals from Bristol Bay into the lake
occurs; (2) there are resident seals in the lake, and some seals from
Bristol Bay migrate to the lake during the summer, but there is no
interbreeding of seals from the two regions and the Bristol Bay seals
do not stay in the lake during winter; (3) Iliamna Lake contains a mix
of seals born in the lake and those born in the marine environment but
who migrated to the lake (either temporarily or permanently), and these
seals are interbreeding; or (4) there is no self-sustaining population
of seals in the lake and migration is necessary to sustain the
population of seals in the lake. The BRT found three of the four
scenarios to be plausible, favoring explanations 1 and 2, but not
ruling out 3. None of the BRT members considered the forth scenario
likely (Boveng et al., 2016). For our DPS analyses, we recognize that
questions remain regarding whether there is migration, and references
below to seals in or from Iliamna Lake are not meant to imply that
their birth location (either in Iliamna Lake or the marine environment)
is known, but rather are an indication of the seals' location in
Iliamna Lake at time of observation or sampling.
Discreteness
We first sought to determine whether the harbor seal population in
Iliamna Lake is discrete in relation to the remainder of the taxon to
which it belongs (i.e., the eastern North Pacific harbor seal
subspecies, P. v. richardii). A population segment of a vertebrate
species may be considered discrete if it satisfies either one of the
following conditions specified in our DPS policy: ``(1) it is markedly
separated from other populations of the same taxon as a consequence of
physical, physiological, ecological, or behavioral factors.
Quantitative measures of genetic or morphological discontinuity may
provide evidence of this separation; or (2) it is delimited by
international governmental boundaries within which differences in
control of exploitation, management of habitat, conservation status, or
regulatory mechanisms exist that are significant in light of section
4(a)(1)(D) of the ESA.'' Because Iliamna Lake is entirely within the
United States, the second discreteness criterion identified above is
not relevant. Thus, we focused our assessment of discreteness on
whether the harbor seals in Iliamna Lake are markedly separated from
other harbor seals in the subspecies P. v. richardii, with emphasis on
the nearest harbor seal stock in adjacent Bristol Bay. In addition to
examining four categories of factors (i.e., physical, physiological,
ecological, or behavioral factors) as mechanisms with the potential for
providing marked separation by limiting the dispersal of breeders
between populations, the BRT recognized that dispersal rates often
cannot be directly measured in natural populations. As such, the BRT
also decided to separately review the available genetic information for
evidence of separation.
Physical Factors: Iliamna Lake is located at the base of the Alaska
Peninsula, where it drains through the Kvichak River into Bristol Bay.
Thus, harbor seal habitat in Iliamna Lake is separated from the nearest
habitat commonly used by harbor seals in Bristol Bay by the Kvichak
River. Reports regarding the length of the Kvichak River vary, with
some older documents reporting the river is approximately 80 km (50 mi)
in length (e.g., Orth 1971; BLM 2004), whereas more recent reports
suggest it is closer to 115-120 km (71-75 mi) (e.g., Withrow and Yano
2009; Boveng et al., 2016; validated by a measurement of the river path
between Kogging and Iliamna Lake using a high resolution topographic
map). The discrepancy in reported distances of the river could be
explained by changes in the river itself over time, variances in the
starting and ending measurement points, or by using
[[Page 81079]]
straight-line measurements on a map versus tracing the path of the
river.
Although seals are found predominantly in the northeast region of
Iliamna Lake, the most recent studies indicate harbor seals are found
throughout Iliamna Lake, in rivers draining into the lake (Iliamna,
Newhalen, and Gilbralter rivers), and throughout the Kvichak River
(Alvarez 2013; Burns et al., 2011; Burns et al., 2012; Burns et al.,
2013; Igiugig Tribal Village Council 2013; Mohr 2013; Van Lanen et al.,
2013; Wilson 2013). The distance that seals would have to travel from
the lake to Bristol Bay is well within the known distances that harbor
seals travel (see previous discussion in ``Habitat Use and
Movements''). Thus, the evidence available does not indicate that the
length of the Kvichak River nor the distance to the northeast region of
Iliamna Lake (approximately 180 km from Bristol Bay) would be a
physical barrier separating seals in Iliamna Lake from those in Bristol
Bay.
Physical factors that could impede harbor seal passage in the
Kvichak River include shallow braided sandbars and ice cover during
winter. Although poorly adapted for travel on land, harbor seals in
other areas have been suspected to cross land up to 0.15 km long and on
inclines as steep as 25 degrees to get from one body of water to
another (COSEWIC 2007), so it is reasonable to assume harbor seals have
the capability to cross shallow braided sandbars in the Kvichak River.
Millions of sockeye salmon enter Iliamna Lake from marine waters
annually via the Kvichak River along with other species of anadromous
salmon. Also, another marine mammal species has been reported to travel
to Iliamna Lake via the Kvichak River. Beluga whales, which are less
agile and much larger than harbor seals, have been documented in the
Kvichak River (Frost et al., 1983; Quakenbush 2002) in the spring,
summer, and fall (Chythlook and Coiley 1994) and have been observed
near Igiugig (Burns et al., 2013; Wilson 2013) and in Iliamna Lake
(Mohr 2013). Thus, the available evidence suggests the Kvichak River is
passable for harbor seals, at least part of the year when the river is
not frozen over.
Individual BRT members were not in agreement regarding the
scientific support for discreteness due to physical factors, but
concluded ``no strong evidence was found either for or against marked
separation by physical barriers between harbor seals in Iliamna Lake
and those in Bristol Bay'' (Boveng et al., 2016). When we considered
the best available information indicating that there is access between
Iliamna Lake and Bristol Bay via the Kvichak River, which is passable
at least part of the year, and that the distance between the two
locations is within documented migration distances of harbor seals,
along with with the opinion of the BRT, we concluded that the best
available information does not support a conclusion that there is
separation due to physical factors. As such, we find that harbor seals
in Iliamna Lake are not markedly separated from other harbor seals of
the subspecies P. v. richardii as a consequence of physical factors.
Physiological Factors: Unlike the Lacs des Loups Marins harbor
seals in Canada, a landlocked population that lives exclusively in
freshwater lakes and rivers and has documented physiological
differences from the adjacent harbor seal population in marine waters
(Smith et al., 1994), no studies exist suggesting there are
statistically significant morphological or physiological differences
between harbor seals in Iliamna Lake and other members of the
subspecies P. v. richardii. Consequently, our discreteness analysis
considered other types of evidence which may suggest physiological
differences. Specifically, we considered observations obtained
primarily from those with LTK of seals in Iliamna Lake having a
different size, taste, pelage, and timing of pupping as compared to
seals in Bristol Bay.
The concentration and availability of salmon to seals in Iliamna
Lake in the summer may account for perceived differences reported by
LTK in size and taste of seals in Iliamna Lake compared to seals in
Bristol Bay. For example, several respondents of a recent LTK survey
indicated that the ``physical size of the seals grows every year
following the salmon runs'' (Burns et al., 2013), suggesting high
availability and consumption of energy-rich salmon results in growth of
seals during the summer. While the well-fed seals may have experienced
salmon-fueled growth, the flavor of the harvested seals has been
reported to become less desirable after the salmon runs, which is
reportedly why seals in Iliamna Lake are not normally hunted in fall
(Burns et al., 2013). The LTK perception of differences in pelage
pattern and coloration is conflicting (see Burns et al., 2013), and no
formal studies have been conducted to determine if there are
significant differences in pelage patterns for harbor seals in Iliamna
Lake versus elsewhere. Burns et al., (2013) speculate that the timing
of the harvest of harbor seals in relation to the timing of the annual
molt may play a role in the perceptions of difference in pelage texture
or coloration. The observed variances in taste, body size, and pelage
traits are more likely a reflection of seasonal diet, normal phenotypic
plasticity, and individual variation rather than an indication that the
seals in Iliamna Lake are physiologically distinct from those in the
adjacent marine environment.
The timing of pupping for eastern North Pacific harbor seals ranges
from March to September (Bigg 1969; Temte et al., 1991; Sease 1992). In
Iliamna Lake, LTK reports about the timing of pupping are variable,
with some reports of seal pups born on the lake ice during March and
April, and other reports indicating pups are born during the first half
of June (Burns et al., 2013). LTK observations of seal pup sightings in
Iliamna Lake ranged from February to September, with the majority of
pup sightings between April and August (Burns et al., 2013). Between
2009 and 2013, aerial surveys of Iliamna Lake documented newborn pups
in June, July, and August (Burns et al., 2013). Both aerial survey
observations and local resident observations of newborn seal pups in
Iliamna Lake are within the normal range of pupping dates for the
eastern North Pacific harbor seal subspecies.
Jemison and Kelly (2001) and Reijnders et al. (2010) showed that
the timing of harbor seal pupping in the same location can shift by as
much as several weeks over the course of a few decades. A review of
data from 1975-2006 for harbor seals in Nanvek Bay, Alaska, (which is
the main location within Bristol Bay for which harbor seal pupping data
are available) indicates that the average peak pupping date can vary by
a couple of weeks over just a few years (e.g., June 18 in 2002 vs. July
3 in 2006; see Table 1 in Boveng et al., 2016). This observed natural
variation in timing of harbor seal pupping, along with scarcity of
available data, may account for seemingly conflicting information in
the scientific literature about the timing of pupping in Iliamna Lake
relative to other harbor seals in Alaska (e.g., Burns et al., 2013
states ``when compared to Bristol Bay seals only, the timing of pupping
in Iliamna does not appear to be substantially delayed'' versus Withrow
et al. (2011) which states ``Elsewhere in Alaska we observe harbor
seals pupping much earlier, in May and June''). According to the BRT
report (Boveng et al., 2016), the latest peak pupping date estimated
for the Nanvek Bay region of Bristol Bay was July 5 (1990). Iliamna
Lake aerial surveys flown in 2010, 2011, and 2013 indicate that the
earliest peak pupping date was July 9 (2010). Sparse data
[[Page 81080]]
about pupping dates in both Bristol Bay and Iliamna Lake lead us to
conclude that while we do not know the precise timing of peak pupping
of harbor seals in either region, we do know that timing of peak
pupping can vary by a couple of weeks among years within a given
location. Therefore, an overlap of the timing of pupping between seals
in Bristol Bay and Iliamna Lake is possible, even though there may be a
15-day delay in the average peak pupping date in Iliamna Lake (July 12)
versus the average peak pupping date in Nanvek Bay (June 27) (see
Boveng et al., 2016). Burns et al. (2013) also concluded that compared
to Bristol Bay, the timing of pupping in Iliamna Lake does not appear
to be substantially delayed. A model developed to estimate the
abundance and trend of harbor seals in Iliamna Lake (Boveng et al., in
prep as cited in Boveng et al., 2016) predicted a peak pupping date of
July 20 (versus the July 12 peak pupping date suggested by a simple
average of the dates of maximum pup counts presented in Table 1 of the
BRT Report); however, there was substantial imprecision in the model's
estimate for the peak of pup counts in the lake.
Individual BRT members were not all in agreement regarding the
degree of scientific support for discreteness based upon marked
separation due to physiological factors. Regarding differences in
physiological traits such as pelage coloration or texture and seal size
and taste, the BRT report stated ``whether any of these differences
truly reflect physiological differences or separation is not clear, and
the BRT was unaware of any documentation that these traits are
heritable and would indicate separation or novel genetic diversity''
(Boveng et al., 2016). Regarding physiological separation based on the
notion that pupping in Iliamna Lake is potentially delayed by two to
six weeks when compared to nearby populations, the BRT stated, ``The
sparsity of information currently available for Iliamna Lake,
imprecision in determining the timing for any of the comparison
populations, and the length of the harbor seal pupping period
(approximately 6-10 weeks), reduce the confidence that can be placed on
the apparent difference'' (Boveng et al., 2016).
When we considered all the evidence currently available to us,
including the lack of direct measures of physiological factors, the
possibility that perceived differences in seals' appearance may be the
result of natural individual variation, the imprecision of estimating
pupping dates due to limited data, the potential overlap of pupping
seasons between Iliamna Lake and Bristol Bay, and the large timeframe
(March to September) for typical pupping times across the eastern North
Pacific harbor seal taxon, we concluded that the available information
is too weak for us to make a determination that there is separation
based on physiological factors. As such, based on the available
evidence, we find that harbor seals in Iliamna Lake are not markedly
separated from other harbor seals of the subspecies P. v. richardii as
a consequence of physiological factors.
Ecological Factors: Harbor seals are known to pursue and aggregate
around concentrations of anadromous prey, particularly salmon (e.g.,
London et al., 2001, Orr et al., 2004, and Wright et al., 2007, as
cited in Peterson et al., 2012; Middlemas et al., 2006; Hauser et al.,
2008). Changes in distribution of seasonally abundant prey in the
Pacific Northwest have been suggested as a possible explanation for
seasonal movements of harbor seals in that area (Peterson et al.,
2012), as harbor seals may move deliberately to exploit regions of
higher prey availability (Hardee 2008). In Alaska, movements of 125 km
by adult female harbor seals have coincided with seasonal eulachon runs
in the Copper River Delta (Lowry et al., 2001). Savarese and Burns
(2010) documented peak harbor seal numbers coincident with peaks in
regional salmon abundance in the Bering Glacier region, and contended
the salmon attracted large numbers of harbor seals to the region.
Peterson et al. (2012) speculated that the observations of harbor seals
using spatially separated haul-out sites on a seasonal basis may be
related to seasonal changes in prey distribution and foraging
opportunities.
Hauser et al. (2008) examined foraging by harbor seals in Iliamna
Lake during July and August, when salmon are very abundant in the lake,
and reported that the seals predominately fed on large salmonids
(salmon, trout, char, and graylings) during the summer months. In
addition to salmonids, Hauser et al. (2008) documented lampreys,
smelts, sculpins, whitefishes, sticklebacks, and other unidentified
prey items in the scat samples of harbor seals in Iliamna Lake. Thus,
harbor seals in Iliamna Lake appear to be opportunistic feeders,
consistent with the general pattern of harbor seals foraging on a wide
variety of fish and invertebrate prey across their range, with regional
differences in diet diversity (Jemison 2001; COSEWIC 2007). The prey
items and seasonal concentration of salmon in the diet of seals in
Iliamna Lake are consistent with those documented for harbor seals in
other freshwater systems. For example, Middlemas et al. (2006)
documented a summer peak in the contribution of salmonid prey to the
diet of harbor seals observed in a Scottish river system; Beck et al.
(1970) documented a seal in Edehon Lake, Canada with both trout and
whitefish in its stomach; and Power and Gregoire (1978) reported that
harbor seals in lakes ate various freshwater fish present in the lakes,
including trout. Smith et al. (1996) examined stomachs of four harbor
seals from the Lacs des Loups Marins which contained in large part lake
whitefish, lake trout, and brook trout. Scat collected in the Nanvak
Bay region of Bristol Bay also showed that harbor seals have a diverse
diet, including some of the same types of prey species consumed in
Iliamna Lake (e.g., salmon, smelts, sculpins) as well as other prey
species (e.g., codfishes, herring, squid/octopus) (Jemison 2001).
Stable isotope analyses of whiskers and muscle tissue can provide
some insights about harbor seal diets from several months prior to the
date the samples were collected. Samples collected from a small number
of subsistence harvested harbor seals from Iliamna Lake provide
preliminary evidence that those specific seals consumed freshwater fish
during the previous winter (Burns et al., 2013). These preliminary data
and the typical timing of ice melt in the Kvichak River and Iliamna
Lake (May-June) suggest that these samples were most likely collected
from seals which had overwintered in the lake. However, these
preliminary stable isotope data are not especially revealing due to the
lack of data on whisker growth rates, tissue turnover times, and direct
measures of the isotopic signature of potential prey resources (Burns
et al., 2013).
If ecological factors prevented harbor seals in Iliamna Lake from
mixing with other harbors seals during mating season, then there could
be marked separation as a result of lack of opportunities for
interbreeding. However, when considering the timing of the annual ice
melt in the Kvichak River and Iliamna Lake, the sockeye salmon runs
into Iliamna Lake, and the presumed mating seasons of seals in Bristol
Bay and in Iliamna Lake, the BRT concluded that the timing of these
events would not preclude opportunities for interbreeding by seals
migrating from Bristol Bay to Iliamna Lake (Boveng et al., 2016).
The BRT members were in general agreement regarding the degree of
scientific support for discreteness based upon marked separation due to
ecological factors, and concluded there
[[Page 81081]]
was ``no strong evidence for separation'' as a result of any of the
ecological factors considered. Based on the available evidence, we find
that harbor seals in Iliamna Lake are not markedly separated from other
harbor seals of the subspecies P. v. richardii as a result of
ecological factors.
Behavioral Factors: There are no scientific or LTK data available
to assess whether mating behaviors (e.g., vocalizations or mate
attraction displays) differ for seals in Iliamna Lake relative to those
in Bristol Bay or other areas of the eastern North Pacific harbor seal
range. Absent data available regarding mating behaviors of harbor seals
in Iliamna Lake, the BRT construed the selection of relatively remote
pupping sites in the northeastern region of Iliamna Lake (nearly 200 km
from pupping sites in Bristol Bay) to be a behavior, and suggested the
selection of the unusual location was evidence of some degree of
separation, especially given harbor seals' site fidelity to breeding
locations. The selection of distant pupping sites could be interpreted
to mean that harbor seals in Iliamna Lake are not freely breeding with
harbor seals in Bristol Bay, and lead to a conclusion there is marked
separation. However, even a small amount of breeding dispersal from
marine populations of harbor seals into Iliamna Lake could render the
degree of genetic differentiation insignificant (Boveng et al., 2016),
suggesting there may not be marked separation. The available LTK does
not resolve this question, as opinions vary regarding whether seals in
the lake are residents, migrants, or a mix of both (see Burns et al.,
2013).
Previously we mentioned that harbor seals commonly follow
anadromous prey into freshwater environments, such as rivers and lakes.
Thus, we do not consider the mere presence of harbor seals in Iliamna
Lake to be a behavioral adaptation suggestive of marked separation from
harbor seals in the marine environment. However, some Alaska Natives in
the Iliamna Lake region, including subsistence hunters, have postulated
that the seals overwinter in the lake by using under-ice air gaps and
haul-outs (Burns et al., 2013), although such winter habitats have not
been documented in Iliamna Lake. Lack of data complicates a
determination of whether use of under-ice shelters would be a special,
learned behavioral adaptation that is unique to harbor seals over-
wintering in freshwater environments, or if this behavior would be one
that any harbor seal in a similar environment may adopt. Similar under-
ice habitats in the Lacs des Loups Marins in Canada have been suggested
as potential harbor seal lairs or breathing chambers (e.g., Smith and
Horonowitsch 1987; COSEWIC 2007). This, in turn, suggests that use of
such under-ice habitats may be an example of the behavioral plasticity
that results in harbor seals using a range of behaviors and habitats in
response to environmental conditions (Komers 1997; Vincent et al.,
2010).
The Lacs des Loups Marins harbor seal population has shown evidence
of modifying typical harbor seal behavior and adapting to its
environment. It is postulated that, because no pups have been observed
being born on the ice during that species' pupping time period (April,
when the lakes are frozen), the Lacs des Loups Marins harbor seals have
learned and adapted to their situation by whelping in under-ice
shelters similar to subnivean birth lairs (snow caves) used by ringed
seals (Consortium Gilles Shooner & Associes et al., 1991 as cited in
Smith 1997). On the contrary, Burns et al. (2013) include information
from local residents near Iliamna Lake who suggest some harbor seal
pups may be born in Iliamna Lake in March and April, when the lake is
still frozen, but pup on the ice, not under it. Due to this reported
on-ice pupping, even if the harbor seals in Iliamna Lake utilize under-
ice habitats as shelters or breathing chambers, such behavior would not
be an adaptation necessary for successful pupping by seals that use the
lake. Thus, unlike the Lacs des Loups Marins harbor seals, the evidence
suggests that harbor seals in Iliamna Lake have not developed novel
behaviors to facilitate pupping in a lake environment.
The BRT members were in general agreement regarding the degree of
scientific support for discreteness based upon marked separation due to
behavioral factors, as determined by selection of pupping locations far
from those in Bristol Bay, and the ambiguity regarding the degree of
migration and breeding dispersal (if any). Their judgment suggests
behavioral separation is possible, but the available evidence is not
strong, or is contradicted by other evidence. Our review of behavioral
factors indicates that the observed harbor seal behaviors in Iliamna
Lake are not uncommon; harbor seals in Iliamna Lake have not been
documented to display behaviors outside the range of normal harbor seal
behaviors (e.g., no unique mating, pupping, or foraging behaviors
reported), although there are unresolved questions about migration and
use of under ice shelters. There is no information available to suggest
that harbor seals living in ice conditions year-round in a freshwater
system would require different behavioral adaptations from harbor seals
living in ice conditions in a saltwater or estuarine system. Despite
the lack of these obvious indications of potential behavioral
separation, we recognize the possibility that the selection of pupping
locations distant from other known pupping locations could be construed
as a behavior and indicate marked separation as a result of the
selection of pupping sites limiting the potential for interbreeding.
Therefore, we find that the best available evidence is not conclusive
but indicates that harbor seals in Iliamna Lake may be markedly
separated from other harbor seals of the subspecies P. v. richardii as
a consequence of behavioral factors.
Genetics: To further consider whether harbor seals in Iliamna Lake
are markedly separated from other populations of eastern North Pacific
harbor seals as a consequence of physical, physiological, ecological,
or behavioral factors, we examined available genetic evidence which may
be indicative of separation. Genetic samples available from harbor
seals in Iliamna Lake were compared to genetic samples available from
harbor seals in the Egegik and Ugashik regions of eastern Bristol Bay.
Bristol Bay has the nearest concentration of seals to Iliamna Lake, and
the BRT determined ``the seals in eastern Bristol Bay would be expected
to be the most similar to the Iliamna Lake seals if there is breeding
dispersal between the two areas, and therefore would be expected to
pose the most stringent test for demonstrating discreteness'' (Boveng
et al., 2016).
Genetic samples have been collected and analyzed from 13 harbor
seals in Iliamna Lake collected in six years from 1996 through 2012.
The mitochondrial DNA (mtDNA) analysis revealed that 11 of 13 seals
sampled from Iliamna Lake exhibited the same mtDNA haplotype (O'Corry-
Crowe 2013), meaning all 11 seals had the same group of genes inherited
from their female parent. The remaining two DNA samples did not yield
results for this test. This specific mtDNA haplotype (Pvit-Hap#7) is
the most common haplotype found in harbor seals sampled from Bristol
Bay and is observed in roughly 21 percent of harbor seals from the
Egegik and Ugashik regions of Bristol Bay (Burns et al., 2013; O'Corry-
Crowe 2013). Thus, this haplotype is not unique to harbor seals in
Iliamna Lake.
The identification of only one mtDNA haplotype in harbor seals from
Iliamna Lake appears to suggest unusually low genetic diversity. For
comparison, 76 harbor seals sampled from the Egegik
[[Page 81082]]
and Ugashik regions of eastern Bristol Bay exhibited 33 different mtDNA
haplotypes (O'Corry-Crowe 2013; Burns et al., 2013). If seals from the
Egegik and Ugashik regions were immigrating into the lake and staying
year-round, there would be almost an 80 percent likelihood that one of
the other mtDNA haplotypes, not Pvit-Hap#7, would be seen in samples
collected from Iliamna Lake (O'Corry-Crowe 2013). However, because
mtDNA is inherited from the mother, mtDNA diversity analysis cannot
determine if male seals are migrating to and from the lake and breeding
with resident female seals. Hardee (2008) recognized similar
limitations of mtDNA given observations of male harbor seals in the
Pacific Northwest traveling larger distances than previously believed,
possibly to mate in a separate geographic region before returning to
their home site. Therefore, conclusive results about the level of
genetic diversity require analyses using nuclear DNA (nDNA; which also
provides information from the male parent), and more formal analyses of
mtDNA with statistical comparisons to harbor seals sampled from other
regions within the range of the taxon (O'Corry-Crowe 2013). These more
stringent data regarding genetic diversity do not exist.
In addition to examining the existing genetic diversity of the
samples, analyses were conducted to examine the extent of genetic
differentiation between harbor seals sampled in Iliamna Lake from those
sampled in the Egegik and Ugashik regions of eastern Bristol Bay. The
results of analyses examining genetic differentiation using both mtDNA
and nDNA suggest that the harbor seals sampled in Iliamna Lake were
genetically differentiated from harbor seals sampled in the Egegik and
Ugashik regions of eastern Bristol Bay (Burns et al., 2013; O'Corry-
Crowe 2013). The results of these analyses also suggest that male and
female-mediated dispersal between the Egegik and Ugashik regions of
eastern Bristol Bay and Iliamna Lake was restricted (Burns et al.,
2013; O'Corry-Crowe 2013). Although no directed comparisons were
conducted between Iliamna Lake samples and genetic samples collected
from harbor seals in other areas of Bristol Bay or other portions of
the range of the taxon, the measure of mtDNA genetic differentiation
between seals in Iliamna Lake and those in eastern Bristol Bay yielded
results showing substantially greater genetic differentiation than all
previous pairwise comparisons between the other major centers of harbor
seal abundance in Alaska (O'Corry-Crowe 2012; Boveng et al., 2016).
These genetic differentiation results are suggestive of the presence of
a small, isolated population of harbor seals in Iliamna Lake.
O'Corry-Crowe (2013) identifies several limitations of the findings
for the Iliamna Lake samples. He cautions that the sample size is
extremely small and that questions regarding the patterns of kinship
among the collected samples remain unresolved (i.e., if some of the
samples were from related individuals, then the data could be skewed
and not representative of a random sampling of the population), and
indicates that genetic differentiation may be enhanced in small
populations when there is a rapid rate of genetic drift, even when
there is continued gene flow. Although the 13 genetic samples from
seals in Iliamna Lake were collected between 1996 and 2012, most
samples were collected during months when seasonal migrants would not
be expected to be in the lake, thus the power to detect seasonal
migrants may be low. Conversely, the timing of the samples may be
benficial for considering if the resident seals in the lake are
discrete from their marine counterparts because for most samples
seasonal migrants would not be expected to be present in the lake.
O'Corry-Crowe (2013) also provides recommendations for future genetic
research to resolve lingering issues, including analyzing 20
microsatellite loci (only 9-11 loci were analyzed) and updating the
techniques used for the analyses to newer technologies, which would
increase the power to resolve genetic questions. We also note that the
tests for genetic differentiation compared the Iliamna Lake samples
solely against samples collected from the Egegik and Ugashik regions of
eastern Bristol Bay. Thus, the samples used for the comparison group
may not be representative of all the seals that could migrate to
Iliamna Lake.
The genetic data available suggest the harbor seals sampled in
Iliamna Lake have low mtDNA diversity, possess the most common mtDNA
haplotype found in Bristol Bay harbor seals, and are genetically
differentiated from harbor seals sampled in the Egegik and Ugashik
regions of eastern Bristol Bay. Given the concerns about the limited
nature of the available genetic information previously discussed here
and by O'Corry-Crowe (2013), ambiguity remains regarding the degree of
separation, and hence discreteness, of harbor seals in Iliamna Lake.
However, in the absence of more samples collected from a greater number
of seals in Iliamna Lake and the Kvichak River, to include the
potential migration season, and/or completion of additional tests such
as those recommended by O'Corry-Crowe (2013), we consider the existing
genetic results to be the best available data upon which to base our
determination. These genetic results support a decision that harbor
seals in Iliamna Lake are markedly separated from harbor seals in
eastern Bristol Bay, and by assumption, from the remainder of the
taxon.
Discreteness Conclusion
We find the available evidence for discreteness based on physical,
physiological, or ecological factors to be unconvincing. The available
evidence based on behavioral factors is not conclusive, but the
selection of pupping locations distant from other known pupping
locations could be construed as a behavior and indicate marked
separation as a result of the selection of pupping sites limiting the
potential for interbreeding. The strongest evidence for discreteness
derives from 13 genetic samples collected from seals in Iliamna Lake.
Analyses of these samples strongly indicate the seals from Iliamna Lake
are genetically differentiated from seals sampled in two locations
within Bristol Bay (Ugashik and Egegik), the nearest concentration of
seals to Iliamna Lake with genetic data available. Genetic comparisons
of samples for the entire taxon do not exist, but this region within
Bristol Bay was expected to provide the most stringent comparison for
discreteness if there is breeding dispersal between the two regions.
The BRT was in strong agreement that the genetic data reflect marked
separation, although the BRT acknowledged that the mechanism of such
separation is unknown and the data are limited. It is possible that the
limited available genetic data may accurately represent the situation
in both Iliamna Lake and all of Bristol Bay, or that additional genetic
analysis from P. v. richardii animals sampled from elsewhere in their
range or from additional seals in Iliamna Lake, could result in a
different conclusion. Nonetheless, the best available genetic
information leads us to conclude that some portion, and perhaps all, of
the harbor seals in Iliamna Lake likely constitute a resident
population that is genetically differentiated from harbor seals in
eastern Bristol Bay, and thus meet the criteria for consideration as a
discrete entity per our DPS policy (61 FR 4722; February 7, 1996).
Significance
Having determined that resident seals from Iliamna Lake are likely
discrete, at
[[Page 81083]]
least from harbor seals in the Egegik and Ugashik regions of nearby
Bristol Bay, we next sought to determine whether they are significant
to the P. v. richardii subspecies.
In carrying out the significance examination per our DPS policy (61
FR 4722; February 7, 1996), we are to consider available scientific
evidence of the population's importance to the taxon to which it
belongs. This consideration may include, but is not limited to, the
following: (1) Persistence of the discrete population segment in an
ecological setting unusual or unique for the taxon; (2) evidence that
loss of the discrete population segment would result in a significant
gap in the range of the taxon; (3) evidence that the discrete
population segment represents the only surviving natural occurrence of
a taxon that may be more abundant elsewhere as an introduced population
outside its historic range; or (4) evidence that the discrete
population segment differs markedly from other populations of the
species in its genetic characteristics.
This determination, however, is highly fact specific and may
consider factors besides those enumerated above. Further, significance
of the discrete population segment is not necessarily determined by
existence of one of these classes of information standing alone.
Information analyzed under these and any other applicable
considerations is evaluated relative to the biological and ecological
importance of the discrete population to the taxon as a whole.
Accordingly, all relevant and available biological and ecological
information is analyzed. As we explained in the DPS policy, ``the
principal significance to be considered in a potential DPS will be the
significance to the taxon to which it belongs'' (61 FR 4722, 4724;
February 7, 1996). Finally,we assessed the biological and ecological
significance of the seals in Iliamna Lake to the P. v. richardii (the
eastern North Pacific harbor seal) taxon in light of Congressional
guidance that the authority to list DPSs be used ``sparingly'' while
conserving the genetic diversity of the species (see Senate Report 151,
96th Congress, 1st Session).
Persistence in an Unusual or Unique Ecological Setting: In
assessing the ``persistence of the discrete population segment in an
ecological setting unusual or unique for the taxon,'' we considered
whether specific characteristics of the Iliamna Lake environment are
unusual or unique; whether persistence in the Iliamna Lake environment
is unusual or unique; and whether there are adaptations as a result of
persistence in an unusual or unique environment which would result in
the discrete population being biologically or ecologically significant
to the taxon P. v. richardii.
The diet of harbor seals in Iliamna Lake is consistent with what we
would expect for the species occupying a freshwater system dominated by
anadromous salmon. Hauser et al. (2008) indicate that harbor seals in
Iliamna Lake consumed large amounts of sockeye salmon when they were
seasonally abundant, and also fed on trout, char, graylings, lampreys,
smelts, sculpins, whitefishes, sticklebacks, and other unidentified
prey items. Burns et al. (2013) examined eight harbor seal stomachs
collected from seals harvested from Iliamna Lake in 2011 and 2012; only
three had identifiable prey items and the remaining five stomach were
either empty, only had worms, or had unidentifiable contents. An
examination of the identifiable prey items found that these seals had
consumed small or young salmonids (salmon and/or trout), threespine
stickleback, and Arctic grayling or lake whitefish (Burns et al.,
2013). The variety and types of prey items in the diet of these sampled
seals in Iliamna Lake reflects harbor seals being opportunistic feeders
(Carretta et al., 2015), and the available data suggest no unusual or
unique prey for the habitat occupied.
We also considered whether the habitat available for use by seals
in Iliamna Lake is unusual or unique. Harbor seals commonly use reefs,
sand and gravel beaches, sand and mud bars, island beaches, and ice
(glacial ice, pan ice, sea ice, or icebergs) as haul-out sites. Harbor
seals in Iliamna Lake are known to haul-out on rocky and sandy
substrates, sand bars, small islands, and ice near pressure cracks or
polynas (Burns et al., 2011; Burns et al., 2012). None of these haul-
out substrates are unique or unusual for harbor seals. Harbor seals in
Iliamna Lake are reported to pup both on ice (Burns et al. 2013) and
other haul-outs in the absence of ice. There is no evidence of seals in
Iliamna Lake pupping in air pockets beneath the ice, which would be
unusual. Such use has been hypothesized for the harbor seals in the
Lacs des Loups Marins (Consortium Gilles Shooner & Associes et al. 1991
as cited in Smith 1997; DFO 2016). According to LTK, pupping in Iliamna
Lake likely occurs at island beaches or sandbars in the northeastern
portion of the lake, which is consistent with the types of substrates
upon which aerial surveys documented pups (i.e., on low-lying islands
and sand spits; Burns et al., 2013). Nothing suggests that harbor seals
in Iliamna Lake display unusual or unique pupping behaviors (including
habitat usage).
Smith and Horonowitsch (1987) studied the ice at one location
within the Lacs des Loups Marins and documented what they refer to as
``shoreline ice-steps'' which they speculated could be used as
breathing chambers for over-wintering seals in the lake. LTK suggests
the presence and use of similar under-ice haul-outs in Iliamna Lake
(Burns et al., 2013). While this would represent unusual habitat use
for harbor seals in general, and unique habitat for harbor seals of P.
v. richardii, it would be consistent with the general observation that
harbor seals exhibit wide variation in habitat use, rather than being
indicative of an adaptation by seals in Iliamna Lake that would be
significant to the P. v. richardii taxon as a whole (see further
discussion of habitat adaptation below).
Harbor seals have the broadest distribution and occur in more
different habitats than any other pinniped species (Burns 2002; COSEWIC
2007), and are frequently and commonly observed in freshwater systems
(Burns 2002). Mansfield (1967) provides information about sightings of
harbor seals in rivers and lakes in Arctic Canada (referencing Doutt
1942 and Harper 1961 for detailed summaries of Arctic harbor seals'
freshwater distribution), indicating that harbor seals have ``a strong
liking for fresh water'' and are often found in estuaries and
freshwater habitats ``far from the sea.'' Beck et al., (1970) report
harbor seals in the Thlewiaza River system and associated lakes west of
Hudson Bay. Smith et al. (1994) and Smith (1997) provide an extensive
list of reports of harbor seals documented in freshwater systems. Smith
et al. (1996) conducted analyses involving both the Lacs des Loups
Marins harbor seals as well as a second group of ``lacustrine'' harbor
seals from Kasegalik Lake in Canada's Northwest Territory. Middlemas et
al. (2006) provide documentation of harbor seals in a Scottish river
system. The Committee on the Status of Endangered Wildlife in Canada
(COSEWIC) reports that harbor seals occasionally ascend the St.
Lawrence River to the Great Lakes (COSEWIC 2007). In the Bristol Bay
region, harbor seals have been observed in other lakes in addition to
Iliamna Lake, such as Lake Becharof and Naknek Lake (Mathisen and Kline
1992). Thus, the presence of harbor seals in freshwater systems or
lakes, including Iliamna Lake, is not unusual or unique for the
species.
Year-round persistence of harbor seals in a lake is less common.
Besides the unknown number of harbor seals
[[Page 81084]]
occupying Iliamna Lake through the winter, the Lacs des Loups Marins
harbor seals are the only other documented instance of harbor seals
persisting in freshwater systems year-round. However, a review of
available literature suggests the possibility this scenario may be more
prevalent than just these two groups of harbor seals. For example,
Mansfield (1967) states that the population of freshwater harbor seals
in the Upper and Lower Seal Lakes east of Hudson Bay (a.k.a. the Lacs
des Loups Marins) is not unique given reports of harbor seals found in
other freshwater systems of Canada. Beck et al. (1970) postulated that
harbor seals may live in the Thlewiaza River and associated lakes year-
round, and documented a pup in the Edehon Lake, leading them to
conclude that harbor seal reproduction is successful in that freshwater
habitat. Beck et al. (1970) also concluded that individual seals in
those lakes may be born and spend most or all of their lives in
freshwater, but there was no reason to believe they were an isolated
population. In Alaska, winter aerial surveys led Savarese and Burns
(2010) to suggest that harbor seals are present year-round in Vitus
Lake, a tidally-influenced lake near the Bering Glacier. No pups were
documented during that study and diet and genetic data indicated seals
from various stocks moved into Vitus Lake to take advantage of local
salmon runs (Savarese and Burns 2010). These reports of potential year-
round presence of harbor seals in various freshwater systems are
sporadic, and do not confirm self-sustaining populations exist in those
other freshwater systems. Therefore, it is reasonable to conclude that
the year-round persistence of a discrete population of harbor seals in
the freshwater environment of Iliamna Lake is at least unusual, if not
unique, to the P. v. richardii harbor seal taxon.
The BRT considered whether the persistence of the population of
harbor seals in this setting is important to the taxon as a whole (see
discussion in Boveng et al., 2016). Specifically, the BRT considered
whether harbor seals in Iliamna Lake exhibit any adaptations to the
environment which would be biologically or ecologically significant to
the P. v. richardii harbor seal taxon. The evidence of such adaptations
is not necessarily required to demonstrate significance; however, the
BRT examined such evidence here in light of harbor seals' widespread
and diverse habitat and diet. The BRT considered the physiology of the
seals in Iliamna Lake as well as their over-wintering strategy as
possible indicators of adaptations of potential importance for the
taxon.
As previously discussed, some local residents of the Iliamna Lake
region have suggested they think the harbor seals harvested from
Iliamna Lake taste, look, or feel different (e.g., seals are fatter;
pelage is softer) from those harvested in the marine environment (Burns
et al., 2013). There was, however, a lack of consensus regarding the
perceived differences (e.g., some say seals from Iliamna Lake are
darker than marine counterparts, others say the seals are lighter)
among the local residents interviewed. Moreover, attributes such as
fatness and softness of the coat, or the way the seals taste when
consumed, are not necessarily inherited traits and could be acquired
during time spent in the lake. Unlike other lake seal species, there
are no data available to document whether morphological (e.g.,
craniometric) differences exist; if such morphological differences are
present, they are not distinct enough to be generally recognized in
traditional knowledge of Alaska Native residents in the area (see
discussion in Boveng et al., 2016). There is no evidence to suggest
these reported physical differences in fatness, softness, or taste are
adaptations that would convey significance of these seals to the taxon.
The use of air gaps under the ice in winter is a potential
adaptation to freshwater life in sub-Arctic regions, and is only
documented among harbor seals in one location (P. v. mellonae of Lacs
des Loups Marins). Whether the use of under-ice shelters would be a
true adaptation to a freshwater environment which freezes over, or
would simply be a response to habitat conditions that may be used by
any harbor seal exposed to those conditions, remains uncertain. On the
importance of this particular behavior relative to significance of
seals in Iliamna Lake to the P. v. richardii subspecies, the BRT
concluded any assessment would ``be in the realm of judgment or even
speculation'' (Boveng et al.,2016). Even though harbor seals in Iliamna
Lake cope with the extensive ice cover in winter, there is no
indication they have adapted or modified their breeding, whelping, or
pup-rearing behaviors in a manner unusual for, or of significance to,
the taxon.
The BRT members were in strong agreement that harbor seals
persisting year-round and breeding in a freshwater lake that freezes
over almost completely nearly every year is unique for the subspecies
P. v. richardii, and unusual for the harbor seal species. However,
there was a lack of consensus amongst BRT members whether the available
evidence reflects physical, life-history, or other adaptations as a
result of persisting in an unusual or unique ecological setting which
would make the harbor seal population in Iliamna Lake biologically or
ecologically significant to the broader taxon. The discrepancies in
opinion stemmed from ``differences in assessing the weights of several
lines of qualitative and indirect evidence'' (Boveng et al., 2016). The
BRT also concluded (1) seals from the marine population would be able
to persist in the Iliamna Lake setting, and (2) even if seals from the
marine population were unable to persist in Iliamna Lake, the ``lack of
`ecological exchangeability' is not important to the persistence of the
taxon as a whole'' (Boveng et al., 2016). Ultimately, the BRT's
assessment favored ``a conclusion that the evidence does not support
significance'' (Boveng et al., 2016). We agree that persistence of a
population of harbor seals in the unusual or unique ecological setting
of Iliamna Lake in and of itself does not confer significance of that
population to the taxon. The absence of evidence suggesting the harbor
seals in Iliamna Lake have adaptations to their environment which would
benefit the taxon to which they belong leads us to determine that the
persistence of a population of harbor seals in Iliamna Lake is not
significant to the subspecies P. v. richardii.
Evidence That Loss Would Result in Significant Gap in Range:
Eastern North Pacific harbor seals range from Mexico northward along
the coastlines of the continental U.S. and Canada and much of Alaska.
In Alaska, harbor seals of this subspecies are distributed almost
continuously throughout the southern coastal waters in the region
surrounding Iliamna Lake. In assessing whether the loss of harbor seals
in Iliamna Lake would result in a significant gap in the range, we
considered a scenario whereby all the seals in the lake were extirpated
and there was no migration into the lake, either because there is no
migration currently occurring or because a future physical barrier
prevents migration. Given the extensive and continuous range of the
eastern North Pacific harbor seals, the loss of the small proportion of
habitat in Iliamna Lake would not result in a significant gap in the
range. Furthermore, the evidence indicating possible seasonal movement
of some harbor seals from Bristol Bay to Iliamna Lake suggests that the
habitat in this portion of the range could be reoccupied.
The loss of harbor seals in Iliamna Lake would not have a
detrimental impact to other harbor seal populations
[[Page 81085]]
that comprise the subspecies P. v. richardii, as this is not an
interstitial population of harbor seals whose loss would isolate
another population from the main group. Additionally, there are only an
estimated 400 harbor seals in Iliamna Lake (Boveng et al., 2016), so
this population represents a minute fraction of the total population
ofeastern North Pacific harbor seals, estimated at 360,000 (DFO 2010).
The BRT was in strong agreement that the evidence is clear that the
loss of the Iliamna Lake segment would not result in a significant gap
in the range of the taxon, and we agree.
Evidence of Only Surviving Natural Occurrence: Harbor seals in
taxon P. v. richardii are currently found throughout their historic
range along the coasts from Baja California, Mexico, northward to
Alaska, and west through the Gulf of Alaska and Aleutian Islands, and
in the Bering Sea north to Cape Newenham and the Pribilof Islands.
There are no known introductions of this species to any place outside
its historic range, thus it is naturally occurring wherever it occurs.
The BRT was unanimous in its assessment that harbor seals in Iliamna
Lake are not the only surviving natural occurrence of the taxon. We
concur in that determination.
Evidence of Marked Difference in Genetic Characteristics: As
discussed above, the limited genetic data available from seals in
Iliamna Lake indicate 11 of 13 (2 samples did not yield results)
sampled seals had the same mtDNA haplotype, an indication of possible
low genetic diversity (O'Corry-Crowe 2013). Unlike the Lacs des Loups
Marins harbor seals, which exhibit mtDNA haplotypes that are only found
in seals from the Lacs des Loups Marins (Smith 1997), the single mtDNA
haplotype exhibited in the harbor seals in Iliamna Lake is not unique
to Iliamna Lake. Rather, it is the most common mtDNA haplotype found in
samples from harbor seals in Bristol Bay (O'Corry-Crowe 2013; Van Lanen
et al., 2013). One plausible explanation for the single haplotype found
in all the harbor seal samples from Iliamna Lake is that these seals
are simply a genetic subset of seals from Bristol Bay, and have lost
rather than gained substantial amounts of genetic diversity since
isolation. An alternative explanation is the seals in Iliamna Lake have
been isolated a long time, during which they may have accumulated
genetic differences at other loci (not currently examined) via
mutation, especially for loci under selective pressure (i.e.,
adaptation). However, as previously discussed, only a small number of
genetic loci were tested and the sample size was small, so the reason
for a single mtDNA haplotype is undeterminable at this time. We
conclude that the best scientific and commercial data available, a
single mtDNA haplotype which is commonly found in other populations of
the taxon and the data used to assess discreteness of the population,
do not indicate that harbor seals in Iliamna Lake have novel genes
which could be significant to the taxon as a whole.
There is no strong evidence to indicate the existence of phenotypic
differences between harbor seals in Iliamna Lake and those in other
portions of the taxon's range. Although there have been some LTK
reports that the seals in Iliamna Lake may taste different or have
pelage of varying appearance from seals in Bristol Bay, there have been
no studies assessing whether these perceived differences are the result
of significant differences in genetics. The BRT members did not reach
consensus regarding this issue, with a slight preponderance of opinion
favoring the conclusion that the genetic characteristics of seals in
Iliamna Lake did not convey significance to these seals in regards to
P. v. richardii. Some members considered the data available as mostly
insufficient for drawing a conclusion regarding significance, and some
considered the evidence against significance slightly more persuasive
than the evidence for significance. Accordingly, we find that the
genetic characteristics (i.e., mtDNA haplotype) found in seals from
Iliamna Lake do not differ markedly from those found in Bristol Bay and
therefore determine that the best available genetic data, albeit
limited, supports a conclusion that harbor seals in Iliamna Lake do not
have genetic characteristics that are significant to the taxon as a
whole.
Overall Significance to the Taxon: We considered several factors
that could indicate whether harbor seals in Iliamna Lake may be
biologically and ecologically significant to the taxon as a whole. Of
the four factors delinated in the 1996 DPS policy, we conclude that
there is evidence of only one: The population persists in an unusual or
unique setting for the taxon. As we explained in our policy,
``occurrence in an unusual ecological setting is potentially an
indication that a population segment represents a significant resource
of the kind sought to be conserved by the'' ESA and in ``any actual
case of a DPS recognized in part on this basis, the Services will
describe in detail the nature of this significance when accepting a
petition or proposing a rule'' (61 FR at 4724). While year-round
persistence in the freshwater environment of Iliamna Lake is unique to
the taxon P. v. richardii and unusual for the entire species, we agree
with the BRT (Boveng et al., 2016) that the best scientific and
commercial data available are limited and suggest that the persistence
of the seals in Iliamna Lake is not significant to the taxon as a
whole. The loss of the Iliamna Lake segment would not result in a gap
in the range of the taxon, and the harbor seals in Iliamna Lake are not
the only surviving natural occurrence of the taxon; thus harbor seals
in Iliamna Lake do not demonstrate significance to the taxon based on
these factors. Further, available genetic data suggest that harbor
seals in Iliamna Lake are not significant to the larger taxon. Although
the best available genetic data indicate that at least some of the
seals in Iliamna Lake are distinct from harbor seals in the eastern
regions of nearby Bristol Bay, the genetic characteristics (e.g., the
single mtDNA observed in samples from seals in Iliamna Lake is the most
common haplotype found in seals frim Bristol Bay) do not appear to
differ in ways that would convey significance to the P. v. richardii
subspecies.
Individual BRT members were not in agreement regarding the degree
of scientific support overall for or against the significance of seals
in Iliamna Lake to the P. v. richardii subspecies, but stated ``the
slight majority judgment against significance of the population segment
. . . summarized a diversity of views about how much weight to place on
the various lines of mostly weak and qualitative evidence'' and that
``the evidence itself must be characterized as mostly indirect,
qualitative rather than quantitative, and equivocal for the purpose of
demonstrating biological or ecological importance to the broader
taxon'' (Boveng et al., 2016). Taking into consideration the totality
of all the information discussed above regarding the possible
significance of harbor seals in Iliamna Lake to the P. v. richardii
taxon, including the qualitative and equivocal nature of the available
information, along with the guidance from legislative history to
identify DPSs ``sparingly,'' we find that the available evidence
supports a conclusion that the harbor seals in Iliamna Lake are not
significant to the remainder of the taxon.
DPS Conclusion
Based on the best scientific and commercial data available, we find
the evidence for marked separation of harbor seals in Iliamna Lake from
the remainder of the taxon based on physical, physiological, ecological
or behavioral factors to be unconvincing or
[[Page 81086]]
weak. The strongest support for marked separation comes from the best
available genetic data which, although limited and preliminary, support
a conclusion that at least some of the harbor seals in Iliamna Lake are
likely isolated from harbor seals in the Egegik and Ugashik regions of
eastern Bristol Bay. Thus, we conclude that the harbor seal population
in Iliamna Lake is separated from other populations of the taxon and
meet the discreteness criterion of our DPS policy (61 FR 4722; February
7, 1996).
Per the second component of our DPS Policy, we are to consider
available scientific evidence of the discrete population's importance
to the taxon to which it belongs (61 FR 4722; February 7, 1996). Our
review of the best available information suggests the only
characteristic which may make this population of harbor seals unique
within its taxon is the fact that they persist year-round in a
freshwater system which freezes over to some degree in most winters.
While that characteristic is unique within the subspecies P. v.
richardii, we determined such persistence is not biologically or
ecologically important to the taxon as a whole. Furthermore, the
information available supports a conclusion that loss of this
population would not be detrimental to the persistence of the taxon or
constitute a gap in the range of the taxon; this population is not the
only natural surviving population; and there are no unique genetic
characteristics conveying significance of this population to the taxon.
After reviewing the best available data as they apply to the
significance criterion, we conclude that the harbor seals in Iliamna
Lake are not significant to the taxon P. v. richardii.
Under our DPS Policy, both the discreteness and significance
elements must be met to qualify as a DPS. Our review has determined
that the seals persisting year-round in Iliamna Lake are discrete but
not significant; therefore, the harbor seals in Iliamna Lake do not
qualify as a DPS and are not a listable entity under the ESA.
Finding
In assessing whether the actions in the petition are warranted, we
reviewed the best available scientific and commercial information
available, including the BRT report, the petition and literature cited
in the petition, published and grey literature relevant to the topic,
correspondence with experts in academic and government institutions,
documentation of LTK, and public comments. On the basis of this review,
we have determined that harbor seals in Iliamna Lake meet the criteria
for discreteness but do not meet the criteria for significance. As
such, the harbor seals in Iliamna Lake do not meet all the criteria
necessary to constitute a DPS, and thus are not a listable entity under
the ESA. Therefore, we find that the petitioned actions to list the
harbor seals in Iliamna Lake as a threatened or endangered species
under the ESA, and to designate critical habitat, are not warranted.
In our 90-day finding (78 FR 29098; May 17, 2013), we indicated we
were commencing a status review of the harbor seals in Iliamna Lake. To
assist our evaluation of whether the seals in Iliamna Lake constitute a
DPS, the BRT prepared a report which compiled background information
about the harbor seals in Iliamna Lake and evaluated the scientific
information relevant to the DPS criteria (Boveng et al., 2016). Upon
our determination that the DPS criteria were not met and the seals in
Iliamna Lake are not a ``species'' under the ESA, there is no need to
complete the status review by conducting a threats assessment or
extinction risk assessment in light of the factors in section 4(a)(1)
of the ESA.
In some instances, where we find a petitioned action is not
warranted because the petitioned population does not constitute a
``species'' under the ESA, we have initiated a status review of a
related or larger population (e.g., the 12-month determination that the
petitioned action to list Lynn Canal Pacific herring was not warranted,
followed by a status review of the Southeast Alaska population of
Pacific herring; 73 FR 19824; April 11, 2008). Here, the scope of the
petition was limited to the seals in Iliamna Lake, and since the most
recent abundance data for the Bristol Bay harbor seal stock (the stock
that includes seals in Iliamna Lake) indicates this stock increased
from an estimated 18,577 seals in 2005 to an estimated 32,350 seals in
2011 (Allen and Angliss 2014; Muto and Angliss 2015), we are not
initiating a status review of the Bristol Bay harbor seal stock at this
time.
References
A complete list of all references cited herein is available upon
request (see ADDRESSES).
Authority
The authority for this action is the Endangered Species act of
1973, as amended (16 U.S.C. 1531 et seq.).
Dated: November 10, 2016.
Samuel D. Rauch, III,
Deputy Assistant Administrator for Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2016-27690 Filed 11-16-16; 8:45 am]
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