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© RUSSIAN JOURNAL OF THERIOLOGY, 2015
Russian J. Theriol. 14(2): 201–215
al., 1964; Fedoseev, 1986; Shpak et al., 2010; Shulezh-
ko et al., 2013).
In the 20th century, this species was hunted inten-
sively by commercial whalers. The first scientific data
on distribution, abundance, migration and seasonal
movements in the Sea of Okhotsk were collected during
this period of commercial whaling (1930–60s). How-
ever, no comprehensive data on distribution were ob-
tained because research primarily focused on areas
accessible to commercial whalers. Beginning in the
early 1970s, scientific studies focused on beluga whale
abundance and distribution in the Sea of Okhotsk were
conducted during ship-based and aerial surveys. These
Introduction
The Sea of Okhotsk is the southernmost portion of
the range occupied by beluga whales (Delphinapterus
leucas Pallas, 1776) in the western North Pacific (Shpak
et al., 2010). Individual beluga whales have been ob-
served in the Sea of Japan (Sato & Ichimura, 2011;
Melnikov & Seredkin, 2014); however, such cases are
rare and should be considered extralimital. Beluga
whales remain year-round in the Sea of Okhotsk, in
relatively deep, ice-covered, offshore waters during
winter and aggregate in coastal regions and river estu-
aries in the summer (Arseniev, 1939; Kleinenberg et
Summer distribution of beluga whales (Delphinapterus leucas)
in the Sea of Okhotsk
Boris A. Solovyev*, Olga V. Shpak, Dmitri M. Glazov, Vyacheslav V. Rozhnov,
Daria M. Kuznetsova
ABSTRACT: The summer distribution of beluga whales (Delphinapterus leucas) in the Sea of Okhotsk is
typical for boreal and subarctic seas that are seasonally ice covered and support numerous anadromous fish
species. During summer in the Sea of Okhotsk, beluga whales aggregate where rivers flow into estuaries,
gulfs, and bays. Beluga whales are currently found in Sakhalinskiy Bay (in an estuary of the Amur River),
the Shantar region, in portions of Shelikhov Bay, and along the northwestern coast of the Kamchatka
Peninsula. Changes in beluga whale distribution have occurred when compared to results of earlier studies.
In particular, summer aggregations have increased in the Shantar region and along the northwestern coast of
the Kamchatka Peninsula.
KEY WORDS: beluga whales, geographic distribution, Sea of Okhotsk
Boris A. Solovyev [solovyev.boris@gmail.com] Faculty of Geography, M.V. Lomonosov Moscow State University, 1-
12, Leninskie Gory, Moscow, 119234, Russia, A.N. Severtsov Institute of Ecology and Evolution, RAS, 33, Leninsky
prospekt, Moscow 119071, Russia; Olga V. Shpak [ovshpak@gmail.com] A.N. Severtsov Institute of Ecology and
Evolution, RAS, 33, Leninsky prospekt, Moscow 119071, Russia Marine Mammal Council 36, Nakhimovskiy
prospekt, Moscow, 117997, Russia; Dmitri M. Glazov [dglazov@yandex.ru] A.N. Severtsov Institute of Ecology and
Evolution, RAS, 33, Leninsky prospekt, Moscow 119071, Russia Marine Mammal Council 36, Nakhimovskiy
prospekt, Moscow, 117997, Russia; Vyacheslav V. Rozhnov [rozhnov.v@gmail.com] A.N. Severtsov Institute of
Ecology and Evolution, RAS, 33, Leninsky prospekt, Moscow 119071; Daria M. Kuznetsova [belyi.kit@gmail.com]
A.N. Severtsov Institute of Ecology and Evolution, RAS, 33, Leninsky prospekt, Moscow 119071 .
Летнее распределение белухи (Delphinapterus leucas)
в Охотском море
Б. А. Соловьёв, О.В. Шпак, Д.М. Глазов, В.В. Рожнов, Д.М. Кузнецова
РЕЗЮМЕ: Летнее распределение белухи Охотского моря типично для бореальных и субарктичес-
ких морей с сезонным ледовым покровом и значительными запасами анадромных видов рыб. В
летний сезон белуха концентрируется в вершинах заливов, эстуариях в некоторых районах, таких
как Сахалинский залив (Амурский лиман), Шантарский район, отдельные районы залива Шелихо-
ва, северо-западное побережье Камчатки. Существуют некоторые различия между современным
распределением белухи в Охотском море и распределением, описанным по результатам предыду-
щих исследований. Так, роль Шатарского района и северо-западного побережья Камчатки, как мест
летних концентраций белухи, согласно представленному исследованию увеличилась.
КЛЮЧЕВЫЕ СЛОВА: белуха, географическое распределение, Охотское море.
202 B.A. Solovyev, O.V. Shpak, D.M. Glazov, V.V. Rozhnov, D.M. Kuznetsova
Fig. 1. The areas of the shore/ship based and aerial surveys conducted in 2007–2014 to study the distribution of beluga whales.
extensive studies were interrupted in the early 1990s,
and resumed only in recent years.
Based on these earlier studies, during the first half
of the 20th century summer aggregations of beluga whales
regularly occurred in the northern part of the Tatarskiy
Strait, the estuary of the Amur River, Sakhalinskiy Bay,
Nikolaya Bay, Ulbanskiy Bay, Tugurskiy Bay and Ud-
skaya Bay, the estuary of the Ayan River, and Tauyskaya
Bay, Gizhiginskaya Bay and Penzhinskaya Bay (Arse-
niev, 1939; Kleinenberg et al., 1964). In the second half
of the 20th century, researchers enumerated the main
aggregations of beluga whales. They determined that in
the western Sea of Okhotsk, the Amur River and its
estuary provided habitat for the largest population of
beluga whales in this region (Berzin et al., 1988; Berzin,
Vladimirov, 1989). In the eastern Sea of Okhotsk, the
largest population occupied Shelikhov Bay, including
Gizhiginskaya and Penzhinskaya Bays (Berzin at al.,
1988; Berzin, Vladimirov, 1989; Vladimirov, 1994).
Although a significant amount of commercial whaling
203
Distribution of beluga whales
Table 1. The time periods and areas of ship and shore based observations.
Area of observations Year Time period of observations
Aerial observations
2009 05.08–08.08
Sakhalin-Amur region, Shantar region, North-Western region 2009 11.09–13.09
Western Kamchatka and Shelikhov region 2009 16.08–02.09
2010 07.08–08.08
Sakhalin-Amur region, Shantar region, North-Western region 2010 23.08–24.08
Western Kamchatka and Shelikhov region 2010 10.08–19.08
Ship and shore based observations
Gizhiginskaya Bay 2004 27.05–08.06
Penzhinskaya Bay 2010 28.06–08.07
2010 30.07–31.08
2011 13–14.08, 14–18.09
Estuaries of the Khairyuzova and Belogolovaya Rivers
2012 13.07–28.08
Estuary of the Moroshechnaya River 2011 17.08–13.09
2007 19.08–24.08
2008 06.07–16.07
2009 24.07–07.08
2010 09.06 -4.09
2011 07.07–27.08
2013 20.07–21.09
Sakhalinskiy Bay and Amur Estuary
2014 27.07–28.08
2009 27.06–20.07
2010 17.07–02.08
Nikolaya Bay
2012 17.08–26.08
2010 07.07–09.08
2011 08.07–18.08
2012 15.07–03.09
Ulbanskiy Bay
2013 12.08–27.08
2010 21.06–12.09
Tugurskiy Bay 2013 02.08–11.08
2010 23.08–12.09
2012 29.06–14.07, 04–13.09
Udskaya Bay
2013 09.07–01.08
occurred in Tauyskaya Bay, beluga whales were not
observed there in the second half of the 20th century
(Melnikov, 2001).
Our study updates information on the summer dis-
tribution of beluga whales in the Sea of Okhotsk and
identifies changes in distribution that have taken place
within this region.
Material and Methods
This paper presents the results of studies conducted
by the authors in the Sea of Okhotsk in 2007–2014 (Fig.
1). The overall pattern of present-day beluga whale
distribution was based on data collected during summer
aerial surveys in 2009–2010. Information about the
presence of belugas in specific bays and estuaries was
obtained from shore-based observations, observations
from ships and boats, and interviews with local resi-
dents and fishermen. Satellite-linked transmitters at-
tached to individual beluga whales provided location
data that were used to learn about their movement and
migration patterns.
Aerial surveys. Bays of the Sea of Okhotsk and its
coastline were surveyed in 2009 and 2010, excluding
the Kuril Islands chain. Surveys were conducted from
August 3 to September 13 in 2009 and from August 4 to
August 24 in 2010, covering the Sakhalin-Amur and
Shantar regions two times each year. Bays were sur-
veyed using standard line transect methodology while
other coastal areas were surveyed during a single coast-
al transect [see Glazov et al. (2012) for methodology
and beluga whale abundance estimates].
Shore based and ship-based surveys. Shore based
observations and studies of behaviour and local move-
ments of belugas were conducted during the summer in
the western Sea of Okhotsk in the Amur Estuary, Sa-
khalinskiy, Nikolaya, Ulbanskiy, Tugurskiy and Ud-
skaya bays. In the northeastern region, shore and ship
based surveys were conducted in the estuaries of the
Moroshechnaya, Khairyuzova and Belogolovaya riv-
ers, as well as in Penzhinskaya Bay. Research efforts
varied among regions and years (Table 1).
Interviews. Over the seven year period, local resi-
dents, fishermen, and fishing inspectors were inter-
204 B.A. Solovyev, O.V. Shpak, D.M. Glazov, V.V. Rozhnov, D.M. Kuznetsova
viewed about marine mammals seen near settlements
and shore-based industrial facilities. The respondents
were asked the following questions: ‘Did you observe
beluga whales in the vicinity of your place of residence/
work, and if so, when do they arrive in the area and
leave it?’ Answers to the latter question also included
more general responses such as ‘after ice break-up’,
‘they came with fish’ etc. In addition to beluga observa-
tions, the respondents were asked to provide informa-
tion about when fish approach the coast and in what
quantity. Over 150 persons were interviewed in 28
locations, including more than 100 persons in 19 loca-
tions in the western region.
Other sources of information. Earlier published data
co-authored by the authors of this paper — results of
genetic analysis (Meshchersky et al., 2012, 2013; Yazyk-
ova et al., 2012), satellite tracking (Shpak et al., 2010,
2012; Shpak & Glazov, 2013), and photo-identifica-
tion — were included to provide additional insights
about beluga whale distribution. Nikon D90 and D300
cameras with Nikkor 28–300mm f/3.5–5.6 VR and
Nikkor 70–200 mm f/2.8G AF-S VR IF-ED lens were
used for photo-identification. Photographs were ana-
lyzed with IrfanView 4.37 software.
Distribution mapping and analyses. Ship/shore
based and aerial survey data were transformed into a
format compatible with GIS processing software. To
visualize beluga whale distribution patterns, Adobe Il-
lustrator CS 3 and Adobe Photoshop CS 3 PC software
were used with the Natural Earth vector map available
in the public domain as a map base.
For convenience, we divided the Sea of Okhotsk
into the following regions (Fig. 1):
• Shelikhov region — Shelikhov Bay including
Penzhinskaya and Gizhiginskaya Bays;
• Western Kamchatka region — the coastal area
between Lopatka Cape and the Palana River in the north;
• Sakhalin-Amur region — Sakhalinskiy Bay in-
cluding Baikal Bay, Amur Estuary and the northern part
of the Tatarskiy Strait;
• Shantar region — the so-called ‘Shantar Sea’
including Udskaya Bay, Tugurskiy Bay, Nikolaya Bay
and Ulbanskiy Bay;
• Northwestern region — the coastal area between
Udskaya Bay and Shelikhov Bay.
ANOVA was carried out using results of repeated
counts (n=36) as depending variable and geographical
place as grouping variable. The analysis was conducted
in R 2.14 software (2008).
Results
Shelikhov region
Gizhiginskaya Bay — According to evidence of
respondents residing near Nayakhanskaya Bay, beluga
whales approach the coast during the herring (Clupea
pallasii) spawning season in May after sea ice retreats.
The main aggregation of several scores of belugas is
observed in the northern part of the bay with individual
whales or small groups of 2–3 belugas seen along the
eastern coast.
In late May or the first half of June, the smelt
(Hypomesus sp.) run begins lasting 10–14 days (Che-
reshnev et al., 2001). During this time, beluga whales
gather in Avekova Bay. Belugas are also present in
Avekova Bay in July and August during pink (Onco-
rhynchus gorbuscha) and chum salmon (O. keta) runs.
The maximum number of belugas in Gizhiginskaya
Bay observed during the aerial surveys in 2009 and
2010 reached 250–300 whales (Fig. 2). Similar to the
bays of the Shantar region, belugas stay in the estuaries
of the Gizhiga and Avekova rivers in August. However,
during one aerial survey flight in mid-August, a dense
aggregation of over 120 beluga whales was discovered
opposite a rocky shore on the east coast of the Gizhigin-
skaya Bay. Further north another group of over 20
whales was observed, also near the shore on the east
coast of the Gizhiginskaya Bay.
Penzhinskaya Bay — According to evidence of
respondents in Manily, an inland settlement on the
Penzhina River which flows into Penzhinskaya Bay,
beluga whales come here twice a year — in late May or
early June during the smelt run and in the second third
of July and August during the salmon run. Beluga
whales often go from the estuary up the Penzhina River,
reaching Manily settlement and sometimes continuing
upriver. Extreme high tides are observed in Penzhin-
skaya Bay, especially at the river estuaries where tidal
range is up to 12 m. Respondents noted beluga whales
have to adapt to quick changes in water levels and rapid
tidal currents.
As beluga whales move along the coast, they drive
fish to nearshore waters within numerous capes. Water
levels near these capes remain deep enough to allow
whales to stay during low tide. During the rising tide,
whales move along river channels, also searching for
prey near capes. According to evidence of respondents,
beluga whales are occasionally trapped on the mud flats
at low tide and their skin becomes dry, but they remain
calm and wait for the next high tide then swim away.
During summer, local residents in Manily usually
see groups consisting of 15–20 belugas. In general,
beluga whales do not gather in large, dense aggrega-
tions in Penzhinskaya Bay but spread out relatively
uniformly in small groups in the area near the estuary of
the Penzhina River.
During boat and shore based surveys in late June
and early July 2010, individual belugas and small groups
up to 3 whales were observed in the Penzhina River
estuary (Fig. 2). One day a group of ca. 10 belugas of
different ages was observed. The belugas appeared in
the river near Manily settlement at the beginning of
high tide and left at the beginning of low tide. Behav-
iours observed included resting and feeding. Aerial
surveys conducted in the second half of August found
up to 250 beluga whales in the estuaries of the Penzhina
and Talovka Rivers. Belugas were also observed along
205
Distribution of beluga whales
Fig. 2. Summer sightings of beluga whales in Shelikhov Bay during the aerial surveys in 2009–2010.
the entire coast of the bay. Up to 40 beluga whales were
seen along the east coast of the bay, with several juve-
niles among them, and over 120 belugas along the west
coast. The densest aggregation of belugas (about 60
whales) was observed in the central area of the bay near
the Elistratov Peninsula. According to our observations
and interviews of local residents, a small but permanent
aggregation exists in the estuary of the Paren’ River in
the northwestern part of the bay. Up to 20 belugas were
seen here during the aerial surveys.
The pattern of beluga whales distribution in Pen-
zhinskaya Bay is somewhat different from whales in the
Shantar region and Gizhiginskaya Bay — belugas are
sighted not only in the river estuaries but also along the
entire coastline of the bay.
Western Kamchatka region
According to evidence of interview respondents,
beluga whales come to the west coast of Kamchatka
Peninsula twice — the first time in the second half of
May or beginning of June when smelt arrives and the
second time in mid-summer remaining until the begin-
ning or even the end of October; feeding on different
salmon species spawning runs.
Aerial surveys found the largest aggregation of bel-
uga whales in the estuary of the Khairyuzova and Bel-
ogolovaya rivers (Fig. 3: inset), where we spotted up to
250–300 belugas. Smaller aggregations were observed
in the estuaries of the Moroshechnaya River (up to 90
belugas), Tigil River (up to 115 belugas) and Voyam-
polka River (up to 140 belugas). In the estuaries of all
other rivers northeast along the coast from the
Khairyuzova River to the Palana River, beluga whales
were seen only in relatively small numbers. We infre-
quently observed individual belugas and belugas in small
groups along the coast between river estuaries. Often
they were travelling along the shore near the surf line.
Boat and shore based observations occurred in July-
September 2010–2012 in the estuaries of the Khairyuzo-
va, Belogolovaya and Moroshechnaya Rivers (Table
1). Numbers of belugas in the Moroshechnaya River
estuary fluctuated notably — from 11 to 111 belugas
during the observation period. In the Khairyuzova and
Belogolovaya estuary and the adjacent coastal area, the
numbers of beluga whales remained more stable (250–
300 whales).
According to evidence of fishermen and local resi-
dents, beluga whales are not seen in large numbers or
with any regularity south of the Moroshechnaya River.
During our observation flights the 1st week of Septem-
ber 2009, belugas were observed near almost every
river mouth within this area (Fig. 3), but in August
2010, only few were detected along this part of the
206 B.A. Solovyev, O.V. Shpak, D.M. Glazov, V.V. Rozhnov, D.M. Kuznetsova
Fig. 3. Summer sightings of beluga whales near the coast of the Kamchatka Peninsula during ship-based and aerial surveys in
2009–2012.
Kamchatka coast. Sightings in 2009 occurred near the
Sopochnaya River (29 belugas), Icha River (1 beluga),
Oblukovina River (6 belugas), Kolpakova River (8
belugas), Bolshaya Vorovskaya River (17 belugas),
Kikhchik River (35 belugas), and Bolshaya River (ca.
50 belugas). Almost three times as many beluga whales
were observed along the peninsula north of Moroshech-
naya River when compared to the peninsula south of the
river (ANOVA, p = 0.05).
Beluga whales were not seen along the eastern coast
of Kamchatka from Avachinskaya Bay to Lopatka Cape
during the aerial surveys in 2009. Also, no sightings
were reported in these areas by other researchers or
respondents.
Photo-identification studies confirmed movement
of some individuals between estuaries, supporting the
idea that belugas aggregating in the river estuaries of
western Kamchatka may move along the coast during
the summer season, presumably, depending on feeding
conditions.
Similar data were collected using satellite tracking
in the Khairyuzova River estuary (3 transmitters de-
ployed in 2011) and the Moroshechnaya River estuary
(1 transmitter deployed in 2011) (Tarasyan et al., 2013;
Shulezhko et al., 2013). When we compared tracking
data to fish run strength (data based on interviews),
whales appeared to travel to another estuary only if
there was a shortage of fish. In late August 2010, beluga
207
Distribution of beluga whales
whales temporarily left en masse the estuary of the
Khairyuzova and Belogolovaya rivers after the pink
salmon run ended, traveling north to the Kovran River
where the pink salmon run was beginning. Also, ac-
cording to evidence of residents of Ust-Khairyuzovo
settlement, it seemed that beluga whales left their favou-
rite feeding places in the estuaries of the Khairyuzova
and Belogolovaya Rivers in September 2011, when
there were few fish in the river, and came back later that
year. In October – early November 2010 and 2011, the
satellite tracked belugas began slowly moving north to
the estuary of the Lesnaya River (in 2010) or the south-
ern part of Penzhinskaya Bay (in 2011).
Sakhalin-Amur region
According to evidence of numerous interview re-
spondents, beluga whales arrive in Sakhalinskiy Bay
and the Amur Estuary right after the area becomes clear
of ice. Typically this happens in the middle or the
second half of May or sometimes later — in the begin-
ning of June (depending on ice conditions) ‘together
with various species of fish’ (usually herring or smelt).
The whales arrive from the north — northern part of
Sakhalin Island (the area near Rybnovsk) and from the
south — the direction of Tatarskiy Strait. If there is no
fish, the belugas leave the region. Fishermen reported
that the belugas remain in the area during several days,
“blocking” the channel at the southernmost point of
Zotova Bank, then leave following the fish into the
Amur Estuary. After the end of the fish run, the whales
move north before returning to the bay with the next
fish run. In May, groups composed of ‘several dozen’
of beluga whales feed on herring as far south as De-
Kastri settlement.
According to our observations, in summer the main
aggregation of belugas is located on the boundary be-
tween Sakhalinskiy Bay and the Amur Estuary — be-
tween Baydukova Island and Zotova Bank (Fig. 4).
Here the bay is at its most narrow, forcing returning
salmon run into a confined space which belugas take
advantage of from July to September. Aerial survey
results show over 1000 beluga whales aggregate there
in the beginning of August (Glazov et al., 2012). South
of the main aggregation, two significantly smaller and
less dense aggregations were observed (up to 100–200
animals) spreading out along the navigation channels
along the eastern and western shores of the Amur Estu-
ary and entering the lower course of the Amur River. In
the beginning of September, individual belugas were
spotted south of the Amur Estuary near Pronge Cape.
During salmon runs, aggregations of beluga whales
were seen regularly at the entrance of Baikal Bay on the
northwestern coast of Sakhalin Island. Aerial survey
results from 2009 and 2010 show 30 to 90 animals in
this area (Fig. 4).
Fig. 4. Summer sightings of beluga whales in the Sakhalinskiy Bay and the Estuary of the Amur River according to the results
of the ship-based and aerial surveys in 2007–2013.
208 B.A. Solovyev, O.V. Shpak, D.M. Glazov, V.V. Rozhnov, D.M. Kuznetsova
Fig. 5. Summer sightings of beluga whales in Shantar Region according to the results of the ship-based and aerial surveys in
2009–2013.
We observed no beluga whales in the northern quar-
ter of Sakhalinskiy Bay, and according to the satellite
tracking data, in summer beluga whales did not go to
the deep-water areas of the bay. Fishermen mentioned
that beluga whales leave the Sakhalin-Amur region
when autumn chum salmon run ends in September or
October, and this was also confirmed by the satellite
tracking data. Some individual whales traveled from
Sakhalinskiy Bay west to the adjacent bays in the Shan-
tar region (Nikolaya and Ulbanskiy, Fig. 5) and spent
autumn there before joining others, which moved di-
rectly northeast to offshore wintering areas (Shpak et
al., 2010, Shpak et al., 2012; Shpak & Glazov, 2013).
Winter migration started upon beginning of ice-forma-
tion; before that tracked beluga whales kept in shallow
waters of the bays and near the coastline.
Shantar region
Nikolaya Bay — According to evidence of inter-
view respondents, beluga whales arrive in Nikolaya
Bay in the second half of May or early June when
capelin (Mallotus villosus), smelt and herring are spawn-
ing. In the second half of June, the whales return to the
Usalgin River estuary when salmon runs start.
According to our data, from late June to mid-July,
individual whales are seen in Nikolaya Bay (up to 10–
15 animals), and in the second half of July the numbers
increase up to 50 whales as pink salmon and chum
salmon runs start. Beluga whales were primarily in the
southern part of the bay in the Usalgin River estuary
(Fig. 5). They disperse in the area in small groups or
individually. With the beginning of high tide they may
go 3 or 4 kilometres (km) up the river, but they return to
the bay before the falling tide.
Aerial survey observations included 37 and 54
beluga whales in Nikolaya Bay in the first half of
August 2009 and 2010, respectively; however, only 6
belugas were spotted there in the beginning of Sep-
tember 2009. In the second half of August 2012, only
individual whales were spotted in the bay. As men-
tioned in the Sakhalin-Amur region section, satellite
tracking data show that from late August to early
September some beluga whales move from Sakhalin-
skiy Bay to Nikolaya Bay. Some tracked individuals
remain there until the end of autumn when ice forma-
tion begins. Also, some beluga whales move further
west to Ulbanskiy Bay. Two beluga whales, tagged a
year earlier in Sakhalinskiy Bay, were photographed
in Nikolaya Bay next summer. Thus, there is some
exchange between the aggregations of beluga whales
summering in these regions. It is not clear whether
beluga whales migrate between the two bays within
one season, or if they spend the entire summer within
209
Distribution of beluga whales
one bay before moving to another the following sum-
mer.
Ulbanskiy Bay — According to evidence of respon-
dents, beluga whales appear in small number in Ul-
banskiy Bay in May, immediately after breakup of the
fast ice even though the entrance of the bay remains
covered with dense ice up to the middle of July. Beluga
whales move south into the bay in large numbers in the
3rd–4th week of July and disperse within 10 km of the
Ulban and Syran River mouths. Groups of 10–30 belu-
gas are also seen along the western coastline of Ul-
banskiy Bay and the southern coast of Konstantina Bay.
During the aerial surveys in August, an estimated
1200 beluga whales occupied the estuary where the
Ulban and Syran Rivers enter Ulbanskiy Bay (Fig. 5).
Whales were in several large aggregations of 100–300
belugas.
Beluga whale behaviours were noted during ship
and shore based observations collected in Ulbanskiy
Bay. These behaviours included traveling, feeding, rest-
ing, and socializing. Feeding and movements between
different feeding areas within the estuary, or in connec-
tion with high and low tides were observed most often.
A specific behaviour associated with avoiding preda-
tors — killer whales (Orcinus orca (Linnaeus, 1758)),
was observed twice — the beluga whales moved rapid-
ly in surf keeping as close to shore as possible.
The age structure of the beluga aggregations in
Ulbanskiy Bay (based on skin colour: Kleinenberg et
al., 1964; Matyshev & Ogniotov, 2006) is similar to
large summer aggregations from other bays in the Shan-
tar region and along the Kamchatka Peninsula. White
belugas (adults) make up 60–80% of the total number
of belugas in the aggregation (Fig. 5). The dark-grey
belugas (yearlings and young-of-the-year) make up about
10%, but the age-ratio in certain groups of the aggrega-
tion may be very different: there are groups formed
almost entirely by females with juveniles.
Tugurskiy Bay — According to results of coastal
observations, in the Tugur River estuary beluga whales
are seen in small numbers, no more than 20 belugas, in
June or the first half of July. By the last days of July or
early August, their numbers begin to increase. Results
of the aerial surveys in August 2010 show up to 750
beluga whales in this area (Fig. 5). Results from our
ship-based surveys in early August 2013 were similar —
at the peak of the salmon run the river mouth was
“blocked” by an aggregation of over 500 beluga whales
lined in ranks. Whales were observed in the estuary
primarily during high tide as the low tide exposes thou-
sands of meters of mudflats.
The number of beluga whales reduces notably, to
several dozen only, a few days after the end of the peak
of the salmon run. We observed beluga whales in the
area until the end of September (no observations were
carried out later). According to evidence of our respon-
dents, in the eastern ‘pocket’ of the bay near the Tugur-
skiy Isthmus, beluga whales are present in mid-Octo-
ber; and small groups of belugas (up to 20 whales) are
frequently seen in Mamga Bay (70 km north of the
Tugur River estuary). During the aerial surveys in 2009
and 2010, individual belugas were seen along the entire
west coast of Tugurskiy Bay. It is presumed that beluga
whales prefer the western part of the bay because the
eastern shore lacks spawning rivers (S. Kulbachniy,
KhabTINRO, pers. comm.).
Similar to other bays and regions, the age structure
of large aggregations in Tugurskiy Bay included all age
classes during all years of observation (Fig. 6) while
small groups of adult beluga whales were not uncom-
mon.
Udskaya Bay — Respondents from the settlement of
Chumikan (on the Uda River) noted that in spring,
beluga whales enter the river estuaries as soon as the
sea ice retreats. For example, in 2014 the sea ice cover
broke unusually early, and beluga whales were seen
near the settlement in late April for the first time. We
hypothesize that beluga whales enter the bay along the
west coast because these coastal areas are the first to
become ice-free within Udskaya Bay. In other years,
beluga whales first arrive in early or mid-June. Beluga
whales are always present in Udskaya Bay at the start of
the smelt run. Based on our observations and inter-
views, in the middle or by the end of July, the number of
beluga whales peaks and remains at this level, with
some fluctuations, until early or mid-September. In
autumn, the number of belugas near Chumikan gradual-
ly falls. Based on coastal observations, up to 100–120
beluga whales remain until ice slush starts to form,
usually in the middle of November.
During the first third of August 2009, aerial surveys
found beluga whales concentrated in two areas: 850 whales
in the estuary of the Uda River near Chumikan and 100
whales in the Torom River estuary (Fig. 6). During this
same period in 2010, all beluga whales (about 1200
animals) were observed in the Uda River estuary.
The surveyed aggregation of beluga whales appears
to be a large reproductive herd (Fig. 6). In groups
concentrating near the coast, the number of juveniles
reaches 30–40%. Many newborns are observed in the
second half of July. Re-sightings of photographed indi-
viduals with scars from biopsy tips confirm the recur-
ring presence of the same individuals in these coastal
groups. Similar to the adjacent bays in the Shantar
region, beluga whale movements within the estuaries of
Udskaya Bay are effected by tides. Extreme low tides
expose mudflats and displace whales several kilome-
tres from the river mouths.
Thus, according to evidence of respondents, small
numbers of beluga whales appear immediately after
break up of shorefast ice in the Shantar region. Evident-
ly, they are able to traverse ice fields that may block the
entrances to some bays (e.g., Nikolaya Bay, Ulbanskiy
Bay). The early arrival of beluga whales coincides with
the beginning of herring, smelt and capelin runs.
By the middle of summer beluga whales gather in
and near the estuaries of large rivers. The largest aggre-
gations of beluga whales (over 1000 animals) were
210 B.A. Solovyev, O.V. Shpak, D.M. Glazov, V.V. Rozhnov, D.M. Kuznetsova
Fig. 6. The age distributions of several summer aggregations of beluga whales in the Sea of Okhotsk determined during the
ship and shore based observations in 2010–2013.The areas and methods of observation are shown on the vertical axis.
observed in Ulbanskiy Bay and Udskaya Bay, while the
smallest were in Nikolaya Bay.
Movement patterns within the estuaries are related
to tidal sea level changes, which vary in this region in
summer from 3.5 m to 7.8 m.
The social structure of the aggregations was vari-
able and depends, most likely, on the pattern of food
distribution. Large groups were comprised of hundreds
of whales that included subgroups of whales of 3–30
animals in close proximity to one another. Isolated
small groups consisting of several belugas were also
seen along the coastlines outside of the river estuaries.
Northwestern region
Seemingly, at present there are no permanent sum-
mer aggregations of beluga whales anywhere along the
northwestern coast of the Sea of Okhotsk between Ud-
skaya Bay and Shelikhov Bay (Fig. 7). During the 2009
aerial survey, only individual whales or small groups
travelling along the coast have been observed. Beluga
whales were seen in two locations: in the estuary of the
Inya River on the administrative border between Kha-
barovskiy Territory and Magadanskaya Region (one
adult with a juvenile) and in the western part of
Tauyskaya Bay (two groups comprised of 9 adults with
2 juveniles and one adult with a juvenile).
In late May – early June 2008, when ice was still
present near the coast, up to 1500 beluga whales en-
tered Tauyskaya Bay and dispersed over the entire area,
later leaving and heading east. Almost no feeding be-
haviour was observed during this time. The same year
in late June, a group of 100–150 beluga whales ap-
peared to be feeding on the bottom, probably on flatfish
(Pleuronectidae), near Ola settlement, about 6–8 km
from the river estuary (A.I. Grachev, Magadan NIRO,
pers. comm.).
According to evidence of respondents (from Tuk-
chi, Ayan and Okhotsk), beluga whales approach the
northwestern coast in the spring following the breakup
of fast ice, as spawning herring and smelt return. For
example, 12 belugas were seen in Ayan Harbor on
April 30, 2014.
On May 26–27, 2011, an aerial ichthyologic survey
team observed ca. 270–300 belugas in groups of 10–30
animals in the herring spawning areas near Okhotsk
town (A.I. Grachev, Magadan NIRO, pers. comm.).
Discussion
According to our data, beluga whales are not uni-
formly distributed along the entire coast of the Sea of
Okhotsk during summer, but aggregate primarily in
river estuaries and associated bays located in several
211
Distribution of beluga whales
Fig. 7. The summer distribution of beluga whales in the Sea of Okhotsk.
regions. These regions include Sakhalinskiy Bay (and
the estuary of the Amur River), continental bays of the
Shantar region, some areas of Shelikhov Bay, and the
northwestern coast of the Kamchatka Peninsula. Only
rare occasional sightings of beluga whales occurred
along the northwestern coast of the Sea of Okhotsk
between the Shantar and Shelikov regions, in Tatarskiy
Strait south of De Kastri settlement, near the eastern
coast of Sakhalinskiy Island, and along the coast of the
Kamchatka Peninsula south of the Moroshechnaya Riv-
er. Similar to studies conducted in the 20th century, we
have not seen beluga whales in summer near the east
coast of Kamchatka, the coast of East Sakhalin and in
the northern part of the Sea of Japan. Although, beluga
whales have been observed near the continental coast
of Primorye region, the coasts of Hokkaido Island, and
even the coast of Honshu Island (Sato & Ichimura,
2011; Melnikov & Seredkin, 2014), most sightings
were of lone whales, “outliers” encountered between
seasons in the period of active migrations.
Numbers of beluga whales found within each region
have changed since the time of commercial whaling and
scientific studies conducted during the 20th century.
When comparing the Shantar region to the Sakhalin-
Amur region, we found that in early–middle 1980s the
numbers in these regions were in a ratio of 1:4, by the end
212 B.A. Solovyev, O.V. Shpak, D.M. Glazov, V.V. Rozhnov, D.M. Kuznetsova
of the 1980s and in the 1990s this ratio changed as 1:2;
however, by 2011 this ratio was 2:1 (Vladimirov, 1985,
1994, 1995; Berzin et al., 1988, 1990; Glazov et al.,
2012). According to V.L. Vladimirov (1994, 1995), the
ratio change in late 20th century was a result of a decline
in number of beluga whales in the Sakhalin-Amur region.
Within the Shantar region, present day estimates in
each bay also differ greatly from beluga counts ob-
tained 30–50 years ago. For example, more beluga
whales were present in Tugurskiy Bay than in Ul-
banskiy Bay (Berzin et al., 1988), while current esti-
mates show almost three times as many whales in Ul-
banskiy compared to Tugurskiy Bay. Although visual
observations support the hypothesis of seasonal resi-
dency, the movements of Shantar belugas between the
bays can not be excluded, given that satellite-tagged
and photo-identified whales from the Sakhalin-Amur
summer aggregation are known to travel to the eastern
Shantar bays. Genetic analysis (Yazykova et al., 2012;
Meshchersky et al., 2013) indicates that there are statis-
tically significant differences in maternal lines in sepa-
rate bays, except Nikolaya and Sakhalinskiy bays. As
summer aggregations of beluga whales demonstrate
pronounced philopatry fixed at the genetic level, it is
unlikely that these aggregations travel from bay to bay
and mix in numbers. We suppose there may be certain
exchange between Sakhalinskiy and Nikolaya bays in
summer months; but for most aggregations, highly res-
idential behavior is characteristic during the months of
chum and pink salmon runs, when most genetic sam-
pling took place. In autumn, belugas become more
mobile. We know that individual whales start moving
from bay to bay, and that the number of belugas in
places of summer concentration gradually decreases.
The autumn movements of entire aggregations remain
unknown, and it is possible that large summer aggrega-
tions disperse breaking into smaller groups, some of
which remain on summer grounds until the winter mi-
gration starts.
According to the papers of the late 20th century
(Berzin et al., 1985; Berzin, 1995), most beluga whales
observed in the northeastern Sea of Okhotsk were in the
Shelikov region, concentrated in Penzhinskaya and Gizh-
iginskaya bays, with only small groups and individual
belugas seen along the western coast of Kamchatka.
Our studies show the numbers of beluga whales near
the western coast of Kamchatka and in the Shelikhov
region are approximately the same (Glazov et al., 2012).
Tagged whales from the Kamchatka coast were ob-
served traveling to the Shelikov region after the sum-
mer season suggesting there may be migratory move-
ments between these two regions. Genetic analyses of
samples collected along the west coast of Kamchatka
(Meshchersky et al., 2012, 2013) compared to whales
from the Sakhalin-Amur and Shantar regions suggest
these populations are distinct one from another. Along
the northwestern and northern coast of the Sea of
Okhotsk, summer aggregations are no longer present.
Prior to the 1930s, several hundred beluga whales were
regularly seen in Tauyskaya Bay (Govorkov, 1930,
1931). Commercial whaling of belugas began in this
bay in 1929, with 388 belugas caught in 1929, and 147
in 1930 (Govorkov, 1930, 1931). It is unknown wheth-
er the aggregation in Tauyskaya Bay was extirpated, or
belugas left this area. Haplotypes characteristic of the
northeastern population are present in belugas aggre-
gating in Udskaya Bay (Meshchersky et al., 2012, 2013).
It is possible that Tauyskaya aggregation abandoned its
summer ground and mixed with belugas from Udskaya
Bay. Intensive commercial harvest in Sakhalin-Amur
region, Tugurskiy and Udskaya bays and Gizhigin-
skaya Bay, which lasted longer than whaling in
Tauyskaya Bay, did not result in the disappearance of
the respective summer aggregations of beluga whales
suggesting that these areas play key role as feeding or/
and breeding habitats similar to distribution pattern of
Cook Inlet belugas described below.
The summer distribution and behaviour of beluga
whales in the Sea of Okhotsk is typical for the subarctic
and temperate zone seas. For example, belugas form
similar summer aggregations in the Anadyr Estuary of
the Bering Sea. Anadyr belugas also approach the coast
after the ice breaks up in late spring or early summer,
feed intensively on anadromous species of fish, and
leave the bay when in October ice starts to form (Litov-
ka et al., 2002).
The same pattern is also observed in the eastern part
of the Bering Sea — in Norton Sound, where beluga
whales aggregate in the estuary of the Yukon River
from May to November, also feeding on salmon (Low-
ry et al., 1986). In Bristol Bay, Alaska, belugas gather
in the Kvichak and Nushagak rivers from April to
August following anadromous fish runs (Lowry et al.,
2008). A small isolated population of beluga whales in
Cook Inlet, Alaska, currently remains in the inlet year-
round and aggregates in river deltas during the summer
months to feed on anadromous runs of eulachon (Tha-
leichthys pacificus) and salmon and find shelter from
killer whales (Shelden et al., 2003, Goetz et al., 2007,
2012). Similar to the whales in some regions of the Sea
of Okhotsk, these whales were intensively hunted but
did not abandon the Susitna delta area (Rugh et al.,
2010). Instead as their numbers declined overall, num-
bers in this particular area remained fairly stable sug-
gesting this aggregation area provided habitat preferred
over other areas, even though this is where most hunt-
ing occurred. In the subarctic and northern seas of the
Atlantic, beluga whales are similarly distributed in sum-
mer in the Gulf of Saint Lawrence where they go up the
estuary of the Saint Lawrence River after the ice re-
treats (Kingsley, 1999; COSEWIC, 2004), as well as in
the western part of Hudson Bay, where beluga whales
gather in the estuaries of the Churchill, Nelson and Seal
rivers to feed on salmon (Sergeant, 1973).
The distribution patterns and behaviour of beluga
whales in the Arctic seas are notably different. Estua-
rine and coastal waters play much less role in belugas
summer distribution in the Arctic seas. When beluga
213
Distribution of beluga whales
whales enter a bay, river estuary, or near shore waters,
they go in small numbers or for a short period of time,
without forming stable aggregations gravitating to cer-
tain locations. These whales also typically undertake
much longer migrations covering thousands of kilome-
tres from their summering grounds to their wintering
areas. For example, this is the case in such locations as
Obskaya Bay, Yeniseiskiy Bay, the Lena River estuary,
Kasegalyuk Bay, Kotzebue Sound, the Mackenzie Riv-
er estuary, and Somerset Island (Norton & Harwood,
1986; Frost et al., 1993; Smith & Martin, 1994; Rich-
ard et al, 2001a, b; Solovyev et al., 2011; Udovik et al.,
2012). These regions do not host large, anadromous
fish runs and remain ice-covered for longer periods
than the subarctic and boreal seas. In a warming arctic,
there is already evidence of salmon numbers increasing
farther north than ever documented in the past (Dun-
mall et al., 2013). It will be interesting to see how these
changes may affect the behaviour of these northern
populations of belugas as sea ice disappears and anadro-
mous fish become a part of their ration.
ACKNOWLEDGEMENTS. The study was conduct-
ed as cooperative research of the Program ‘The White
Whale’ of the Permanent Expedition of the Russian
Academy of Sciences (RAS) (A.N. Severtsov Institute
of Ecology and Evolution, RAS) and the Project ‘Cur-
rent status of the Sakhalin-Amur beluga aggregation
(Sea of Okhotsk, Russia): sustainability assessment’
(“Dolphin and I” Ltd, Marine Mammal Council) with
the financial support of the Russian Geographical Soci-
ety, Ocean Park Corporation (Hong Kong); Georgia
Aquarium Inc., Seaworld Parks & Entertainment, Mys-
tic Aquarium and Institute for Exploration (USA); Ka-
mogawa Sea World (Japan).
The authors would like to thank all the participants
of the field studies on the distribution of beluga whales
in the Sea of Okhotsk — V.I. Chernook, E.A. Nazaren-
ko, A.N. Vasiliev, A.I. Grachev, A.Yu. Paramonov,
D.I. Ivanov, D.A. Udovik, T.C. Shulezhko, K.K. Tara-
syan, F.V. Kazansky, A.N. Kanzeparova, S.E. Kul-
bachniy, E. Molchanova, R. Michaud, M. Moisan. The
authors would like to thank Kim Shelden and Olga
Filatova for comments and suggestions, which have
greatly improved the manuscript, and Dr. Shelden’s
extraordinary help on English correction.
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