ISSN 1062-3590, Biology Bulletin, 2006, Vol. 33, No. 1, pp. 53–58. © Pleiades Publishing, Inc., 2006.

Original Russian Text © V.V. Krasnova, V.M. Bel’kovich, A.D. Chernetsky, 2006, published in Izvestiya Akademii Nauk, Seriya Biologicheskaya, 2006, No. 1, pp. 63–69.

ZOOLOGY

Mother–Infant Spatial Relations in Wild Beluga

(Delphinapterus leucas) during Postnatal Development
under Natural Conditions
V. V. Krasnova, V. M. Bel’kovich, and A. D. Chernetsky
Shirshov Institute of Oceanology, Russian Academy of Sciences, Nakhimovskii pr. 36, Moscow, 117997 Russia
e-mail: adcher@sio.rssi.ru
Received May 18, 2005

Abstract—Elements of behavior under natural conditions, their duration, and frequency are described in three
age groups of belugas calves: newborn, one-month-old, and two-month-old. The quantitative and qualitative
indices of the recognized behavioral elements allowed us to evaluate the mother–infant contacts and to analyze
their dynamics during calf growth. The most common calf positions relative to the mother during this period
were “at the cow’s tail” and “at the cow’s side.” The importance of behavioral responses of calf for the devel-
opment of social behavior in adult animals is emphasized.
DOI: 10.1134/S1062359006010079

Beluga is one of the most typical representatives of
Arctic marine mammals. The initial interest to beluga
was provoked by its large-scale fishery. As a result, fish-
ery aspects of their distribution, abundance, and migra-
tion were covered in numerous publications in 1950s–
1970s. As beluga fishery was discontinued, the interest
to its research diminished. Recent decade revived the
interest to beluga as a unique mammalian species with
behavioral and morphological adaptations to life in the
Arctic Seas.
It is common knowledge that social behavior of
mammals is formed from innate and acquired behav-
ioral patterns, while specific development of behavioral
responses during early development determine social
behavior of adult animals (Menning, 1982). Beluga
calves remain with the mother for three or more years.
A long period of nursing and great mother–infant
attachment are important factors of calf survival and its
timely adaptation to aquatic environment. The mother–
infant relations maintained for several years allow them
to form communities of cows and juveniles. Males at
the age of 4–5 years leave the maternal school and
come back only during the breeding season. Cows keep
to the maternal school during their whole life (Kleinen-
berg et al., 1964; Bel’kovich and Yablokov, 1965;
Yablokov et al., 1972). Hence, studies of parental
behavior, particularly, the mother–infant spatial inter-
actions and their quantitative and qualitative descrip-
tion are important to understand the role of intraspecific
contacts determining their population ecology and
social structure.
The mother–infant relations in dolphins are a com-
pound complex of behavioral elements, the nature and
biological significance of which are subject of different
opinions and speculations. This is primarily due to the
difficulties of material collection, observation tech-
niques, and data processing. Nearly all research on the
subject was carried out in dolphinaria (Bondarchuk
et al., 1976; Bliznyuk and Dzhincharadze, 1978; Voro-
nin et al., 1978; Tomilin and Bliznyuk, 1979; Kara-
bashyan et al., 1982; Reid et al., 1995; Gubbins et al.,
1999; Chechina, 2004). Studies under natural condi-
tions are usually limited to bottlenose dolphin (Tursi-
ops truncatus) (Richards, 1966; Mann and Smuts,
1998, 1999).
The goal of this work was to study the mother–
infant relations during the first months of calf life and
to trace its dynamics.
MATERIALS AND METHODS
Observations were carried out in the White Sea near
the Beluzhii Cape of the Solovetskii Island in the breed-
ing ground of belugas in June–August 2001 from an
11 m wooden tower at the littoral boundary. This breed-
ing ground is uniquely close to the observation point,
which allowed detailed monitoring of the mother–
infant relations. The studied water area was divided
into four zones with stable animal concentration
(Bel’kovich et al., 2002).
Cows with calves of different age were monitored.
Three age groups of calves were recognized by the size,
color, other appearance distinctions, breath-holding
time, and their combination (Table 1). These characters
insignificantly varied within the norm.
The attachment to mother was confirmed by the calf
position relative to mother at different developmental
stages and by the pattern of their locomotor activity.

53
54 KRASNOVA et al.

Table 1. Description of beluga calves of different age

Size in proportion
Breath-holding
Age group Color of mother’s body Appearance
time, s
size
Newborn Brown 1/4 5–7 transverse skin folds, big head (commonly 5–6
dark), not completely stretched pectoral blades
One-month-old Black 1/3 Smooth skin, occasional side folds Up to 120–180 in a series
of short (8–9 s) pauses
Two-month-old Dark gray 1/2 Smooth skin with no folds Same as above

The following calf positions were recognized: (1) “near mother and usually keeps at the cow’s side or tail. No
the cow,” at a distance of no more than 1.5 m from it; observations were carried out under unfavorable
(2) “at a distance from the cow,” from 1.5 to 5 m pro- weather conditions (rain, strong wind, or fog). Under-
viding there was only one calf. Cases when a calf was water recordings from a video camera installed in the
more than 5 m away from the mother and swimming site most visited by belugas were also used in the anal-
with other school numbers were considered as a “group ysis.
behavior.” In total, 11 elements of calf behavior were Within 177 h observations, 1959 behavioral
recognized (Fig. 1). responses were recorded including 581, 475, and 903
We used the methods of total record-keeping, obser- records for newborns, one-, and two-month-old calves,
vation of individual animals and animal groups (focal respectively.
observations), and recording of individual behavioral
responses (Popov and Il’chenko, 1990). Animals were RESULTS AND DISCUSSION
identified by clearly visible long-term markers (Cher-
netsky and Krasnova, 2001). The markers of cows When observation started, the school included dark
allowed us to recognize their calves in a school and to gray yearlings (half size of adult), apparently, born in
monitor their development. The duration and frequency April or May. The first newborn was recorded on July
of behavioral elements as well as proportions between 10; next two were born on July 16; the total number of
them were recorded for each calf age group. During newborns was eight. The last newborn appeared on
total record keeping and focal observation, the behavior August 14. This confirms an extended period of birth in
of the first, second, third, etc. calves of identified cows belugas proposed previously (Kleinenberg et al.,
were recorded. Behavioral responses of calves were 1964). The total number of yearlings in the Solovetskii
only recorded in the absence of stress and anthropo- breeding ground of belugas was around 30.
genic impacts. In danger, belugas demonstrate the fol- Development of motor activity in newborns. Starting
lowing behavioral strategy: a calf is covered by the from birth, newborns permanently follow their mother,
which is one of the most important adaptations—the
behavior of the mother following (Tomilin and
Table 2. Frequency of behavioral elements in each age group Bliznyuk, 1979). The behavior of a mother with a new-
of calves, %
born features swimming close to the water surface,
Calf groups which facilitates the calf’s respiration. The calf move-
Behavioral element
ments are poorly coordinated yet and all behavioral
one- two-month- responses are short-term. The movements are hasty,
newborn
month-old old wasteful, and imperfect. The position relative to the
At the cow’s side 44.8 24 26 mother often changes. In the first days after birth, a calf
is most commonly at the cow’s side or tail (Table 2).
On the cow’s back 10 8 14 Apparently, this position is the most beneficial for the
On the cow’s tail 3 3 5 calf and allows it to overcome water resistance. The
Ahead of the cow 4 7 8 longest stable periods of newborns are at the cow’s side
Above the cow 9 3 3 (2 to 35 s). Movements requiring high coordination and
strength are avoided or very short-term (1 to 4 s). These
Below the cow 0.3 1 2 include riding on the cow’s back or tail favorite among
Behind the cow 0.9 2 3 older calves. One- and two-day-old calves could neither
At a distance from the cow 8 12 6 climb on the mother’s back nor stay on her caudal
At the cow’s belly 1 3 4 peduncle, particularly, in movement. At the same time,
short-term attempts to climb on the back (2 to 13 s) and
At the cow’s tail 17 34 22 tail (1 s) were recorded. The behavioral repertoire of
At the cow’s head 2 3 7 newborns includes nearly no elements when a calf is

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 MOTHER–INFANT SPATIAL RELATIONS IN WILD BELUGA 55

At the cow’s side On the cow’s back On the cow’s tail

Ahead of the cow Above the cow

Below the cow Behind the cow

At a distance from the cow At the cow’s belly

At the cow’s tail At the cow’s head

Fig. 1. Schematic behavioral elements of beluga calf in spatial interactions with the cow.

inaccessible or uncontrollable by the cow, such as
behavioral patterns “behind the cow” or “away from the
cow.”
The data on the duration and frequency of feeding in
beluga available to date were obtained in oceanaria
(Russell et al., 1997). We have obtained original data
on beluga calf feeding using underwater video camera.
Feeding is preceded by the food-begging behavior of a
calf manifested as tight physical contacts. A calf leans
upon the mother’s tail and rubs against her sides and
back. Such behavior can last up to 59 s, after which they
dive. Feeding always occurs underwater; a calf is at the
mother’s belly at an angle of 90 or 45°. The duration of
single feeding his 4–6 s (behavioral pattern “at the
cow’s belly”).
On day 4, mother allows a calf to swim at a distance
of 2–3 m but only under safe conditions, e.g., surrounded
by other cows and calves and in the absence of males and
active gray juveniles. On day 7, calves reliably stay on
the mother’s back (up to 12 s) and tail (up to 17 s).
The behavior of calves becomes more complex as
they physically grow and develop; they become more
independent. Nevertheless, the positions “at the cow’s
side” and “at the cow’s tail” still prevail. The proportion
of these patterns in the total duration of recognized

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56 KRASNOVA et al.

% behavioral elements was 47.46 and 20.77%, respec-

50 47.46 tively (Fig. 2a).
(a)
40 On day 10, newborns demonstrated swimming on
the back for over 10 s away from the cow. On day 20–
30 24, they could form short-term (7 to 32 s) groups with
20.77 other yearlings. They preferred to form groups of two
20 (53%) usually with one-month-old calves. Groups of
11.54
10 6.35
four or three occurred less frequently (27 and 20%,
5.06
2.25 2.99
0.63 0.76 1.3 0.89 respectively).
0 In breeding grounds of belugas, the periods of
At the cow’s side
On the cow’s back
On the cow’s tail
Ahead of the cow
Above the cow
Below the cow
Behind the cow
At a distance from the cow
At the cow’s belly
At the cow’s tail
At the cow’s head
breeding and birth can overlap in time. When excited
males swim to the school, cows with newborns usually
keep apart from the main group. Otherwise, the mother
covers a calf from a close male or calf by itself hides
behind the cow from an approaching male. An inverse
behavior when a cow with a newborn actively swims to
a group of males and females ready for mating is
% extremely rare. In single instances, excited animals
50 (b) drove a cow with calves from the main group. For
40 37.4 instance, on August 6 a cow with a calf joined a group
of excited females and males (9 individuals); after a few
30 25.62 seconds one beluga (of unknown sex) quickly swam up
and drove them away from the group. Such behavior of
20 13.78 a cow with a newborn has no explanation yet.
10 7.25
4.73 The pigmentation of newborns changed 15–20 days
3.04 1.79 2.13 2.66
0
1.09 0.51 after birth. Calves became black from brown.
At the cow’s side
On the cow’s back
On the cow’s tail
Ahead of the cow
Above the cow
Below the cow
Behind the cow
At a distance from the cow
At the cow’s belly
At the cow’s tail
At the cow’s head

Motor activity of one-month-old calves. At the age

of one month (1/3 size of adult), calves demonstrate a
more diverse behavioral repertoire. They can swim on
their side or back for long periods, jump out of water
exposing the body, rotate, etc. Nevertheless, the most
common behavioral elements of calves of this age
include positions “at the cow’s tail” (34%) and “at the
cow’s side” (24%) (Table 2). Contacts with other mem-
bers of the group become more frequent. At the same
% time, the frequency of behavioral element “at a distance
50 (c) from the cow” increased by as low as 4%. More inde-
pendent calves more frequently occurred behind (2%
40 vs. 0.9% in newborns) or ahead of the cow (7 vs. 4% in
30.89 30.27
30 newborns). These behavioral elements in this age group
increased not only in frequency but also in duration
20 (Fig. 2b).
10 9.03
5.54
8.16 We failed to observe beluga-specific kindergartens
2.94 2.5 2.38 2.65 2.64 3 when calves form a separate group overseen by young
0 animals, while adults hunt (Bel’kovich and Shcheko-
At the cow’s side
On the cow’s back
On the cow’s tail
Ahead of the cow
Above the cow
Below the cow
Behind the cow
At a distance from the cow
At the cow’s belly
At the cow’s tail
At the cow’s head

tov, 1990). This can be attributed to the absence of

searching and hunting behavior in the Beluzhii Cape
belugas since they swim away to the sea for feeding.
For rest, two to five cows group with calves and quies-
cently lie on the water surface or move slowly using
their fins. Calves demonstrate different behavior during
this period: usually they keep to their mothers and swim
around her, lean against her body, or group in two or
three without going far from the cows. We observed
independent movement of calves from one cow to
Fig. 2. Proportions of time spent for each behavioral element
another (e.g., one-month-old, July 1; newborn, July 17)
by newborn (a), one- (b), and two-month-old calves (c); and active attempts to communicate with them. They
abscissa: calf position; ordinate: time proportion, %. try to lie on the caudal peduncle and to climb on the

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 MOTHER–INFANT SPATIAL RELATIONS IN WILD BELUGA
back of cows. The cows patiently accepted such behav-
ior.
Motor activity of two-month-old calves. The most
frequent behavioral elements of one-month-old calves
were observed at a similar or slightly lower frequency
in two-month-old ones (Table 2). They need no more
mother’s help for swimming. The behavioral element
“at the cow’s side,” which was extremely important for
newborns, is notably less common in two-month-old
calves (Fig. 2c). They become more independent at this
age. In this context, the behavior with limited mother’s
care becomes more frequent. For instance, swimming
ahead of the cow occurs by 4 and 1% more frequent
compared to newborns and one-month-old calves. Two-
month-old calves appear behind the mother by 2.1 and
1% more frequently, respectively. The duration of their
stay away from the mother increases. For instance, on
July 7 we observed a two-month-old calf swimming far
from the mother for 45 min. The breath-holding time
increased as calves developed (Table 1), which can
explain the increasing frequency and duration of their
stay under the cow (0.3, 1, and 2% in newborn, one-,
and two-month-old calves, respectively). Despite their
independence, yearlings spend a lot of time on the
mother’s back (9.03%) and tail (2.94%). These posi-
tions can represent behavioral elements of rest or game.
Sometimes two-month-old calves “rode” on the back of
gray juvenile belugas (July 14).
It is of interest that the frequency of behavioral ele-
ment “at the cow’s head” increased with calf age. In this
case, a calf at the mother’s head leans against it or
jumps out of water nearby as though attracting her
attention. For instance, the frequency of this element
was 2, 3, and 7% in newborn, one-, and two-month-old
calves, respectively. In dolphins, physical contacts in
the form of touching or rubbing are typical manifesta-
tions of social activity. The head region is preferred for
such contacts (Mann and Smuts, 1999). During hierar-
chical behavior, dolphins use force interactions; e.g.,
they collide at high speed trying to submerge a compet-
itor, try to toss up a competitor, etc. (Pryor, 1975). In
addition, belugas demonstrate “butting” hierarchical
behavior when animals collide with their heads trying
to push away a competitor. Social activity increases
with calf age; it more imitates the adult behavior and
the frequency of this behavioral element accordingly
increases.
As previously, calves spend most of their time near
mothers; however, at the sight of other yearlings a calf
usually joins them. For instance, on July 17 a cow with
a calf lay quiescently aside from other animals for
40 min. When two swimming yearlings approached,
the calf joined them, while the mother remained where
she was.
Note that all described general trends in calf behav-
ior do not exclude pronounced individual distinctions
in both the frequency and duration of particular behav-
ioral elements. According to our data, newborns could
BIOLOGY BULLETIN Vol. 33 No. 1 2006
57
swim at a distance from the cow on average for 12.67 s.
At the same time, we observed a 4-day-old calf swim-
ming at a distance of 4 m from the cow for 28 s; on July
20, a 10-day-old calf was observed separate from the
cow for 48 s. The mean duration of the one-month-old
calf position “at the cow’s tail” was 13.29 s; however, a
calf of this age retained this position for 70 s on July 8.
Apparently, such individual peculiarities depend on the
calf physical state as well as on the behavior of the
mother and the whole animal group, which requires
special investigation.
The unique breeding ground of beluga in the White
Sea near the Beluzhii Cape of the Solovetskii Island
made possible the first investigation of their parental
behavior, particularly, the spatial mother–infant rela-
tions under natural environmental conditions. The
revealed behavioral patterns of newborn belugas under
natural conditions such as high motor activity and
short-term behavioral elements conflict the observa-
tions in captivity. For instance, Schneider et al. (2003)
observed low motor activity of two newborn belugas
within the first week in a Canadian oceanarium: they
were quiescent for up to 80% of total observation time.
Clearly, this was directly due to their living conditions
or physical state. High motor activity in the sea can be
required in strong current conditions, while the absence
of current in an oceanarium makes calf locomotion
unnecessary.
The obtained data demonstrate a decreasing dura-
tion of calf staying with the mother as it grows, which
is typical for the general behavioral strategy of all
mammals. This corresponds to the increasing index of
behavioral element “at a distance from the cow” and to
the decreasing behavioral element “at the cow’s side”
as calf grows (Fig. 1) as well as to the formation of their
short- and long-term game groups. While playing with
other animals of the school, calves learn to recognize
their responses and environmental stimuli, which allow
them to timely notice and respond to danger or changed
behavior of a neighbor. Calves contacting with each
other acquire and share game and orientation skills
(Baskin, 1976; Voronin et al., 1978; Karabashyan
et al., 1982).
All age groups of calves demonstrated the predomi-
nance of two behavioral elements in the interaction
with the mother: “at the mother’s side” and “at the
mother’s tail.” These elements seem to have an adaptive
significance during the most critical postnatal period of
their development, promote swimming, and favor their
survival. The decreasing duration of these behavioral
elements in two-month-old calves can reflect a
decreased capacity of a cow to “carry” a calf in her flow
field due to its growth and independence. The swim-
ming skills of calves improve at the same time (Cock-
roft and Ross, 1990).
We believe that the data obtained shed new light to
the spatial mother–infant relations in belugas under nat-
ural environmental conditions.
58 KRASNOVA et al.
ACKNOWLEDGMENTS
This work was supported by the International Fund
for Animal Welfare (IFAW).
REFERENCES
Baskin, L.M., Povedenie kopytnykh zhivotnykh (Animal
Behavior), Moscow: Nauka, 1976.
Bel’kovich, V.M. and Shchekotov, M.N., Belukha. Povede-
nie i bioakustika v prirode (Beluga. Natural Behavior and
Bioacoustics), Moscow: In-t okeanologii im. P.P. Shirshova,
1990.
Bel’kovich, V.M. and Yablokov, A.V., The Structure of
School of Toothed Whales (Odontoceti), in Morskie mlekopi-
tayushchie (Marine Mammals), Moscow: Nauka, 1965,
pp. 65–69.
Bel’kovich, V.M., Chernetskii, A.D., and Kirillova, O.I., The
Biology of Beluga (Delphinapterus leucas) in the Southern
White Sea, in Morskie mlekopitayushchie. Rezul’taty issledo-
vanii, provedennykh v 1995–1998 gg (Marine Mammals.
Results of Studies in 1995–1998), Moscow: Nauka, 2002,
pp. 53–77.
Bliznyuk, Ya.I. and Dzhincharadze, K.A., Razvitie koordi-
natsii dvizhenii v rannem ontogeneze, in Mater. 7 Vsesoyuzn.
soveshch. “Morskie mlekopitayushchie”. Simferopol’, 20–
23 sentyabrya 1978 (Proc. 7 All-Union Conf. “Marine Mam-
mals.” Simferopol, September 20–23, 1978), pp. 41–42.
Bondarchuk, L.S., Matisheva, S.K., and Skibnevskii, R.N.,
Development of Behavior in Calves of Bottlenose Dolphin
(Tursiops truncatus) in the Black Sea, Zool. Zh., 1976,
vol. 55, no. 2, pp. 276–281.
Chechina, O.N., Behavioral Patterns in the Mother–Infant
Relations in Bottlenose Dolphin (Tursiops truncates ponti-
cus Varabassh, 1940) under Dolphinarium Conditions, Tr. III
Mezhdunar. konf. “Morskie mlekopitayushchie Golarktiki”
(Proc. III Int. Conf. “Holarctic Marine Mammals”), Moscow,
2004, pp. 579–581.
Chernetsky, A.D. and Krasnova, V.V., Identification of White
Whales Using Natural Markers, 14-th Biennial conference
on the Biology of Marine Mammals. Vancouver, Canada, 28
November–3 December, 2001, p. 42.
Cockroft V.G., Ross G.J.B. Observations on the Early Devel-
opment of a Captive Bottlenose Dolphin Calf, The Bottlenose
Dolphin, Leatherwood, S., Reeves, R.R., Eds., New York:
Academic, 1990, pp. 461–478.
Gubbins, C., McCowan, B., Lynn, S., and Reiss, D., Mother-
infant Spatial Relations in Captive Bottlenose Dolphins, Tur-
siops truncatus, Marine Mamm. Sci., 1999, no. 3, pp. 751–
765.
Karabashyan, V.V., Gedzhadze, G.Sh., and Suglobov, S.I.,
Relation of Infants with the Mother and Other Dolphins in
Captivity, in Izuchenie, Okhrana I Ratsional’noe Ispol’zova-
nie Morskikh Mlekopitayushchikh. Mater. 8. Vsesoyuzn.
soveshch., Astrakhan’, 5–8 oktyabrya (Studies and Conser-
vation of Marine Mammals. Proc. 8 All-Union Conf. Astra-
khan, October 5–8) 1982, pp. 154–155.
Kleinenberg, S.E., Yablokov, A.V., Bel’kovich, V.M., and
Tarasevich, M.N., Belukha. Opyt monograficheskogo issle-
dovaniya vida (Beluga. Targeted Investigation of the Spe-
cies), Moscow: Nauka, 1964.
Mann, J. and Smuts, B.B., Natal Attraction: Allomaternal
Care and Mother-Infant Separations in Wild Bottlenose Dol-
phins, Animal Behaviour, 1998, vol. 55, no. 5, pp. 1097–
1113.
Mann, J. and Smuts, B.B., Behavioral Development in Wild
Bottlenose Dolphin Newborns (Tursiops sp.), Behaviour,
1999, vol. 136, no. 5, pp. 529–566.
Manning, A., An Introduction to Animal Behavior, Massa-
chusetts: Addison-Wesley, 1979. Translated under the title
Povedenie zhivotnykh. Vvodnyi kurs, Moscow: Mir, 1982.
Popov, S.V. and Il’chenko, O.G., Metodicheskie rekomen-
datsii po etologicheskim nablyudeniyam za mlekopitayu-
shchimi v nevole (Guidelines to Ethological Observations of
Mammals in Captivity), Moscow: Mosk. zool. park, 1990.
Prior, K., Lads before the Wind, Washington: Sunshine
Books, North Bend, 1975.
Reid, K., Mann, J., Weiner, J.R., and Hecker, N., Infant
Development in Two Aquarium Bottlenose Dolphins, Zoo
Biol., 1995, no. 14, pp. 135–147.
Richards, A.F., Bottlenose Dolphin Female Relationships
and Reproductive Parameters in Shark Bay, Australia: PhD
Dissertation, University of Michigan, 1996, p. 183.
Russell, J.M., Simonoff, J.S., and Nightingale, J., Nursing
Behaviors of Beluga Calves (Delphinapterus leucas) Born in
Captivity, Zoo Biology, 1997, vol. 16, pp. 247–262.
Schneider, L., Schamel, L., and Noonan, M., Behavioural
Landmarks in the Development of Neonatal Beluga Whales
(Delphinapterus leucas), in 17-th Conference of the European
Cetacean Society. Las Palmas de Gran Canaria, 9–13 March
2003, pp. 131–132.
Tomilin, A.G. and Bliznyuk, Ya.I., Behavioral Patterns of
Calves and Cows of Bottlenose Dolphin during the Repro-
ductive Period. The Following Behavior in Cetaceans, Byul.
Mosk. O-va Ispytatelei Prirody. Otd. Biol., 1979, vol. 84,
no. 5, pp. 35–39.
Voronin, L.G., Vannikova, N.R., Vronskaya, S.D., and
Kozarovitskii, L.B., Certain Properties of Early Ontogenesis
of Behavior in the Black Sea Dolphin, Mater. 7 Vsesoyuzn.
soveshch. “Morskie mlekopitayushchie”. Simferopol’, 20–
23 sentyabrya 1978 (Proc. 7 All-Union Conf. “Marine Mam-
mals.” Simferopol, September 20–23, 1978), p. 23.
Yablokov, A.V., Bel’kovich, V.M., and Borisov, V.I., Kity i
del’finy (Whales and Dolphins), Moscow: Nauka, 1972.
BIOLOGY BULLETIN Vol. 33 No. 1 2006