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Structure and Assessment of Beluga Whale, Delphinapterus leucas, Populations in the Russian Far East


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In 2000, the International Whaling Commission conducted a global assessment of beluga whales, Delphinapterus leucas. Following this assessment, five beluga stocks were recognized in Russian Far East waters: Western Chukchi-East Siberian Sea, Anadyr Gulf, Shelikhov, Sakhalin-Amur, and Shantar. This paper provides a revised assessment of beluga abundance, distribution, and population structure in the Russian Far East. This region encompasses the Okhotsk Sea, and the coastline of the Chukotka Autonomous Region (CAR), which includes the western Bering, western Chukchi, and eastern East Siberian seas. Published results of genetic analysis are updated with our original unpublished data. Based on information available to date, we propose recognizing seven beluga stocks in the Russian Far East. Five stocks in the Okhotsk Sea: 1) Sakhalin-Amur, 2) Ulbansky, 3) Tugursky, 4) Udska- ya, 5) Shelikhov, and two stocks in the CAR: 6) Anadyr, and 7) Bering-Chukchi-Beaufort (BCB). Natural and anthropogenic threats to these stocks are described and include ice entrapment, over-fishing of key prey, live captures for aquaria, bycatch in fisheries, exposure to effluent, and seismic and military activities
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72 Marine Fisheries Review
Structure and Assessment of Beluga Whale, Delphinapterus leucas,
Populations in the Russian Far East
Olga Shpak (, Ilya Me-
schersky, Dmitry Glazov, Daria Kuznetsova,
and Viatcheslav Rozhnov are with A. N. Sev-
ertsov Institute of Ecology and Evolution of the
Russian Academy of Sciences, Str. Leninsky
Prospect, 33, Moscow, Russia 119071. Denis
Litovka is currently with the Office of Governor
and Government of the Chukotka Autonomous
Region, Str. Bering 20, Anadyr, Russia 689000.
ABSTRACT—In 2000, the International
Whaling Commission conducted a global
assessment of beluga whales, Delphin-
apterus leucas. Following this assessment,
five beluga stocks were recognized in Rus-
sian Far East waters: Western Chukchi-
East Siberian Sea, Anadyr Gulf, Shelikhov,
Sakhalin-Amur, and Shantar. This paper
provides a revised assessment of beluga
abundance, distribution, and population
structure in the Russian Far East. This re-
gion encompasses the Okhotsk Sea, and
the coastline of the Chukotka Autonomous
Region (CAR), which includes the western
Bering, western Chukchi, and eastern East
Siberian seas. Published results of genetic
analysis are updated with our original un-
published data. Based on information avail-
able to date, we propose recognizing seven
beluga stocks in the Russian Far East. Five
stocks in the Okhotsk Sea: 1) Sakhalin-
Amur, 2) Ulbansky, 3) Tugursky, 4) Udska-
ya, 5) Shelikhov, and two stocks in the CAR:
6) Anadyr, and 7) Bering-Chukchi-Beaufort
(BCB). Natural and anthropogenic threats
to these stocks are described and include
ice entrapment, over-fishing of key prey, live
captures for aquaria, bycatch in fisheries,
exposure to effluent, and seismic and mili-
tary activities.
In the Russian Far East, the belu-
ga, Delphinapterus leucas, or white
whale, occurs in the Okhotsk Sea and
along the coastline of the Chukotka
Autonomous Region in the western
Bering, western Chukchi, and eastern
East Siberian seas (Fig. 1). From late
1920’s to the dissolution of the So-
viet Union in 1991, Soviet scientists
conducted extensive studies on this
species’ abundance and distribution,
mainly because belugas were hunt-
ed commercially (Kleinenberg et al.,
1964; Matishov and Ognetov, 2006).
In 2000, scientific information on be-
lugas in Russian waters was reviewed
and summarized for the global assess-
ment of the species by the Scientific
Committee of the International Whal-
ing Commission (IWC, 2000). At that
time, five beluga stocks in Russian
Far-Eastern waters were recognized:
Western Chukchi-East Siberian Sea
(stock no. 25), Anadyr Gulf (26); She-
likhov (27); Sakhalin-Amur (28), and
Shantar (29). The delineation of Stock
25 was questioned by some experts,
who, based on satellite tracking data,
suggested belugas migrating along
Chukotka Peninsula coast belonged
to United States or Canadian stocks
(i.e., Eastern Bering Sea (stock no. 3),
Eastern Chukchi Sea (4), or Beaufort
Sea (or Eastern Beaufort) (5)). For
stocks 25 and 26, no abundance esti-
mates were provided, and the Okhotsk
Sea beluga population (comprised of
stocks 27, 28, and 29) was estimated
at 18,000–20,000 (IWC, 2000).
Apart from nonexistent or method-
ologically poor abundance estimates,
no genetic information was available
at that time, and migration routes and
winter grounds remained mostly un-
known. Stock delineation was based
solely on summer distribution and lo-
cal movement patterns.
Since the IWC (2000) assessment,
and a break in research in the 1990’s
following the dissolution of the So-
viet Union, extensive beluga studies
have been resumed in the Russian Far
East in 2005. Currently management
of marine resources in Russia, includ-
ing belugas, is not based on the spe-
cies population structure, but rather on
geographically defined fishing zones.
Our studies in the Okhotsk Sea have
led to the Russian Federal Agency of
Fisheries and the Ministry of Agricul-
ture recognizing the necessity to con-
sider stock distribution when issuing
total allowable takes (TAT) in this re-
gion (Ministry of Agriculture, Order
N 533, 27 October 2017). An updated
assessment of beluga population struc-
ture would be important for effective
and sustainable management.
Here, we review and summarize our
previously published research results
and present updated genetic results
on the Far Eastern beluga population
structure and status, to provide a basis
for effective conservation and sustain-
able management. A working version
of this paper was originally presented
and discussed at the Global Review of
Monodontids (GROM)1 meeting held
in March 2017 (Hobbs et al., 2019,
provide an overview of the meeting
Our use of the term “biological
population,” or population, is defined
as a group of individuals who inter-
breed and are genetically isolated
from other biological populations due
to geographical, behavioral, or any
other reasons. In summer months,
many beluga populations seasonally
break into discrete, relatively resident
“nursery aggregations” (a synonym to
the term “reproductive aggregation”
al-review-monodontids-now-available/ accessed
9 Aug. 2018).
81(3–4) 73
Figure 1.—Russian Far East with toponyms mentioned in the text.
sometimes used in the Russian litera-
ture), and small groups, comprised
mostly of males, which are more no-
madic. A “stock” (also see Hobbs
et al., 2019, under “Identification of
Stocks”), is a population unit which
can be defined as discrete and sea-
sonally stable and should be managed
separately from other equivalent units
(synonym of “management unit”).
Thus, a summer nursery aggregation is
considered a stock if its geographical
(spatiotemporal) separation from the
neighboring aggregations of the same
biological population can be proved
by genetics, photo-identification, or
other methods. Following the guidance
in IWC (2000:244), “possible stock
units should be split until evidence is
available to justify combining them.”
A “pool” is defined as a large beluga
whale unit of an unknown demograph-
ic status (presumably made up of mul-
tiple stocks or even multiple biological
populations), within which we cannot
delineate stocks or populations due to
the lack of information on their distri-
bution ranges, movement patterns, and
For the genetic analyses presented
herein, we have combined our pub-
lished (Meschersky et al., 2013, 2018)
and unpublished data to delineate each
stock, population, or pool. The level of
population isolation was evaluated us-
ing allele frequencies of 17 microsat-
ellite loci: (Cb1, Cb2, Cb4, Cb5, Cb8,
Cb10, Cb11, Cb13, Cb14, Cb16, Cb17
– Buchanan et al., 1996; Ev37Mn,
Ev94Mn – Valsecchi and Amos, 1996;
415/416, 417/418, 464/465, 468/469
– Schlötterer et al., 1991; Fullard et
al., 2000). The methods of genotyp-
ing and sex determination were anal-
ogous to those previously described
in Meschersky et al. (2013). Sample
comparison was performed using the
allele frequency based Fst-criterion
(Arlequin 3.1 software, Excoffier et
al., 2005) and a clustering method
(LOCPRIOR-Admixture model, 5
runs of 50,000 MCMC iterations of
burn-in and 200,000 iterations of main
analysis) using Structure 2.3.4 soft-
74 Marine Fisheries Review
ware (Pritchard et al., 2000). Patterns
of spatial distribution within regions
and populations were evaluated using
maternal lineage (mtDNA haplotypes)
distribution (Fst-criterion—haplotypes
frequency only, Arlequin 3.1 soft-
ware). In most cases, a fragment of
559 bp (see Genbank JQ716342, for
instance) was used. Where appropri-
ate, results from O’Corry-Crowe et al.
(1997) were included to expand the
dataset. The original samples (skin bi-
opsies or tissues of dead animals) were
collected during the period 2008–16
(and for Sakhalinsky Bay since 2004).
Data were collected during aerial
and vessel surveys and shore-based
operations. Satellite-linked transmit-
ter tags attached to belugas provided
additional information on movement
patterns and seasonal habitat use. We
summarized information from pub-
lished accounts, technical reports, and
unpublished data.
Natural and Anthropogenic
The Federal Agency of Fisheries
provided some numbers for beluga
live-captures and traditional hunting.
However, due to the obvious incom-
pleteness of reported figures, we also
collected information from the re-
gional Fishery offices, and obtained
records directly from the companies
that captured belugas. Further, we list-
ed major current and potential threats
to belugas in different geographic re-
gions based on our expertise.
Okhotsk Sea
Two multi-year projects were con-
ducted in the Okhotsk Sea during the
period 2007–16. Genetic analyses
(Meschersky et al., 2013), together
with geographic distribution studies
(Shpak et al., 2010; Solovyev et al.,
2015), delineated two biological pop-
ulations: 1) Western Okhotsk, which
includes belugas summering in the
Sakhalin-Amur and Shantar regions,
and 2) Northeast Okhotsk (or Shelik-
hov), which spends the summer in the
Gizhiginskaya and Penzhinskaya bays
of Shelikhov Gulf and along the west-
ern coast of the Kamchatka Peninsula
(Fig. 1).
Genetic analysis, based on a signifi-
cantly increased sample (355 samples
from the western Okhotsk Sea and 79
from the northeast Okhotsk Sea) com-
pared to the earlier study (Meschersky
et al., 2013), confirmed the genetic iso-
lation between these two populations.
The Fst distance based on 17 micro-
satellite loci was estimated at 3.52%.
The high level of isolation between the
two populations was also confirmed
by the clustering method analysis (see
samples 1–7, Fig. 2). Maximal average
LnP(K) and Delta K (Evanno meth-
od, Earl and vonHoldt, 2012) values
were found for K = 2 hypothesis for
the total Okhotsk Sea sample set. No
evidence of further subdivision in the
Okhotsk Sea was found for higher K
(up to 7) hypotheses. Significant dif-
ferences (Fst = 32.04%, p = 0.0000)
in mitochondrial lineage compositions
and frequencies were found between
the two Okhotsk Sea regions.
Isolation of the Okhotsk Sea belu-
gas from whales inhabiting the Anadyr
Estuary (in the Bering Sea) was dem-
onstrated earlier (Meschersky et al.,
2013; Borisova et al.2). With the cur-
rent sample size, we unambiguously
confirmed isolation of the Okhotsk
Sea belugas from Bering Sea belu-
gas. The combined Anadyr-Chukotka
coast sample (n = 83) microsatellite
allele distance was estimated at 4.78%
for the western Okhotsk population,
4.30% for the Northeast Okhotsk
(Shelikhov) population, or 4.15% for
the combined Okhotsk Sea sample
(n = 434). All pair-wise compari-
sons were statistically significant (p
= 0.0000). The isolation between the
Okhotsk Sea and Anadyr-Chukotka
belugas was also confirmed by the
clustering analysis (Fig. 2).
Western Okhotsk Population
Based on aerial surveys and coastal
observations (Solovyev et al., 2015)
in the western part of the Okhotsk
Sea, belugas concentrate in the shal-
low waters of the Sakhalin-Amur and
Shantar regions in several nursery ag-
gregations during the summer (Fig.
3). Nursery aggregations tend to stay
2Borisova, E. A., I. G. Meschersky, O. V. Shpak,
D. M. Glazov, D. I. Litovka, and V. V. Rozhnov.
2012. Evaluation of effect of geographical iso-
lation on level of genetic distinctness in beluga
whale (Delphinapterus leucas) populations in
Russian Far East. Marine Mammals of the Hol-
arctic: Collection of papers presented at the 7th
Int. Conf., Suzdal, 2012, 1:113–117 (http://mar-
Figure 2.—The results of K = 3 hypothesis tested by clustering analysis based on 17 microsatellite loci alleles frequencies.
LOCPRIOR-Admixture model. Samples = 1–Sakhalinsky Bay, 2–Nikolaya Bay, 3–Ulbansky Bay, 4–Tugursky Bay, 5–Udskaya
Bay, 6–Western Kamchatka coast, 7–Gizhiginskaya Bay, 8–Anadyr Estuary, and 9–Chukotka coast. Light gray shaded area =
western-Okhotsk population, dark gray shaded area = northeast Okhotsk population, and black shaded area = Anadyr-Chukotka
81(3–4) 75
Figure 3.—Okhotsk Sea beluga summer ranges based on aerial surveys, boat and coastal observations during the period 2007–16
(modified and updated from Solovyev et al., 2015), and winter ranges based on satellite tracking data (Shpak et al., 2010; Shpak
et al. text footnote 3) for the western Okhotsk population, and on unpublished data for the northeast population.
in the shallower, estuarine areas of the
bays, and the lack of sightings at the
exits of the bays suggests these ag-
gregations are relatively isolated from
each other in summer months. How-
ever, two belugas, one with a tag and
another—with the scars caused by
tagging (apparently, the animals that
were tagged in Sakhalinsky Bay in
August 2007 and/or July 2008) were
photographed in Nikolaya Bay in July
2009 (Shpak and Glazov, 2013). It is
unclear whether these belugas moved
between the bays within one summer
(2009), or, in different years, chose
one or the other bay as a summering
ground. In September–October, be-
lugas begin to migrate. Some whales
travel from Sakhalinsky Bay westward
to Nikolaya Bay and may briefly visit
Ulbansky Bay as well (Shpak et al.,
2010; Shpak et al.3). An individual
tagged in the middle of September in
Ulbansky Bay moved to Nikolaya Bay
within a week (Shpak et al.4), which
led us think we had tagged not an Ul-
3Shpak, O. V., D. M. Glazov, D. M. Kuznetsova,
L. M. Mukhametov, and V. V. Rozhnov. 2012.
Migratory activity of the Okhotsk Sea belugas
Delphinapterus leucas in winter-spring period.
Marine Mammals of the Holarctic: Collection
of papers presented at the 7th Int. Conf., Suz-
dal, 2012, 2:390–395 (
4Shpak, O.V., A. Yu. Paramonov, D. M. Glazov,
I. G. Meschersky, and D. M. Kuznetsova. 2018.
Results of the pilot project on the beluga whale
(Delphinapterus leucas) satellite tagging us-
ing remote tag deployment without capturing
whales. Marine Mammals of the Holarctic: Col-
lection of papers presented at the 9th Int. Conf.,
Astrakhan, 2016, 2:271–278 (
bansky summer resident, but one of
Sakhalin-Amur “autumn nomads.” A
single beluga or small groups, some
with immature individuals, are some-
times observed outside areas of ma-
jor concentration, between the bays
or near the Shantar Islands (Solovyev
et al., 2015; our unpubl. data; Fig. 3).
Similar observations (including moth-
ers with calves) have been reported in
the Sea of Japan (Sato and Ichimura,
2011; Melnikov and Seryodkin5; our
interview data) (Fig. 1), but such cases
are rare and thought to be extralimital.
5Melnikov, V. V., and I. V. Seryodkin. 2014.
Sightings of beluga whales (Delphinapterus leu-
cas) in the Sea of Japan. Materials of the Int.
Appl. Sci. Conf.: Habitats, migrations and other
movements of animals, Vladivostok, p. 196–198
[in Russ.].
76 Marine Fisheries Review
Russian-Japanese ship-based sur-
veys conducted in July–August 2009–
10, primarily in offshore waters of the
Okhotsk Sea outside the 12-mile zone,
resulted in no sightings of beluga
whales (Istomin et al., 2013). This sup-
ported past studies and the hypothesis
that in summer belugas concentrate in
coastal waters and bays (Fig. 3). Data
on winter distribution of western Ok-
hotsk belugas is limited to tracking
belugas tagged in Sakhalinsky Bay in
summer (Fig. 3, Shpak et al., 2010,
Shpak et al.3, Shpak and Glazov6).
6Shpak, O. V., and D. M. Glazov. 2013. Review
of the recent scientific data on the Okhotsk Sea
white whale (Delphinapterus leucas) population
structure and its application to management.
In autumn, Sakhalinsky Bay belugas
move offshore north and northeast, but
remain in the Okhotsk Sea throughout
the year, not entering Shelikhov Gulf
or approaching the Kamchatka west-
ern coast. Based on one individual
tagged in Udskaya Bay (Shpak et al.),
it appears these whales stay in the bay,
traveling along the coast until Decem-
ber when the bay starts to freeze. We
speculate that belugas from Sakhalin-
Amur and Udskaya summer aggrega-
tions may meet in December–January
on the way to wintering grounds.
The most recent abundance aerial
surveys in the Okhotsk Sea were con-
Paper SC/65a/SM23 pres. to IWC Sci. Comm.,
Jeju Island, June 2013, 19 p.
ducted in 2009 and 2010 (Shpak and
Glazov, 2013; Shpak and Glazov6;
Glazov et al.7). With the exception of
the southern part of Sakhalinsky Bay
and the Amur Estuary, which were
surveyed in parallel line-transects, all
other regions were covered with a sin-
gle-line coastal survey (direct count).
At the suggestion of the research team,
an availability correction of 50% was
accepted by an IUCN expert panel re-
view (Reeves et al.8). The abundance
estimate for the western Okhotsk pop-
ulation was based on the results from
the aerial survey conducted in August
2010. The corrected abundance was
9,560 belugas. Due to differences in
survey design across the study area,
no coefficient of variation (CV) was
Genetic analyses showed that all
belugas summering in the western
Okhotsk Sea shared a single nuclear
gene pool and thus represent a single
population (Fig. 2). Microsatellite al-
lele frequencies distances between
samples collected in different bays of
the Shantar region were not signifi-
cantly different (Fst = 0.00–0.22%, p
= 0.0723–0.6984). Belugas of Sakha-
linsky Bay differed from other bays
(except for Nikolaya Bay) to a great-
er extent – Fst = 0.41–0.82%, p =
0.0000–0.0020, but these values are
incomparably smaller than differ-
ences between any sample from the
western Okhotsk bays and northeast
Okhotsk (Shelikhov) population (Fig.
4A). At the same time, it was shown
(see Meschersky et al., 2013) that be-
7Glazov, D. M., V. I. Chernook, O. V. Shpak, B.
A. Solovyev, E. A. Nazarenko, A. N. Vasilev, N.
G. Chelintsev, D. M. Kuznetsova, L. M. Mukha-
metov, and V. V. Rozhnov. 2012. The results of
beluga whale (Delphinapterus leucas) aerial
surveys in the Okhotsk Sea in 2009 and 2010.
Marine Mammals of the Holarctic: Collection
of papers presented at the 7th Int. Conf., Suz-
dal, 2012, 1:167–172 (
8Reeves, R. R., R. L. Brownell Jr., V. N. Bur-
kanov, M. C. S. Kingsley, L. F. Lowry, and B. L.
Taylor. 2011. Sustainability assessment of belu-
ga (Delphinapterus leucas) live-capture remov-
als in the Sakhalin–Amur region, Okhotsk Sea,
Russia. Rep. of Independ. Sci. Rev. Panel. Oc-
casional Pap. of the SSC, No. 44. IUCN, Gland,
Switzerland, 34 p. (
Figure 4.—UPGMA-cladogram of Fst-distances found for beluga whales in the
Okhotsk Sea. A) allelic composition of 17 nuclear microsatellite loci, B) presence
and frequencies of the mtDNA control region (559 bp) haplotypes.
81(3–4) 77
lugas summering in different bays
of the western Okhotsk Sea may dif-
fer significantly in mitochondrial lin-
eages composition. Analysis based
on significantly increased sample sets
confirmed these differences (Fig. 4B,
Table 1).
Thus, based both on distribution pat-
terns and results of genetic analysis,
we propose the following subdivision
of the western Okhotsk population
into summer stocks.
Sakhalin-Amur Stock. Belugas of
Sakhalinsky Bay and the Amur Estu-
ary significantly differ by mitochon-
drial-lineage composition from other
western Okhotsk region bays (with the
exception of Nikolaya Bay, see below)
and are designated as the Sakhalin-
Amur stock. This stock is the largest
and best studied of all western Ok-
hotsk summer aggregations. An abun-
dance estimate of the Sakhalin-Amur
stock based on three line-transect
surveys conducted in 2009 and 2010
was 1,977 (CV = 0.24) belugas. Add-
ing a correction for availability bias
(50%) resulted in an overall estimate
of 3,954 (CV = 0.24) belugas (Shpak
and Glazov, 2013).
During the summer, Nikolaya Bay
is occupied with a relatively low num-
ber of belugas (usually less than 100),
and it is unclear whether this aggre-
gation is seasonally resident or if dif-
ferent groups visit the bay throughout
the summer. Belugas of Nikolaya Bay
differ by mitochondrial-lineage com-
position from Ulbansky, Tugursky, and
Udskaya bays but not from Sakha-
linsky Bay (Fig. 4b; Table 1). These
results should be interpreted with
caution, however, due to a very small
sample size that is skewed towards
males from Nikolaya Bay (8 males and
1 female). Nonetheless, further evi-
dence of relatedness between Nikolaya
belugas and the Sakhalin-Amur sum-
mer stock was obtained from photo-
identification studies and behavioral
observations. Therefore, until the sta-
tus of belugas in Nikolaya Bay is con-
firmed with data of sufficient power,
we propose that animals observed in
Nikolaya Bay during the summer be
assigned to the Sakhalin-Amur stock.
Ulbansky Stock. The identity of the
Ulbansky beluga summer stock as a
separate demographic unit within the
western Okhotsk population is based
on the multi-year summer and autumn
observations in Ulbansky Bay and ge-
netic analysis (Table 1). In Septem-
ber–October, before winter migration,
some belugas from Sakhalinsky Bay
move to Nikolaya Bay and may also
visit Ulbansky Bay, but overall beluga
numbers in the inner part of Ulban-
sky Bay seem to decrease in autumn.
Winter migratory routes and feeding
grounds are unknown. In August 2010,
1,167 belugas were counted during a
direct count aerial survey (Shpak and
Glazov, 2013). Abundance, corrected
for availability bias, was estimated at
2,334 belugas.
Tugursky Stock. Although genetic
testing did not separate belugas found
within Tugursky and Udskaya bays
(Fig. 4, Table 1), nevertheless, un-
like the situation with Nikolaya Bay,
we propose the Tugursky belugas as
a separate demographic unit within
the western Okhotsk population. This
designation is supported by historical
information and opportunistic obser-
vations. During summer, belugas are
regularly seen in the southern (estua-
rine) part of Tugursky Bay and some-
times along the western coast, and no
belugas have been observed travelling
between Tugursky and Udskaya bays;
although, small groups have been re-
ported near the south coast of Big
Shantar Island and along the north-
east coast of Tugursky Bay. We noted
behavior differences (e.g., reaction to
boat presence) between beluga groups
in Tugursky and Udskaya bays. Winter
migratory routes and feeding grounds
are unknown. In August 2010, 753
belugas were counted during a direct
count aerial survey (Shpak and Glazov,
2013), with a corrected abundance es-
timate of 1,506 belugas.
Udskaya Stock. Designating the
Udskaya summer stock as a separate
demographic unit within the western
Okhotsk population is based on histor-
ical information, multi-year observa-
tions of beluga summer aggregations
in the bay, and genetic analysis (with
the exception of Tugursky Bay, men-
tioned above). Belugas are present in
the estuarine area from June to Octo-
ber and often enter the Uda River. Be-
lugas are also known to concentrate in
the estuary of the Torom River, about
40 km east of the Uda mouth. There
are no genetic samples from the To-
rom concentration area, but frequent
beluga sightings between the two riv-
ers suggest that all animals belong to
the same stock. Upon ice formation in
the Uda estuary, belugas move further
from the estuary, but remain coastal.
Winter migratory routes and feeding
grounds are unknown. In August 2010,
1,232 (2,464, corrected for availability
bias) belugas were counted during a
direct count aerial survey (Shpak and
Glazov, 2013).
Northeast Okhotsk (Shelikhov)
Recent information on the Shelik-
hov beluga population is very limited.
Most research effort has concentrated
on the western coast of the Kamchat-
ka Peninsula. In the Shelikhov Gulf,
belugas are known to approach river
estuaries during herring, Clupea pal-
lasii; smelt, Osmeridae; and salmon,
Salmonidae, runs. Larger aggregations
were observed in the lower parts of the
bays (Solovyev et al., 2015), but over-
all distribution was more dispersed
along the coastline (Fig. 3) than, for
Table 1.—Conventional F-Statistics from haplotype frequencies: Fst (below diagonal) and P (above diagonal) val-
ues estimated for belugas of dierent bays of the western Okhotsk Sea and the Northeast Okhotsk (Shelikhov)
population. Bold indicates bays that were not statistically dierent.
Sakhalinsky Bay, Nikolaya Bay, Ulbansky Bay, Tugursky Bay, Udskaya Bay, Shelikhov popul.,
n = 183 n = 9 n = 91 n = 45 n = 88 n = 80
Sakhalinsky Bay
Nikolaya Bay
Ulbansky Bay
Tugursky Bay
Udskaya Bay
Shelikhov popul.
0.3253 0.3149
78 Marine Fisheries Review
example, in any of the Shantar region
bays. In summer 2016, a large nurs-
ery aggregation of over 400 belugas
was observed feeding on salmon in
the estuarine part of Gizhiginskaya
Bay, and smaller groups of up to 20,
mostly adults, were encountered along
its eastern coast (Filatova et al.9), the
pattern very much resembling the dis-
tribution observed on aerial surveys
during the period 2009–2010 (Fig. 3).
Beluga form relatively stable summer
aggregations within particular river es-
tuaries (e.g., the Khairyuzova, see Ust-
Khairyuzovo on Fig. 1) on the central
west coast of the Kamchatka Penin-
sula. Only occasional sightings are re-
ported along the southern west coast.
No belugas have been encountered at
the south tip or along the east coast of
the peninsula (Solovyev et al., 2015).
Very little is known regarding
Northeast Okhotsk beluga winter dis-
tribution. In the 1980’s, belugas were
found along the ice edge in Shelik-
hov Gulf and off western Kamchatka
in January–February (Vladimirov and
Melnikov, 1987). Fedoseev (1984) en-
countered belugas in Shelikhov Gulf
also in April a satellite-tagged beluga
remained in Shelikhov Gulf through-
out the winter until April (our unpubl.
data). The ice edge extent largely var-
ies from year to year and often stretch-
es south from Shelikhov Gulf. We
suppose in such cases, Shelikhov belu-
gas remain near the ice edge or along
the west coast of Kamchatka and,
when ice conditions allow, they may
9Filatova, O. A., O. V. Shpak, A. Ju. Paramonov,
D. M. Glazov, A. I. Grachev, and I. G. Mescher-
sky. 2018. Cetacean encounters in the coastal
waters of the northern Okhotsk Sea in the sum-
mer 2016. Marine Mammals of the Holarctic:
Collection of papers presented at the 9th Int.
Conf., Astrakhan, 2016, 2:218–222 (http://mar-
return to the gulf. During a coastal sin-
gle-line survey in August 2010, 1,333
belugas were counted and after cor-
rected for availability bias, abundance
of the Northeast Okhotsk population
was estimated as 2,666 belugas (Shpak
and Glazov, 2013).
Reproductive isolation of belugas
summering in the northeast part of the
Okhotsk Sea, in Shelikhov Gulf and
along the west coast of Kamchatka
Peninsula, from whales summering
in the western Okhotsk Sea was con-
firmed by genetics studies (Fig. 2, 4).
Sightings in Shelikhov Gulf in winter
suggest these whales do not overwin-
ter with the western Okhotsk stocks.
In our study, sample set “Kamchatka”
consisted of 57 individuals (39 males,
15 females, sex of 4 individuals is un-
known) biopsied or found dead dur-
ing the period 2009–12. Whales were
from the west coast of the Kamchatka
Peninsula ranging from the Khairyu-
zova estuary (n = 52) to the lower part
of Penzhinskaya Bay. The sample set
“Gizhiginskaya” consisted of 22 bi-
opsy samples (21 males and only one
female) collected in 2016 in the Gizhi-
ga River estuary in the north and along
the east coast of Gizhiginskaya Bay.
Both sample sets differed from the
Sakhalinsky Bay and Shantar regions
by similarly significant level (Fig. 2;
Table 2).
Although summer aerial surveys
showed discontinuity in the coastal
distribution of belugas between She-
likhov Gulf and western Kamchatka
(Solovyev et al., 2015), no strong evi-
dence for delineating the stocks within
the Northeast Okhotsk population are
available. Differences between the two
population sample sets were found to
be significant, although at a low level:
Fst = 1.46% (p = 0.0006) for microsat-
ellites, and Fst = 22.86% (p = 0.0007)
for mtDNA. However, this may be the
result of inadequate sampling. This in-
cludes multi-year versus single year
collecting, unusual predominance of
males, and low mtDNA lineage di-
versity in the “Gizhiginskaya” sample
set (90% of individuals had the same
haplotype, H = 0.1775 vs. H = 0.5650
found for “Kamchatka”). Moreover,
clustering method analysis showed
only minor differences between “Kam-
chatka” and “Gizhiginskaya” belugas
and determined the two sample sets
were a single genetic cluster. Future
studies will likely find some intrapop-
ulation structuring, but at present all
belugas from the Northeast Okhotsk
population are considered a single
stock (Shelikhov).
Thus, in the Okhotsk Sea, five be-
luga summer stocks should be desig-
nated based on information available
to date: Sakhalin-Amur, Ulbansky, Tu-
gursky, Udskaya, and Shelikhov.
We note that the current gap in be-
luga distribution in the northwest por-
tion of the Okhotsk Sea is a relatively
new pattern. Numerous Soviet litera-
ture sources state that in the first half
of the 20th century, belugas regularly
aggregated in Tauyskaya Bay (Fig. 1),
but after several years of commercial
harvest in early 1930’s, they disap-
peared. The aggregation was either
extirpated or abandoned this summer
ground. In 2008, two unusual sight-
ings of about 1,500 and 100–150 be-
lugas were reported in Tauyskaya Bay
by A.I. Grachev10. Apart from these
two observations, there were very
few encounters of single whales or
small groups in the bay. Without ge-
netic data, we are unable to determine
which of the two Okhotsk Sea popu-
lations may have seasonally occupied
Tauyskaya Bay in the early 20th cen-
tury and during the summer of 2008.
Chukotka Autonomous Region
In Russian waters of the western
Bering and western Chukchi seas, be-
lugas occur near Wrangel Island, along
the entire coast of the Chukotka Pen-
insula, in Anadyr Gulf, and, in winter,
around Cape Navarin (Fig. 1). Only
in Anadyr Gulf, are belugas known to
form a summer resident aggregation
(Arsenyev, 1939; Kleinenberg et al.,
1964). Genetic isolation of Anadyr be-
lugas from the populations of the Ok-
hotsk Sea was shown by Meschersky
et al. (2013), but their original stock
10Grachev, A. I. (deceased) MagadanNIRO, Rus-
sia, unpubl. data.
Table 2.—Frequency based Fst distances (%/%) found
between each of the two Northeast Okhotsk popu-
lation sample sets (Kamchatka and Gizhiginskaya)
and two regions of the western Okhotsk population
(Sakhalinsky Bay/Shantar). All dierences were statis-
tically signicant at p = 0.0000 level.
Marker Kamchatka Gizhiginskaya
Alleles of 17 microsat. loci
mtDNA haplotypes
4.71 / 3.59
31.42 / 31.68
4.55 / 3.08
42.22 / 36.50
81(3–4) 79
designation in 2000 was based mostly
on distribution studies within the Ber-
ing-Chukchi-Beaufort (BCB) region
(IWC, 2000). The status of other be-
lugas observed along Chukotka Pen-
insula in the Chukchi and Bering seas
remains to be confirmed.
In the eastern part of the East Sibe-
rian Sea, belugas are rare. Most coast-
al sightings were near the mouth of
the Kolyma River (Kleinenberg et al.,
1964). Belikov and Boltunov (2002)
mentioned only twelve offshore sight-
ings of belugas during sea ice recon-
naissance for the period 1958–1995.
Satellite tracking of eastern Beaufort
Sea belugas in the 1990’s showed a
westward autumn migration during
which whales travelled as far west as
Wrangel Island (Richard et al., 2001).
In multiple literature sources, belugas
from the BCB region and the East Si-
berian Sea were either grouped into
one population (for ex., Klumov, 1939;
Kochnev, 2003; Belikov et al., 200211),
or subdivided into multiple stocks or
populations (Berzin and Yablokov,
1978; IWC, 2000; and many others),
but either delineation lacked sufficient
grounding. Until the question of popu-
lation structure is clarified, we prefer
to use the term “pool” for BCB belu-
gas. The following sections summarize
Russian studies conducted within this
region, supplementing data collected
on the other stocks in the BCB pool
that have been published in English
and that are readily accessible.
Anadyr Stock
Belopol’sky12 and Pikharev13 de-
scribed the movements of belugas into
the Anadyr Estuary after ice breakup,
and stated that the whales were com-
11Belikov, S. E., A. N. Boltunov, and Yu. A. Gor-
bunov. 2002. Distribution and migrations of ce-
taceans in Russian Arctic according to multiyear
aerial reconnaissance of sea ice and information
from “North Pole” drift station. Marine mam-
mals: Results of research conducted in 1995–
1998. Collection of Pap. Moscow: Mar. Mamm.
Counc., p. 21–51 [in Russ.].
12Belopol’sky, L. O. 1931. Short preliminary re-
port on marine mammals research in Anadyr re-
gion. TINRO archive, cat. #25, 25 p. [in Russ.].
13Pikharev, G. A. 1943. The beluga whale and
pinnipeds in coastal waters of Anadyr region.
TINRO archive, cat. #2585, 29 p. [in Russ.].
monly seen in groups ranging in size
from dozens to thousands. In the
Anadyr Estuary during the ice-free
period, whales concentrated in shal-
low waters and traveled up to 200
miles into the estuary (Litovka, 2002).
Short-term pilot photo-identification
study in 2013 revealed few re-sight-
ings within a season (Prasolova et al.,
2014). Our observations support a hy-
pothesis that in summer belugas form
a resident aggregation in the Anadyr
Estuary (Fig. 5). Telemetry studies
of Anadyr belugas during the period
2001–10 (Litovka et al., 2013; Citta
et al., 2017; Litovka et. al.14,15; Hobbs
14Litovka, D. I., R. C. Hobbs, K. L. Laidre, G.
M. O’Corry-Crowe, J. R. Orr, P. R. Richard, R.
S. Suydam, and A. A. Kochnev. 2002. Research
of belugas Delphinapterus leucas in Anadyr
Gulf (Chukotka) using satellite telemetry. Ma-
rine Mammals of Holarctic: Abstracts of The
2nd Int. Conf., Lake of Baikal, 2002, p. 161–
163 (
15Litovka, D. I., R. C. Hobbs, K. L. Laidre, G.
M. O’Corry-Crowe, J. R. Orr, P. R. Richard, and
R. S. Suydam. 2004. Studying of dive patterns
of belugas (Delphinapterus leucas) in Anadyr-
Navarin region of Bering Sea using satellite
telemetry. Marine Mammals of Holarctic: Col-
lection of papers presented at the 3rd Int. Conf.,
Koktebel’, Ukraine, 2004, p. 327–331 (avail.
et al.16) confirmed and clarified dis-
tribution and movement patterns in
the Anadyr Estuary and Anadyr Gulf,
which had previously been based on
coastal and aerial counts (Litovka,
Belugas spend the ice-free period
(5–6 months) in the Anadyr Estuary.
The latest reported sighting occurred
in late November. As ice forms in
the estuary, belugas leave the Anadyr
River mouth. They move northeast to
feed on smelt, and later migrate to the
exit and southern part of Anadyr Gulf
(Litovka et al., 2013). Results from
telemetry studies (Citta et al., 2017;
Litovka et al.14) and aerial surveys
conducted in April (Litovka et al.17)
suggest Anadyr belugas may mix with
other BCB stocks while in feeding ar-
eas off Cape Navarin in winter–spring
(December–April) (Fig. 5).
No abundance surveys have been
conducted in Anadyr Gulf in sum-
16Hobbs, R. C., D. I. Litovka, and G. M.
O’Corry-Crowe. 2007. Bering Sea wintering
grounds of beluga whales. Seattle, WA. N. Pac.
Res. Board Final Rep. 324:1–12
Figure 5.—Summer and winter ranges of Anadyr belugas based on observations
and satellite-tracking data from published literature sources (see text) and beluga
sightings and group sizes during aerial surveys in April 2005 and 2006 (redrawn
from Litovka et al., text footnote 17).
80 Marine Fisheries Review
mer. Coastal observations suggest two
peaks in sighting rates: 1) from the
end of June to the beginning of July,
and 2) in the beginning of August,
both of which coincide with salmon
spawning. The largest concentration,
241 whales, was recorded in late June
(Litovka, 2002). Pacific walrus, Odo-
benus rosmarus, aerial surveys with
opportunistic beluga counts were con-
ducted in April 2005 and 2006 (Fig.
5). The surveys did not cover the area
south of Cape Navarin, where, ac-
cording to satellite tracking data, part
of the Anadyr beluga stock may have
resided. In April 2005, 162 groups of
410 whales total were counted, and in
April 2006, 195 groups with 403 in-
dividuals total (Litovka et al.17). The
estimated beluga abundance using a
correction for availability (2.86) was
15,127 (7,447–30,741) (Litovka18).
This estimate represents “a theoretical
abundance calculated as direct extrap-
olation of the estimated mean density
17Litovka, D. I. , V. I. Chernook, A. A. Kochnev,
A. N. Vasiliev, A. V. Kudriavtsev, and V. G.
Myasnikov. 2006. Distribution of beluga (Del-
phinapterus leucas) and bowhead (Balaena mys-
ticetus) whales in the northwestern Bering Sea
according to aerial survey performed in April,
2005 and 2006. Marine Mammals of Holarc-
tic: Collection of papers presented at the 4th
Int. Conf., Saint-Petersburg, 2006, p. 323–327
18Litovka, D. I. 2013. Ecology of Anadyr stock
of the beluga whale Delphinapterus leucas (Pal-
las, 1778). Ph.D. Dissert., Voronezh State Univ.,
Voronezh, Russia, 149 p. [in Russ.].
to the unsurveyed areas” (Litovka18)
and should be viewed with caution. In
addition, some of the beluga whales
observed in April may have belonged
to one or several BCB region stocks
given their overlap in winter–spring
range. An estimate of the Anadyr
summer stock of about 3,000 belugas
(Litovka, 2002) appears more realistic
and should be used for management
purposes until a dedicated survey of
the summering area is conducted.
Meschersky et al. (2013) show that
some Anadyr belugas possess mito-
chondrial haplotypes phylogenetically
different from those of maternal lines
common elsewhere in the BCB re-
gion. The differences observed in the
Anadyr sample set (n = 76) from other
BCB stocks, when compared to data
from O’Corry-Crowe et al. (1997), are
strong, although with varying degrees
of significance. For the Beaufort Sea
sample (n = 96) the Fst = 4.48% (p =
0.0011), while for the eastern Chukchi
Sea (n = 113) and Norton Sound sam-
ples (n = 74) the Fst = 31.4%–44.0%
(p = 0.0000), respectively. These re-
sults, together with data on distribu-
tion, support delineation of Anadyr
Gulf belugas as a separate stock.
We also conducted analysis of 17
microsatellite loci for the above men-
tioned Anadyr Estuary sample, a
sample from the Chukotka Peninsula
coast (n = 10), and samples from oth-
er stocks in the BCB region (provided
by the Mammal Genomic Resources
Collection, Univ. of Alaska Museum
of the North (UAM)). These included
three samples from the Beaufort Sea
(UAM:Mamm: 76608, 86886, and one
sample from A. N. Severtsov Institute
collection), eight samples from the
eastern Chukchi Sea (UAM:Mamm:
52221–25, 70322–24, 70326), three
samples from the Diomede islands
(UAM:Mamm: 66416, 98081–82),
five samples from Norton Sound
(UAM:Mamm: 70519–20, 70522–23,
70525), and two samples from Bris-
tol Bay (UAM:Mamm 66633, 83385).
Although, the Anadyr sample was not
significantly different from the Chu-
kotka Peninsula and eastern Chukchi
Sea samples (n > 5) (Fst = 0.97%, p
= 0.0692 and 0.86%, p = 0.12474, re-
spectively), a cluster analysis of the
entire BCB sample suggests two clus-
ters, one represented by Anadyr belu-
gas (Fig. 6).
O’Corry-Crowe et al. (2018) also
obtained similar results showing: 1)
substantial mtDNA differences be-
tween Anadyr and five other major
summer coastal concentration areas
within the BCB region (Bristol Bay,
Norton Sound, Kotzebue Sound, Kas-
egaluk Lagoon, Mackenzie-Amund-
sen); 2) few differences in allelic
composition of eight microsatellite
loci; and 3) moderate differentiation of
Anadyr belugas revealed by clustering
analysis. These authors also concluded
that the belugas of the Anadyr sum-
mering aggregation should be defined
Figure 6.—Results of K = 2 hypothesis (most probable solution) tested by LOCPRIOR-Admixture model for allele frequencies of
17 microsatellite loci. 1 – Anadyr Estuary, 2 – Chukotka Peninsula coast, 3 – Beaufort Sea, 4 – Diomede Isl., 5– Eastern Chukchi
Sea, 6 – Norton Sound, 7 – Bristol Bay.
81(3–4) 81
as a separate stock. Mixing of Anadyr
belugas with other BCB stocks, if it
exists, is more likely to occur with the
eastern Beaufort Sea belugas. This hy-
pothesis is supported both with satel-
lite tracking data (Citta et al., 2017),
which showed that Beaufort belugas
migrate closer to Chukotka Peninsula
and Anadyr Gulf than the other stocks,
and comparatively lower genetic dif-
ferences between Anadyr and eastern
Beaufort Sea samples (Fig. 6).
Bering-Chukchi-Beaufort Pool
In summer, beluga sightings along
the Chukotka Peninsula are relatively
rare. It is not until autumn that most
whales concentrate in the western
Chukchi Sea, and winter and early
spring in the Bering Strait and west-
ern Bering Sea. The sightings in the
eastern part of the East Siberian Sea
are rare, and most occurred during the
period September–October (Kochnev,
2003; Belikov et al.11). The most re-
cent (late September 2002) western-
most sighting of “many” belugas (at
about lat. 69.75° N, long. 164.1° E)
was provided to Kochnev (2003) by
local hunters. This timing coincides
with results from satellite-tracked be-
lugas from the eastern Beaufort Sea
stock (Richard et al., 2001; Hauser
et al., 2014), that in autumn moved
across the Chukchi Sea to the East
Siberian Sea, some approaching Chu-
kotka while others remained offshore
(around Wrangel Island). Kochnev
(personal commun.) noted that such
approaches were irregular and may
be linked to the ice conditions: for
example, they were frequent in the
1950’s–1960’s, but not again until the
1990’s. Belugas may enter the East
Siberian Sea from the Chukchi Sea
by following the mainland coastline,
or approach from the north (Arctic
Ocean), the latter route is supported by
observations and tracking data (Rich-
ard et al., 2001; Hauser et al., 2014;
Melnikov, 2014; Belikov et al.11).
Solovyev et al. (2013) created a
good representative picture of belu-
ga seasonal distribution and numbers
based on the coastal observations from
twelve villages along Chukotka Pen-
insula’s northern, eastern, and south-
ern coasts. The presence of belugas
was low in June, July, and August.
Kochnev (2003) also noted the lack of
large numbers of belugas summering
in the western Chukchi Sea, along the
northern coast of the Chukotka Penin-
sula, and the waters around Wrangel
Island. According to Kochnev (2003),
belugas are absent near Wrangel Is-
land in summer, and along the north-
ern coast of the peninsula most of the
year, except for autumn. The lack of a
historic traditional beluga harvest to
the west of long. 172°W further sup-
ports this observation (Kochnev, 2003;
Bogoslovskaya and Krupnik19). Mel-
nikov (2014) also states that in sum-
mer belugas are rarely observed along
the Chukotka coast, both in Chukchi
and Bering seas.
From all reviewed sources it is clear
that most beluga sightings in terms
of frequency and number occur dur-
ing the spring and autumn migra-
tions. According to Melnikov (2014),
spring migration along the southern
and eastern peninsula coasts starts in
April, and most observations suggest
that belugas approach the southeast
of the peninsula from the Bering Sea
rather than from Anadyr Gulf. These
observations correspond well with the
satellite tracking analysis (Citta et al.,
2017). It is important to note that most
of the data in Solovyev et al. (2013)
and Melnikov (2014) were collected
not by the authors personally but by
“Beringia” National Park local em-
ployees and hired local residents. Al-
though the remoteness of the study
area and lack of comprehensive re-
search make such data invaluable, they
should be treated with caution, and
cross-checking of information should
be applied whenever possible. In addi-
tion, Melnikov (2014) does not always
cite the sources of the data used in the
text and figures.
19Bogoslovskaya, L. S., and I. I. Krupnik. 2000.
Aboriginal harvest of the beluga whale in the
Far East. Marine Mammals of Holarctic: Col-
lection of scientific papers presented at the Int.
Conf., Arkhangelsk, 2000, p. 34–36 [In Russ.]
Whether belugas can winter in the
Chukchi Sea in the leads and polyn-
yas is unknown, and different opinions
have been expressed. Solovyev et al.
(2013) acknowledges this possibility
by noting a lack of “the high level of
herding” as a sign of “migration activ-
ity” (cited from Matishov and Ogne-
tov, 2006) along the northern coast of
the peninsula in late autumn, and the
rapid emergence of belugas follow-
ing the Arctic cod, Boreogadus saida,
in open water areas in winter months.
In contrast, Kochnev (2003) and Mel-
nikov20 do not support this idea, and
Kochnev (2003) specifically notes
that from December to June belugas
are absent along the northern coast
of the Chukotka Peninsula. Sightings
are limited to the northwest cape of
the mainland—Cape Dezhnev (Fig.
1). During autumn months, the home
range of eastern Beaufort Sea belugas
also shifts from north of Wrangel Is-
land toward the Bering Strait (Hauser
et al., 2014).
There are no abundance estimates
for belugas approaching the Chukotka
Peninsula in different seasons, except
for the counts by local observers (So-
lovyev et al., 2013) and opportunistic
counts in the mouth of Anadyr Gulf
during the walrus aerial surveys in
2005–06 (see Anadyr Stock section).
Meschersky et al. (2018), when com-
paring the belugas taken in autumn and
winter in different parts of Chukotka
Peninsula (n = 10) with the data pro-
vided by O’Corry-Crowe et al. (1997),
found that the mtDNA haplotype com-
position of the Chukotka sample was
significantly different from the eastern
Chukchi Sea and Norton Sound sam-
ples, but that it did not differ from the
Beaufort Sea sample. No differences
(Fst = 0.89%, p = 0.2747) in haplo-
type composition were found between
our Chukotka Peninsula and Anadyr
samples. Also, the Chukotka Peninsula
sample microsatellite allele frequen-
cies did not differ either from Anadyr
20Melnikov, V. V. 2012. Cetaceans (Cetacea) of
the Pacific sector of the Arctic: modern distri-
bution, migrations, abundance. Doctor of Biol.
Sci. Dissert. V. I. Ilyichov Pacific Oceanol. Inst.,
Vladivostok, 305 p. [In Russ.].
82 Marine Fisheries Review
or eastern Chukchi Sea samples (Fst =
0.97%, p = 0.0692, and Fst = 0.00%;
p = 0.7614, respectively). Nonethe-
less, according to the results of the
clustering analysis (Fig. 6), Chukotka
Peninsula belugas do not belong to the
Anadyr cluster.
In their recent study, O’Corry-Crowe
et al. (2018) also analyzed 10 beluga
samples from the Chukotka Peninsula
described as northbound spring mi-
grants and obtained similar results. For
the mtDNA, the animals from Chukot-
ka differed from all other BCB sample
sets except the Mackenzie-Amundsen
whales (eastern Beaufort Sea). No sta-
tistically significant differences were
found for microsatellite loci alleles,
but according to different methods of
the assignment testing, Chukotka belu-
gas were assigned to the Beaufort Sea
and not to the eastern Chukchi Sea.
In summary, no definitive conclusions
can be made regarding the status of
belugas migrating along the Chukotka
Peninsula. Therefore, we recommend
they continue to be classified as part of
the Bering-Chukchi-Beaufort pool for
management purposes in Russia, until
larger samples are obtained that sug-
gest otherwise.
Anthropogenic and Natural
Hunting and Live Captures
Total Allowed Takes (TAT) of aquat-
ic biological resources, including ma-
rine mammals, are issued annually in
autumn by the Ministry of Agriculture
and reflect the maximal theoretically
sustainable yield. These documents
are publicly available. Corrections and
changes may be provided to the issued
yields of TAT’s if they comply with
all necessary legal procedures. The
number of licenses requested for tra-
ditional harvests or live-captures may
be obtained from the Federal Agency
of Fisheries, but the actual capture
numbers of marine mammals (at least,
beluga whales), to our experience, are
not reported properly by licensees or
regional fisheries offices.
Okhotsk Sea. Until recently, belu-
ga stock structure in the Okhotsk Sea
has not been taken into account in the
process of calculating TAT’s, which in
the Russian Federation are distribut-
ed across fishing zones and subzones,
whether the latter coincide with stock
boundaries or not. Shpak and Glazov
(2013) presented abundance estimates
and recommended to use PBR with a
recovery factor (F) from 0.5 to 0.65 for
beluga summer stocks of the western
Okhotsk population, and to minimize
beluga takes (limit them to tradition-
al harvests and scientific takes) in the
northeast part of the sea until more data
on the Shelikhov population are avail-
able. The PBR of 42 (F = 0.65) was
suggested for the Sakhalin-Amur stock
with a note that, depending on the actual
2013 take, PBR may have to be recalcu-
lated with F=0.5. It was also recom-
mended that no takes should be allowed
in the little-studied Nikolaya Bay. These
recommendations were partly imple-
mented. Starting in 2017, The Ministry
of Agriculture recommended distribut-
ing the TAT in the Okhotsk Sea among
different summer aggregations/regions
as follows: in the North Okhotsk fishing
subzone, Sakhalinsky Bay and Amur
Estuary – 40, Nikolaya and Ulbansky
bays – 40, Tugursky Bay – 20, Udskaya
Bay – 40, northern coast – 10; and in
the West Kamchatka subzone, Gizhi-
ginskaya Bay – 7, Penzhinskaya Bay –
6, and West Kamchatka coast – 12. We
have compiled information on beluga
live-captures in the Okhotsk Sea start-
ing in 2000 (Table 3).
All beluga captures (Table 3) in the
North Okhotsk subzone were con-
ducted in southern Sakhalinsky Bay
and, possibly in Nikolaya Bay in 2014
and 2015. Boltnev et al. (2016) men-
tions that on average, live-captures of
Sakhalin-Amur belugas do not exceed
30 whales annually, but that the maxi-
mum take was “over 100 animals.” It
is unclear if the authors were provid-
ed information unavailable to us, or if
they summed the numbers of belugas
captured and drowned in 2013 that
were provided in Shpak and Glazov21.
The majority, if not all, live-captured
belugas are immature two- to three-
year old individuals. Usually, the pre-
ferred sex for takes is female.
Quotas for beluga harvest are sel-
dom requested. A quota for tradition-
al hunting of 90 belugas in the North
Okhotsk subzone was issued in 2012,
but no whales were harvested under
this permit (Shpak and Glazov, 2013).
To our knowledge, no belugas in the
Okhotsk Sea have been landed under
traditional harvest quota since 2000.
Based on interviews with local people
in the Shantar region (where three set-
tlements are located) “several” belugas
per settlement are harvested without
permit annually. Most people reported
beluga meat was used to feed dogs in
the winter, and in one village, to be
21Shpak, O. V., and D. M. Glazov. 2014. Update
report on the white whale (Delphinapterus leu-
cas) live captures in the Okhotsk Sea, Russia.
Paper SC/65b/SM14 pres. to IWC Sci. Comm.,
Bled, Slovenia, May 2014, 4 p.
Table 3.—The annual beluga Total Allowed Takes (TAT) for North-Okhotsk/West Kamchatka subzones, and sum-
mary data for actual permanent removals by live-capture (LC, no. of whales) from Sakhalinsky Bay, North-Ok-
hotsk subzone (from Shpak and Glazov text footnotes 6, 21). Supplemented with information received from
the Federal Government-Financed Institution Centre of Fishery Monitoring and Communications (CFMC) for the
years 2000, 2014, and 2015.
Year 2000 2001 2002 2003 2004 2005 2006
10 (161)
Year 2007 2008 2009 2010 2011 2012 2013
Year 2014 2015 2016 2017 2018
1According to the CFMC, all belugas taken in 2000 were landed under commercial harvest quota, while our data repre-
sent live capture numbers. Thus, in total, 26 belugas may have been taken in 2000.
2The number reported (Shpak and Glazov text footnote 21) was estimated based on direct observations and reports by
capture teams. To compare, the beluga take for 2013 reported to the CFMC was eight whales.
81(3–4) 83
consumed by people. In the Sakhalin-
Amur region, we did not hear about
harvesting belugas for food, but there
was mention of several cases of fish-
ermen shooting beluga when the
whales entered salmon traps or ap-
proached nets. We do not have any
information from Shelikhov Gulf and
western Kamchatka. Bogoslovskaya
and Krupnik19 relied on information
from an ethnographer, who reported
that beluga takes in this region were
“occasional” and did not exceed 10
whales per year. Human-caused belu-
ga incidental mortality, as bycatch in
salmon traps, gillnets, and poachers’
sturgeon nets, and ship-strikes, cannot
be estimated. We are aware of several
cases of beluga bycatch in the Sakha-
lin-Amur and Shantar regions. Ship
strikes were not recorded or reported.
Chukotka Autonomous Region. The
beluga harvest during 1915–98 for
Chukotka was summarized by Bo-
goslovskaya et al. (2007). For the 65
years analyzed, an average of 34.2 (SE
= 6.3, median = 18) belugas were tak-
en annually, and only twice (in 1925
and 1950) did the harvest exceed cur-
rently set TAT (200 belugas). Although
the authors mentioned lack of catch
data from one of the villages for many
years, the data in general reflect belu-
ga takes in the Soviet Union period.
In recent decades, TAT’s for Chu-
kotka waters have been calculated as
the abundance estimate multiplied by
a theoretical growth rate of 4% and
precaution coefficient of 0.5 due to
the absence of sufficient data on be-
luga abundance and population pa-
rameters. Abundance estimates of
10,000 belugas for the western Bering
Sea and 4,000 whales for the western
Chukchi Sea (Vladimirov, 1994) have
been used for these TAT calculations.
These estimates were based on differ-
ent sources and should be considered
as an “expert opinion.” In the absence
of exact figures and given possible er-
rors, the abundance of 10,000 belugas
(Boltnev et al., 2016 based on Vladi-
mirov, 1994) was used, and the TAT
for Chukotka waters was set to 200
whales (Table 4).
Based on the data presented above,
we recommend the Anadyr beluga
stock be managed separately. Due to
the lack of information on BCB pool
stock structure and movement pat-
terns, belugas from the BCB pool in
Chukotka waters (Bering Sea, Chuk-
chi Sea, East Siberian Sea) cannot be
further subdivided into management
units. Until more data are available,
we recommend management of the
BCB pool as a single stock and main-
taining the currently used distribu-
tion of the TAT across the four fishing
zones (Table 4).
For the entire Chukotka region, Na-
tive hunters harvested 205 belugas
during the period 2000–17, or, on av-
erage, a little over 11 animals per year.
Apart from 2003, for which no break-
down is available, the majority (83%)
of whales were harvested in the Ber-
ing Strait area (CFZ) and along the
Arctic coast of the Chukotka Peninsula
(CSFZ). Belugas were taken only dur-
ing the spring and fall migrations. The
beluga harvest significantly decreased
after the Chukotka Native harvest be-
gan including larger species of whales
(bowhead whales, Balaena mystice-
tus, and gray whales, Eschrichtius ro-
bustus) and walruses from 2008–10.
The illegal harvest of belugas in
Chukotka is considered insignificantly
small. Beluga hunting requires Native
skin boats, special skills and equip-
ment; however, no more than 25% of
marine mammal hunters possess them.
According to other sources, in 2006,
Chukotka hunters landed 13 belugas,
and none were taken in 2007 (Zdor
and Mymrin22). In 2009, according
to the same authors, six belugas were
landed in the region (Mymrin and
Zdor23). These numbers do not coin-
cide with the numbers available to us
(Table 4). Despite the differences in
the numbers of landed whales, it is
clear that beluga harvest in Chukotka
is far below the TAT set by the Russian
Ministry of Agriculture.
Not included in the numbers pre-
sented above are the whales harvested
during ice entrapments. In Chukotka,
belugas sometimes become entrapped
in ice in Seniavin Strait (Fig. 7). The
most dramatic occasion happened in
December 1984 when about 3,000
22Zdor, E. V., and N. I. Mymrin. 2008. Results of
marine mammals hunting in Chukotka in 2006–
2007 and environment notes. Marine Mammals
of Holarctic: Collection of papers of the 5th Int.
Conf., Odessa, Ukraine, p. 617–619 (http://mar-
23Mymrin, N. I., and E. V. Zdor. 2010. Results
of the marine mammals harvest in Chukotka in
2009. Marine Mammals of Holarctic: Collection
of papers presented at the 6th Int. Conf., Kalin-
ingrad, p. 412–415 (
Table 4.—The annual beluga Total Allowed Takes (TAT) for Chukotka Autonomous Region and total actual land-
ings in each of the four Chukotka shing zones, 2000–16 (from Boltnev et al., 2016, updated by D. Litovka); har-
vest data in 2003 was provided by the CFMC.
Year TAT WBSZ1 CFZ2 CSFZ3 ESSFZ4 Total take
2000 200 2 0 4 0 6
2001 200 3 0 4 0 7
2002 200 1 3 2 0 6
2003 200 ? ? ? ?
2004 200 12 2
12 0
2005 200 10 0
10 0
2006 200 1 1 0 0 2
2007 200 0 3 0 0 3
2008 200 0 6 2 0 8
2009 200 0
50 0 0
2010 200 0 8 0 0 8
2011 200 0 0 0 0 0
2012 200 1 9 8 0
2013 200 0
11 3 0
2014 200 0 0 0 0 0
2015 200 0 3 0 0 3
2016 200 0 2 0 0 2
2017 200 0 2 8 0
2018 200 0 1 4 0
1WBSZ - Western Bering Sea Fisheries Zone (from Koryak coast to long. 175°W).
2CFZ - Chukotskaya Fisheries Zone (from long. 175°W to Cape Dezhnev).
3CSFZ - Chukchi Sea Fisheries Zone (from Cape Dezhnev to Longa Strait).
4ESSFZ - East-Siberian Sea Fisheries Zone (from Longa Strait to the Kolyma River).
5Incomplete data, only spring take is included.
84 Marine Fisheries Review
Figure 7.—Belugas entrapped in ice in Seniavin Strait (photo provided by I. Zagrebin).
whales became entrapped. Mymrin24
described the entrapment and actions
taken by locals and authorities from
13 December when a local hunter first
spotted belugas, until 5 June when
a few belugas were seen for the last
time. Over 500 individuals from the
entrapped aggregation were harvested
by locals during that winter.
Another instance of beluga ice en-
trapment in Seniavin Strait took place
in December 2011. Approximately
100 whales spent the winter in polyn-
yas in Seniavin Strait, and most (if not
all of them) died (Zagrebin25). On 12
January 2012 (Fig. 7), seven belugas
were harvested, and one was found
dead. All harvested whales had empty
stomachs. The next month, two whales
were found dead and three were taken.
24Mymrin, N. I. 2006. Beluga whales (Delphin-
apterus leucas P.) in the ice trap. Bering Strait,
Chukotka. Marine Mammals of Holarctic: Col-
lection of papers presented at the 4th Int. Conf.,
Saint- Petersburg, 2006, p. 377–380 (http://mar-
25Zagrebin, I. A. 2012. The beluga whales (Del-
phinapterus leucas Pallas) in the Senyavina
strait: again in ice trap. Marine Mammals of
Holarctic: Collection of papers of the 7th Int.
Conf., Suzdal, 1:254–257 (
Several belugas still remained trapped
in early April.
Other Potential Threats
Social and industrial development
(fisheries, gold mining and ore trans-
port, oil and gas exploration and pro-
duction) in the Far East, together with
climate change, pose new threats to
belugas, who closely associate with
ice in winter and demonstrate a high
site-fidelity to coastal waters and shal-
low estuarine parts of the bays in sum-
mer. These arising challenges to the
species need be considered when man-
aging the stocks. The Sakhalin-Amur,
Tugursky, Udskaya, and Shelikhov
stocks were commercially harvested
in the Soviet Union (mostly before
the 1950’s). Information on harvest
and original abundances is limited,
and it is difficult to assess whether all
stocks have completely recovered. The
Anadyr stock has never been over-
harvested and is considered stable.
Sakhalinsky Bay and Amur Estuary as
well as Anadyr Gulf are areas inten-
sively exploited by salmon fisheries.
This type of industry has also devel-
oped in all of the bays of the Chukotka
and Shantar regions. Belugas compete
with fishermen, and enter salmon traps
causing conflicts. The current salmon
catch (together with illegal operations)
is likely decreasing the carrying ca-
pacity, at least, in Sakhalinsky Bay. We
do not have information on the inten-
sity of salmon fisheries in the Shelik-
hov region.
Amur River flooding, leading to
washing of human/pet/livestock waste
and chemicals into the estuaries, is
also of concern. Alekseev et al. (2017)
showed that, after a catastrophic flood
in 2013, the health of belugas was af-
fected by infectious and invasive dis-
eases, whose causative agents were
associated with terrestrial animals.
Flooding in the populated flat-coast
lower parts of Uda and Anadyr river
estuaries is also of similar concern.
The gold-mining industry is highly
developed across the entire Okhotsk
Sea coast with the potential threat of
river and estuarine pollution by toxic
substances. Ship strikes have not been
reported, but direct disturbance and
noise should be considered as threats
in, at least, the Sakhalin-Amur area,
Udskaya Bay, Anadyr Estuary, and
Bering Strait. Future climate change
and sea ice reduction will extend the
81(3–4) 85
period of open water and increase the
flow of marine traffic in the Bering
Strait. Whether this will affect belu-
gas is unknown, since their migration
routes and timing may also change.
It is also anticipated that seismic and
military activities in the BCB region,
including the eastern part of the East
Siberian Sea, will increase with longer
ice-free periods.
Revised assessment of beluga abun-
dance, distribution, and population
structure in the Russian Far East led
to us recognizing seven beluga stocks,
five stocks in the Okhotsk Sea: 1)
Sakhalin-Amur, 2) Ulbansky, 3) Tu-
gursky, 4) Udskaya, 5) Shelikhov, and
two stocks in the Chukotka Autono-
mous region waters: 6) Anadyr and 7)
Bering-Chukchi-Beaufort (BCB) pool.
These new designations will help im-
prove management of these stocks
when addressing concerns about natu-
ral and anthropogenic threats.
We are thankful to A. Paramonov, B.
Solovyev, A. Burdin, S. Gorin, I. Za-
grebin, and inspectors of the “Berin-
gia” NP for providing beluga genetic
samples. The samples from Gizhigin-
skaya Bay (Okhotsk Sea, 2016) were
collected by Olga Filatova during the
Pew Marine Fellowship project “Pro-
tecting critical habitats of whales in
the Russian Pacific.” Our unpublished
data presented in this paper were
collected under “The White Whale
Program” funded by the Russian
Geographic Society, and the Project
“Western Okhotsk Beluga Population
Assessment” (2013–16) funded by the
Ocean Park Corporation, Hong Kong.
We thank all GROM participants for
valuable advice and fruitful discussion
during the meeting. We sincerely thank
Kim W. Shelden for a thorough review
and greatly improved language of this
manuscript, an anonymous reviewer
for the aimed comments that helped
to clarify the text, and the Editor for
scrupulous checking and corrections.
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Technical Report
Full-text available
This document provides an update on the status of marine mammals in the circumpolar Arctic from 2015– 2020
Full-text available
The monodontids—narwhals, Monodon monoceros, and belugas, Delphinapterus leucas—are found in much of the Arctic and in some subarctic areas. They are hunted by indigenous subsistence users. In the past, some populations were substantially reduced by commercial hunting and culling; more recently, some populations have declined due to uncontrolled subsistence hunting and environmental degradation. Monodontids are impacted increasingly by human activities in the Arctic including ship and boat traffic, industrial development, icebreaking, seismic surveys, competition with fisheries, and alteration of habitat due to climate change. Since comprehensive reviews in the 1990’s, substantial new information has become available on both species and on changes to their habitat as a result of human activities and climate change. Thus NAMMCO and partners undertook an updated review in 2017. The review recognized 21 extant beluga stocks, 1 extirpated beluga stock, and 12 stocks of narwhals. The available information on each stock regarding population size, depletion level, current and past removals, and trends in abundance was reviewed to determine status. Concern was expressed where the lack of information prevented reliable assessment, removals were thought to be un- sustainable, or the population was deemed at risk of declining even without direct removals by hunting. Beluga stocks of greatest concern are the small stocks in Ungava Bay (possibly extirpated), Cook Inlet (ca 300), St. Lawrence Estuary (ca 900), and Cum- berland Sound (ca 1,100), and the stocks with uncertainty in Eastern Hudson Bay and the Barents-Kara-Laptev Seas. Narwhal stocks of greatest concern are those in Melville Bay and East Greenland.
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Definition of individual beluga whales by the method of photo identification and selection of the main types of markers are presented for beluga whales in the Anadyr estuary, western Bering Sea. The study was conducted in the aggregation of beluga whales in three different sites at Anadyr — in the Anadyr sea port and at ferry docks № 8 and № 10 in the period from August 14 to September 6, 2013 (summer feeding of the whales). The maximum number of whales (40–70 animals) was observed at the ferry dock № 10, 2–6 whales were at the ferry dock № 8, and 1–2 ones — at the port. Mother-calf pairs were observed in all sites, but the portion of adult whales was the highest at the ferry dock № 10. Photocamera Nikon D90 and Nikkor lens (70–200 mm) were used for the photo ID. Mechanical skin damages, infections, spinal ridge traces, and age spots were used as individual markers. In total, 4400 images were obtained, including 155 ones with sides of beluga whales. The markers were observed both on the left side (32 cases) and right side (97 cases) of the whales, but both sides were identified for 13 whales only, including 3 females with calves. Taking into account the risk of reassessment in case of using two sides for identification, the identification is considered as successful for 110 individuals of beluga whale. Almost all identified individuals were met only once during the whole period of observation, only 6 of them were met twice, and only 4 — three times. One whale appeared 3 days in a row: on August 16, 17 and 18, and 5 individuals came 2 days in a row. Such few recurrent meetings may indicate a weak fidelity of beluga whales to certain water areas in the estuary, though maybe the length of study is insufficient, taking into account that a strong fidelity of mother-calf pairs of beluga whales to certain areas of the Anadyr estuary, considered as their hunting areas, was determined by visual observations of Shirshov Institute of Oceanology in the 1980s. Probably, the area at the dock № 10, where the whales form large aggregation, is not assigned to individual animals but is a common feeding area. This large aggregation is similar to the aggregation of beluga whales at Cape Beluzhy of Solovetsky Island in the White Sea. However, these aggregations are fundamentally different by their nature: the reproductive aggregation at Solovetsky Island is distinguished by various forms of social interactions, including sexual activity, while the search and hunting behavior prevail in the feeding aggregation at the ferry dock № 10. The main reason for the beluga whales concentration in this area is dense concentration of freshwater fish and salmons migrating for spawning. Reproductive aggregations of beluga whales are not known yet in the Far-Eastern Seas
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At least five populations (stocks) of beluga whales (Delphinapterus leucas) are thought to winter in the Being Sea, including the Bristol Bay, Eastern Bering Sea (Norton Sound), Anadyr, Eastern Chukchi Sea, and Eastern Beaufort Sea (Mackenzie) populations. Belugas from each population have been tagged with satellite-linked transmitters, allowing us to describe their winter (January–March) distribution. The objectives of this paper were to determine: (1) If each population winters in the Bering Sea, and if so, where? (2) Do populations return to the same area each year (i.e., are wintering areas traditional)? (3) To what extent do the winter ranges of different populations overlap? Tagged belugas from all five populations either remained in, or moved into, the Bering Sea and spent the winter there. Each population wintered in a different part of the Bering Sea and populations with multiple years of data (four of five) returned to the same regions in multiple years. When data were available from two populations that overlapped in the same year, they did not occupy the shared area at the same time. Although our sample sizes were small, the evidence suggests belugas from different populations have traditional winter ranges that are mostly exclusive to each population.
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The results of aerial observations of belugas (Delphinapterus leucas Pall.) in the western Chukchi and eastern East Siberian Seas for the period from April to November were generalized based on Russian literature and unpublished data. The data of observations in the littoral zones of Wrangel Island and Chukotka carried out in 1985 to 2001 are submitted. The author's data, those of native hunters, as well as the published results showed the use of different parts of the seas by beluga, its migrations in the eastern Arctic, and the degree of separating the populations of belugas of the Chukchi and East Siberian Seas. The data obtained attest that the fish does not winter regularly in these seas, and there are no reasons for separation of local Chukchi and Bering populations. A gap in the species range is suggested to take place in the western part of the East Siberian Sea.
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This paper is based on 748 observations of belugas (Delphinapterus leucas) and 382 observations of baleen whales in the Russian Arctic, the majority of the data provided by aerial reconnaissance of sea ice (ARSI). Although the data are not suitable for the estimation of the number and density of the animals, they represent a multi-year (1958-1995) range of observations to update our knowledge on the seasonal distribution and migrations of the species. Belugas inhabit not only shelf waters but also the zone of the shelf slope and the abyssal zone of the Arctic Ocean, where the animals appear mostly in summer. In winter belugas were observed only in the Barents Sea. In June-August, the frequency of beluga observations was highest in the Laptev Sea, which has previously been believed to have considerably lower numbers of beluga than the Kara and Barents seas. Patterns of seasonal distribution and ice cover suggest the existence of a natural border preventing or reducing population exchange between belugas inhabiting the western and eastern parts of the Russian Arctic. A brief review of available data on distribution of the narwhal (Monodon monoceros) in the Russian Arctic is also given. Two species of baleen whales were frequently seen in the Russian Arctic: the bowhead whale (Balaena mysticetus), and the grey whale (Eschrichtius robustus). The majority of such observations were made in the southeastern part of the East-Siberian Sea and the southern part of the Chukchi Sea. In the Bering Sea baleen whales were usually seen near the Chukotka Peninsula, in Anadyr Bay and southeast of it. Whales were usually seen in ice-free water: observations of whales among rarefied ice and near the ice edge were rare. There were considerable annual and seasonal variations in distribution and migrations of baleen whales in the region, probably caused mainly by the dynamics of ice conditions.
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The genetic structure of four summer aggregations of the Beluga Whale, Delphinapterus leucas, in Sakhalin Bay and Udskaya Bay, off the western coast of Kamchatka in the Sea of Okhotsk and in the Anadyr Estuary of the Bering Sea was analyzed through nucleotide sequencing of the mtDNA control region and detection of the allelic composition of nine microsatellite loci in nuclear DNA. It has been shown that each of the aggregations features a unique set of maternal lines, which indicates a high degree of philopatry in this species. Beluga whales of the Anadyr Estuary are genetically isolated from those of the Sea of Okhotsk. Beluga whales of the summer aggregations of Sakhalin Bay and those from Udskaya Bay share a common gene pool and belong to a single population, while the whales that summer off western Kamchatka with great consistency may be attributed to a different population. Comparison of nucleotide sequences of the mtDNA in beluga whales from various waters of the Russian Far East and North America allowed us to propose a hypothesis about how the structure of beluga whale populations formed in the North Pacific during the postglacial period.
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Beluga whales of the eastern Beaufort Sea stock were tagged with satellite-linked time-depth recorders and tracked during summer and autumn in 1993, 1995, and 1997. Whales occupied the Mackenzie estuary intermittently and for only a few days at a time. They spent much of their time offshore, near or beyond the shelf break and in the polar pack ice of the estuary , or in Amundsen Gulf, M'Clure Strait, and Viscount Melville Sound. The movements of tagged belugas into the polar pack and into passages of the Canadian Arctic Archipelago suggest that aerial surveys conducted in the southeastern Beaufort Sea and Amundsen Gulf may have substantially underestimated the size of the eastern Beaufort Sea stock. Ranges of male and female belugas were somewhat segregated in two of the three years of study. In late July of 1993 and 1995, most males were located in Viscount Melville Sound, while females were primarily in Amundsen Gulf. Movement patterns of males tagged in late July in 1997 were different from those of males tagged in early July in 1993 and 1995. In September, belugas migrated westward along the continental shelf and farther offshore in the Alaskan Beaufort Sea. The tracks from 1997 show that the western Chukchi Sea is an autumn migratory destination and that at least some belugas continued their migration south towards the Bering Strait in November. Some conclusions from this study about beluga ecology challenge conventional wisdom, in that estuarine occupation appears to be short-lived, belugas travel long distances in summer to areas hundreds of kilometres from the Mackenzie Delta, an d they do not avoid dense pack ice in summer and autumn.
We describe a model-based clustering method for using multilocus genotype data to infer population structure and assign individuals to populations. We assume a model in which there are K populations (where K may be unknown), each of which is characterized by a set of allele frequencies at each locus. Individuals in the sample are assigned (probabilistically) to populations, or jointly to two or more populations if their genotypes indicate that they are admixed. Our model does not assume a particular mutation process, and it can be applied to most of the commonly used genetic markers, provided that they are not closely linked. Applications of our method include demonstrating the presence of population structure, assigning individuals to populations, studying hybrid zones, and identifying migrants and admixed individuals. We show that the method can produce highly accurate assignments using modest numbers of loci—e.g., seven microsatellite loci in an example using genotype data from an endangered bird species. The software used for this article is available from
The recent evolutionary history, population structure and movement patterns of beluga whales in the western Nearctic were inferred from an analysis of mitochondrial DNA control region sequence variation of 324 whales from 32 locations representing five summer concentration areas in Alaska and north-west Canada. Phylogenetic relationships among haplotypes were inferred from parsimonious networks, and genetic subdivision was examined using haplotypic frequency-based indices and an analysis of variance method modified for use with interhaplotypic distance data. MtDNA relationships were characterized by a series of star-like phylogenies which, when viewed in conjunction with information on haplotype frequency and distribution, suggested a rapid radiation of beluga whales into the western Nearctic following the Pleistocene, and an early divergence of the Beaufort Sea from the Chukchi and Bering Seas subpopulations. Overall nucleotide diversity was low (0.51%) yet all major summering concentrations were significantly differentiated (ΦST= 0.33) from one another. Stratification of samples by gender and age from the three northernmost subpopulations suggested that female cohorts from neighbouring subpopulations were more differentiated than males. Further stratification of adult animals by age revealed that older adults were substantially less subdivided among locations than younger adults, particularly for males, suggesting that dispersal, although limited, is biased toward older adult males. Overall, the patterns of mtDNA variation in beluga whales indicated that the summering concentrations are demographically, if not phyletically distinct. Population structure appears to be maintained primarily by natal homing behaviour, while asymmetries in dispersal may be associated with the type of mating system.