Mammals of the Soviet Union

Part 3

This is the fourth book of Mammals of
the Soviet Union, representing the third
part of the second volume; it is devoted
to descriptions of the orders of Soviet
aquatic mammals — pinnipeds
(Pinnipedia) and, in part, cetaceans
(Cetacea), toothed whales (Odontoceti).
In the sequence of descriptions from the
“higher” to the “lower” orders adopted in
this series, pinnipeds should have pre-
ceded carnivores, 1.е., should have
appeared in the second book. The group-
ing of the orders at a higher level is
given after G.G. Simpson (1945). The
sequence of genera and species within
the orders has been retained as before,
1.е., from the less specialized to the more
specialized.

The order of pinnipeds or seals
(Pinnipedia) is described in this volume.
The cohort of ungulates and carnivores
represented in Soviet fauna by orders of
artiodactyls, and perissodactyls, sirenians,
carnivores, and pinnipeds thus comes to
an end and the cohort of whales (Mutica)
commences. The toothed whales are
described in this volume.

While it has not always been pos-
sible to maintain a totally uniform
description of the genera and species as
in the volumes already published, in spite
of every effort to do so, the sequence
has been adhered to, with some excep-
tions, in the case of Pinnipedia. But, it
was impossible to use sane format in
describing the toothed whales
(Odontoceti). Only a brief morphological
description has been given for many
species, which is more or less adequate
for identifying the species. General
information on their distribution and
fragmentary biological data are also
given. In some cases the total absence of
such information is indicated.

fy
ef

hel
и

Mammals of the Soviet Union

Volume II, Part 3

MAMMALS OF THE SOVIET UNION

In Three Volumes

Edited by
Late V.G. Heptner and N.P. Naumov

Vysshaya Shkola Publishers
Moscow, 1976

Mammals of the
Soviet Union

VOLUME о Part 3

—_

PINNIPEDS AND TOOTHED WHALES
Pinnipedia and Odontoceti

Late V.G. Heptner, К.К. Chapskii, V.A. Arsen’ev
_and У.Е. Sokolov

Edited by
Late V.G. Heptner

Scientific Editor
JAMES G. MEAD

Smithsonian Institution Libraries
and
The National Science Foundation
Washington, D.C.
1996

Smin-B86 SIL 005

Mlekopitayushchie Sovetskogo Soyuza
In Three Volumes
V.G. Heptner and N.P. Naumov, editors

Vysshaya Shkola Publishers
Moscow, 1976

Translator: P.M. Rao
General Editor: Dr. V.S. Kothekar

©1996 Oxonian Press Pvt. Ltd., New Delhi

Library of Congress Cataloging-in-Publication Data
(Revised for v. 2, pt. 3)

Geptner, V.G. (Vladimir Georgievich), 1901 - 1975.
Mammals of the Soviet Union.

Translation of: Mlekopitaiushchie Sovetskogo Soiuza.
Vol. by V.G. Heptner and A.A. Sludskii; illustrators,
A.N. Komarov and N.N. Kondakov.

Vol. 2, pt. 2 includes index.

Includes bibliographical references.

Supt. of Docs. no.: $1 1.2:Ar7/2

Contents: v. 1. Artiodactyla and Perissodactyla—
у. 2 pt. 2 Carnivora (hyaenas and cats)—pt. 3.
Pinnipeds and toothed whales (Pinnipedia and Odontoceti)

1. Mammals—Soviet Union—Collected works.

I. Nasimovich, A. A. П. Bannikov, Andrei Grigor’evich.
Ш. Hoffmann, Robert $. IV. Sludskii, А. A. У. Title.
QL728.S65G4713 1988 599.0947 85-600144

Translated and published for the Smithsonian Institution Libraries, pursuant
to an agreement with the National Science Foundation, Washington, D.C.,

by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001

Printed in India at Baba Barkha Nath Printers, New Delhi

UDC 596.5

We gratefully acknowledge the assistance rendered by the follow-
ing persons, which made the publication of this monograph possi-
ble: V.D. Pastukhov (Listvenichnoe, Baikal), late P.G. Nikulin and
D.I. Chugunkov (Petropavlovsk-Kamchatskii), I.F. Golovlev, V.A. Zem-
skii, late B.A. Zenkovich, M.V. Ivashin, L.A. Popov, and S.K. Klumov
(Moscow), S.L. Delyamure (Simferopol’), A.S. Skryabin, V.V. Treshchev,
and M.V. Yurakhno (and other colleagues at the Helminthological Lab-
oratory, Crimean State University), V.D. Kokhanov (Kandalaksha), late
В.1. Badamshin (Fort Shevchenko), F.Sh. Khuzin, М.Уа. Yakovenko
(Murmansk), А.С. Beloborodov, Уи.Г. Nazarenko, and V.A. Potelov
(Arkhangel’sk), A.A. Berzin, G.M. Kosygin, Yu.V. Kurochkin, and late
Е.А. Tikhomirov (Vladivostok), А.Р. Shustov and С.А. Fedoseev (Maga-
dan), and many others cited at the appropriate places in the text. Further,
S.V. Marakov (Kirov), V.I. Krylov (Moscow), and some others made
available to us their original photographs; due acknowledgments have
been given under the photographs.

We express our appreciation to all these persons, and to all
those others without whose assistance this publication would not have
been possible. The authors are extremely grateful to IP. Mitina and
E.V. Zubchaninova of the Department of Vertebrate Zoology, Moscow
University, who put in much effort toward the preparation of this
manuscript for the press and its publication; and to V.I. Korotkova,
Chief Librarian of the Zoological Museum, Moscow University, who
readily supplied to the authors rare and almost inaccessible reference
materials. The authors are particularly grateful to Prof. Г.1. Barabash-
Nikiforov (Voronezh) and S.V. Marakov (Kirov) who undertook the
formidable task of reading through the manuscript, and for their
extremely valuable suggestions. The authors also express their gratitude
to O.L. Rossolimo, Director, Zoological Museum, Moscow University.
Much of the preparatory work for this publication was completed in this
museum.

Most of the reference materials utilized were published before the
end of the 1960s, although a few books and articles from more recent
years have been useful. References are cited in the-text by the author’s
name without initials and with the year of publication. Initials are used
only when the surnames are identical. References to authors of unpub-
lished private communications are also made in parentheses but with
initials and without the year. The bibliography at the end of the book
covers only the references cited (although many more were reviewed),

У1

excluding those listed under synonyms. As in the previous publications,
the authors have included, quite naturally, their own unpublished mate-
rial in the text.

The preceding volumes of Mammals of the Soviet Union have been
warmly received within and outside the USSR. The first three books in
the series were translated into German. The authors hope that this vol-
ume, too, will be as warmly received although they are aware of its several
shortcomings. An explanation for this, although not a justification, is the
complicated nature of the material, especially that of whales. Save for
some individual species, the museums in the USSR contain practically
no collections of this group.

V.G. Heptner

7 |
Fe А | | |
<
2 — J}
gf и ‘ y=

FOREWORD TO THE ENGLISH
EDITION

The Smithsonian Institution Libraries, in cooperation with the National
Science Foundation, has sponsored the translation into English of this and
hundreds of other scientific and scholarly studies since 1960. The program,
funded with Special Foreign Currency under the provisions of Public Law
480, represents an investment in the dissemination of knowledge to which
the Smithsonian Institution is dedicated.

One of the values of this translated volume is to give English readers
an insight into another philosophical system that devoted nearly a cen-
tury to the studies of marine mammals. This work is important because
it has an abundance of data that were taken from commercial harvest,
particularly from small cetaceans, that have been generally unavailable
to the English-speaking community. A monograph of this magnitude,
with such an extensive bibliography, serves as an excellent entré into the
Russian literature.

This volume is the last to be published under the general editor-
ship of V.G. Heptner and N.P. Naumov; Volume II, part 3, Pinnipeds
and Toothed-whales appeared in 1976, the year after Professor Hept-
ner’s death. Only one more volume of the original series remains to be
published in an English edition, Volume II, Part 1, Sea Cows and Carni-
vores (Dogs, Bears, and Mustelids), but editing of the English language
manuscript is about 60 percent complete, and it is the intention of the
Smithsonian Libraries Translation Program to publish the volume as
soon as possible.

After a lapse of two decades, the series has been revived with the
sponsorship of the Russian Academy of Sciences’ Institute of Evolu-
tionary Animal Morphology and Ecology in Moscow, with a new series
title reflecting recent political changes—Mammals of Russia and Border-
ing Regions. The first volume in this new series is, as had been antici-
pated, Usatye Kity [Baleen Whales], by V.E. Sokolov and V.A. Arsen’ev
(1994, Nauka, Moscow, 208 pp.). A second volume on Zaitseobraznye
[Lagomorphs] is in production. It is hoped (but by no means certain)
that English editions of the revived series can be made available.

Vill

English readers interested in Dr. Heptner’s contributions as a mam-
malogist should refer to the Foreword to the English Edition of Volume:
I. Conventions used in rendering geographic names, first stated there,
are reprinted here for the convenience of the reader. Geographic names
are generally transliterated directly, but a few exceptions were permitted
(e.g. Moscow instead of Moskva, translation rather than transliteration
of certain modifiers of place names, such as Northern, rather than Sever-
naya Dvina). Soviet administrative units are numerous, and the following
equivalents were employed in translation: Krai, territory; oblast’, district;
raion, region; guberniya (archaic), province. Also, in the original Russian
text, rivers, mountain ranges, and cities are often not explicitly identified,
the Soviet reader being presumed sufficiently familiar with the geography
of the country to be able to understand from the context of the sentence
what sort of place is referred to. Complicating the matter is the lack
of articles as parts of speech in Russian. To assist the English reader,
the following conventions have been adopted: if a river is referred to,
an article precedes it; if a mountain range is referred to, it is translated
as a plural; if a city is referred to, it is singular, and lacks the article.
Examples are: the Ural (river); the Urals (mountains); Ural’sk (city).
Geographic place names are also inflected in Russian, and these have
been simplified by omitting transliteration of the inflected ending. For
example, the Russian phrase у Yaroslavskoi 1 Kostromskoi oblastyakh is
translated “in the Yaroslavl’ and Kostroma districts.” In cases where the
nominative form of the place name has an -sk ending this is, however,
transliterated (e.g., Omsk); when a Russian “soft sign” is employed in a
place name, this is transliterated as an apostrophe (e.g., Khar’kov).

As in any translation, particularly from one orthography into
another, there arise problems of standardization of names. For
geographic names the National Geographic Atlas of the World, 4th edition,
1975 was used for this volume (principally the map on pages 122 - 123).
Some of the geographic terms like gulf, bay, inlet, strait [zaliv, bukhta,
guba, proliv] are used in slightly different contexts in the original Russian
text. Because of the large number of place names in this volume, it was
not possible to verify all of them, and some inconsistencies are likely to
occur. We would appreciate it if readers would bring any errors they may
notice to our attention.

The usage of the common terms “dolphin” and “porpoises” varies
geographically. In North America these terms are used interchangeably
in reference to members of the family Delphinidae (sensu Simpson 1945) |
but only the term “porpoise” is used to refer to members of the fam-
ily Phocoenidae (sensu Simpson 1945). In most other English speaking

ix

countries, including England, the term “dolphin” is restricted to refer-
ence to the family Delphinidae. In Russian, the term “del’fin” (dolphin)
is used to refer to all of the members of the family Delphinidae and the
term “morskaya svinya” (Sea pig), which is equivalent to the meaning
of the Latin roots of “porpoise” (porcus = pig + piscis = fish) is used
to refer to the members of the family Phocoenidae. We have therefore
chosen to use the more restrictive English usage in this volume.

A few of the vernacular names, those where the Russian name was
vastly different from the generally accepted English name, have been
changed, but translations of Russian vernacular names were employed
where there was not a generally accepted English term (e.g. the vernac-
ular name of the genus Lagenorhynchus). There is no English vernacular
name for members of the genus Mesoplodon; Mead prefers to trans-
late the Russian term “remnezubov” (sword-toothed) as Mesoplodon,
rather than a literal translation. In addition, recent systematic studies
have shown that the appropriate scientific name for the Malay dolphin
(Stenella dubia) is Stenella attenuata (= Pan-tropical spotted dolphin)
and the English common name of the Bridled dolphin (Stenella frontalis)
is Atlantic spotted dolphin.

Some terms placed in brackets [ ] indicate that they are additions
of the scientific editor and are not found in the Russian text. In statisti-
cal references, x replaces М (mean), a much more accepted symbol for
“mean” in English works.

One further point of confusion not apparent when Volume I was
translated also requires clarification; that is the English transliteration
of the senior author’s surname. This begins with the fourth letter of the
Cyrillic alphabet, which usually has а “С” sound in Russian. However,
the surname was originally German, and in the original German began
with the letter “Н” of the Latin alphabet. Since Cyrillic has no equivalent
of “H” this is usually transliterated шо “С” in Russian. However, Hoff-
mann, from conversations learned that Dr. Heptner preferred to use the
original Germanic form of his surname rather than the transliterated ver-
sion, which is rendered as Geptner. The rules of transliteration employed
by the Library of Congress do not permit such flexibility, and the atten-
tive reader may notice that Library of Congress cataloging employs the
latter.

The Series Scientific Editor expresses particular thanks to the edi-
tor of this volume, Dr. James G. Mead, Curator of Marine Mammal
Project, Department of Vertebrate Zoology, National Museum of Nat-
ural History, Smithsonian Institution. Dr. Mead’s broad knowledge of
all aspects of marine mammal biology was critical to the quality of this
English edition, and I am grateful to him for the considerable time he has

xX

devoted to ensuring the accuracy of the volume. Thanks are also due to
P.M. Rao and Dr. V.S. Kothekar, Translator and General Editor respec-
tively of Amerind Publishing Company, New Delhi, India, for producing
the original translation under the provisions of U.S. Public Law 480.
Credit also must be given to the general editorial staff of Amerind Pub-
lishing Co., who confirmed all of the technical names of the prey species
and parasites, checked the bibliographic references and translated many
obscure Russian scientific terms.

Robert S. Hoffmann

Series Scientific Editor

Assistant Secretary for the Sciences
Smithsonian Institution, Washington, DC

James G. Mead

Volume Scientific Editor

Curator of Marine Mammals
National Museum of Natural History

FOREWORD

This is the fourth book of Mammals of the Soviet Union, represent-
ing the third part of the second volume; it is devoted to descriptions
of the orders of our [Soviet] aquatic mammals—pinnipeds (Pinnipedia)
and, in part, cetaceans (Cetacea), toothed whales (Odontoceti). In the
sequence of descriptions from the “higher” to the “lower” orders adopted
in this series, pinnipeds should have preceded carnivores, 1.е., should
have appeared in the second book. The microsystem of the class is given
below to elucidate the natural sequence and the relationships between
orders. This is a simple and presently more widely used system, but pin-
nipeds have been assigned the status of an order and not a suborder as
commonly accepted. The grouping of the orders at a higher level is given
after G.G. Simpson (1945). The sequence of genera and species within
the orders has been retained as before, i.e., from the less specialized to
the more specialized.

The order of pinnipeds or seals (Pinnipedia) is described in this vol-
ume. The cohort of ungulates and carnivores represented in our [Soviet]
fauna by orders of artiodactyls and perissodactyls, sirenians, carnivores,
and pinnipeds thus comes to an end and the cohort of whales (Mutica)
commences. The toothed whales are described in this volume and the
baleen whales will be taken up in the next volume.

While it has not always been possible to maintain a totally uniform
description of the genera and species as in the volumes already published,
in spite of every effort to do so, the sequence has been adhered to, with
some exceptions, in the case of Pinnipedia. But, it was impossible to use
the same format in describing the toothed whales (Odontoceti). Firstly,
the biology of nearly all the species, except the Black Sea species, is not
adequately known even in respect of principal features, and information
on their distribution is more scant. Only a brief morphological descrip-
tion has been given for many species, which is more or less adequate
for identifying the species. General information on their distribution
and fragmentary biological data are also given. In some cases the total
absence of such information is indicated. Secondly, the faunal compo-
sition of the Soviet Pacific waters is not yet clear. While the presence

ХИ

of some species already reported calls for confirmation, several species
known from the coastal waters of Japan have not been reported in our
waters. Some of them are found on the American coasts at the same lati-
tudes. Their presence in our waters even as “Stray” finds is highly proba-
ble; such probable species have been indicated, included in the keys, and
briefly described. In this regard the present volume has achieved its aim,
not only to describe the known species, but also to draw attention to
the unknown. Such species total 14 while the number of relatively well-
known species in the Soviet Pacific waters is quite large. While using the
keys and morphological characteristics of the Pacific dolphins, reported
here for the first time from our waters, their identification should be
checked with the available literature and museum specimens. Interest-
ing finds could be anticipated in the Soviet Far East and each case of
a “stray” report or capture should be studied carefully. The probable
species and genera have been given without numbers and in small print.

The ranges of distribution of every species have been compiled on
the same principle as in the preceding books. However, in view of the
specificity of the biology of marine species (long migrations), in many
cases they are given in a more general form. The range extending beyond
our waters is mainly based on the works of Van den Brink (1958), Schef-
fer (1958), Hall and Kelson (1959), King (1964), Hershkovitz (1966),
Siivonen (1967), and some others, mostly of Japanese authors. The lat-
ter have been cited in the text. The entire synonymy has been selected
on the principles adopted earlier (see Foreword to Vol. I and Vol. II,
parts 1 and 2). In view of the scant indigenous literature on whales, their
synonymy is given very briefly. A more complete list of the synonyms of
the species of this order can be found in the monograph by Hershkovitz
(1966).

The total number of species constituting this class is roughly 3,500,
of which about 300 are represented in our fauna (Heptner, 1956). Almost
all the illustrations in this book are original with the exception of those
of the toothed whales taken from Hall and Kelson (1959). The original
drawings are by the well-known Russian artist N.N. Kondakov who, apart
from being a talented artist, is an experienced zoologist. Other sources
of the drawings have been acknowledged at the appropriate place.

The contribution of each author to this volume is as follows.
K.K. Chapskii wrote the general outline of the order Pinnipedia and
the family of seals (Phocidae), gave the keys to the families of the order
and species of the family of seals, and wrote the text on all the species of
this family except for the Baikal seal, Phoca sibirica, and the ribbon seal,
Phoca fasciata. V.A. Arsen’ev wrote the characteristics of the walruses
(Odobenidae) and the eared seals (Otariidae), and the entire description

хШ

of walruses, sea lions, fur seals, and the ribbon and Baikal seals. He also
wrote the sections on the distribution, biology, and economic importance
of all the species of toothed whales (Odontoceti), the characteristics of
the suborder of baleen whales (Mysticeti), and the outlines of all the
species of this suborder which, as mentioned above, will be included in
the next volume. V.E. Sokolov wrote the general outline of the order
of cetaceans, suborder of toothed whales, and the description of all
the genera of this suborder and the morphological characteristics of
its species. V.G. Heptner wrote the introduction to the book, keys to
the species of eared seals (Otariidae), and an outline of the genera and
species of sea lions (Zalophus californianus). Moreover, he worked out
the entire synonymy and participated in the compilation of notes on the
distribution of many species. As in the preceding books, the individual
authorship has been shown by initials within parentheses at the end of
the pertinent section. The overall format, the system adopted, selection
and sequence of the species, their scope, and the numbers and scope of
the subspecies have been confirmed by V.G. Heptner.

TABLE OF CONTENTS:

FOREWORD TO THE ENGLISH EDITION
FOREWORD

CLASSIFICATION OF CLASS MAMMALIA

KEY FOR IDENTIFYING ORDERS OF MAMMALS

PART I. ORDER OF PINNIPEDS
ORDER PINNIPEDIA ILLIGER, 1811

Cohort Ferungulata Simpson, 1945
Superorder Ferae Linnaeus, 1758
Order Pinnipedia Illiger, 1811
Family Odobenidae Allen, 1880 (Walruses)
Genus Odobenus Brisson, 1762: Walruses
Odobenus rosmarus (Linnaeus, 1758): Walrus
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Superfamily Otarioidea Smirnov, 1908
Family Otariidae Gill, 1866 (Eared Seals)
Subfamily Otariinae Boetticher, 1934 (Sea Lions)
Genus Eumetopias Gill, 1866: Steller’s Sea Lion
Eumetopias jubatus (Schreber, 1776): Steller’s
Sea Lion
Diagnosis
Description
Taxonomy

XXVII

* Pages 713-718 in the Russian original. The English table of contents is not a literal

translation.

Geographic Distribution 69
Geographic Variation 72
Biology WZ
Economic Importance 87

Genus Zalophus Gill, 1866: Northern Sea Lions 89
Zalophus californianus (Lesson, 1828): 92
California Sea Lion

Diagnosis 92

Description 92

Geographic Distribution and

Geographic Variation 93

Biology 96
Subfamily Arctocephalinae Boetticher, 1934: 96
(Fur Seals)

Genus Callorhinus Gray, 1859: Northern Fur 96

Seals
Callorhinus ursinus (Linnaeus, 1758): 98
Northern Fur Seal

Diagnosis 98

Description 98

Taxonomy 101

Geographic Distribution 101

Geographic Variation 105

Biology 108

Economic Importance 141

Superfamily Phocoidea Smirnov, 1908 142

Family Phocidae Gray, 1825 (True Seals) 142

Subfamily Phocinae Gill, 1866 (True, or 160
10-incisored, Seals)

Genus Erignathus Gill, 1866: Bearded Seals 164
Erignathus barbatus (Erxleben, 1777): 166
Bearded Seal

Diagnosis 167
Description 167
Taxonomy 174
Geographic Distribution 174
Geographic Variation 181
Biology 182
Economic Importance 209
Genus Phoca Linnaeus, 1758: True Seals and 212

Ringed Seals
Subgenus Pusa Scopoli, 1777: Ringed Seals 218

Phoca (Pusa) hispida Schreber, 1775:
Ringed Seal
Diagnosis
Description
Taxonomy
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Phoca (Pusa) caspica Gmelin, 1788:
Caspian Seal
Diagnosis
Description
Taxonomy
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Phoca (Pusa) sibirica Gmelin, 1788: Baikal
Seal or Baikal Ringed Seal
Diagnosis
Description
Taxonomy
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Subgenus Phoca Linnaeus, 1758: True Seals
Phoca (Phoca) vitulina Linnaeus, 1758:
Common Seal, Larga
Diagnosis
Description
Taxonomy
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Subgenus Pagophilus Gray, 1844: Harp or
Greenland Seals
Phoca (Pagophilus) groenlandica
Erxleben, 1777: Harp or Greenland Seal
Diagnosis

XVil

218

219
219
223
225.
291
234
256
260

260
261
266
267
269
269
286
290

290
290
295
295
295
295
304
307
307

308
308
313
315
323
330
364
369

369

370

ХУШ

Description

Taxonomy

Geographic Distribution
Geographic Variation
Biology

Economic Importance

Subgenus Histriophoca: Ribbon Seals
Phoca (Histriophoca) fasciata
Zimmermann, 1783: Ribbon (Banded) Seal

Diagnosis

Description

Taxonomy

Geographic Distribution
Geographic Variation
Biology

Economic Importance

Genus Halichoerus Nilsson, 1820:
Gray Seals
Halichoerus grypus (Fabricius, 1791): Gray Seal

Diagnosis

Description

Taxonomy

Geographic Distribution
Geographic Variation
Biology

Economic Importance

Subfamily Monachinae Trouessart, 1904
(Monk Seals or 8-incisored Seals)
Genus Monachus Flemming, 1822: Monk Seals
Monachus monachus (Hermann, 1779): Monk

Diagnosis

Description

Taxonomy

Geographic Distribution
Geographic Variation
Biology

Economic Importance

Subfamily Cystophorinae Gray, 1866 (Hooded

Syl
381
382
387
389
429
436
436

437
437
440
440
442
442
451
452

454
455
455
466
467
469
472
493

495
499
502

502
502
507
508
512
512
519
519

Seals and Elephant Seals, or 6-incisored
Seals)
Genus Cystophora Nilsson, 1820: Hooded Seals
Crystophora cristata Erxleben, 1777: Hooded
Seal
Diagnosis
Description
Taxonomy
Geographic Distribution
Geographic Variation
Biology
Economic Importance

PART II. ORDER OF WHALES
ORDER CETACEA BRISSON, 1762

Cohort Mutica
Order Cetacea Brisson, 1762
Suborder Odontoceti Flower, 1867
Superfamily Delphinoidea Flower, 1864
Family Delphinidae Gray, 1821 (Dolphins)
Genus Steno Gray, 1846: Rough-toothed
Dolphins
Steno bredanensis Lesson, 1828: Rough-
toothed Dolphin
Genus Stenella Gray, 1866: Spotted Dolphins
Stenella coeruleoalba Meyen, 1833:
Blue-white Striped Dolphin
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology

Stenella dubia [=айепиаа] С. Cuvier, 1812:

Malay [Pan-tropical Spotted] Dolphin
Stenella frontalis G. Cuvier, 1829: Bridled
[Atlantic Spotted] Dolphin
Stenella longirostris Gray, 1828: Long-snout
[Spinner] Dolphin
Genus Delphinus Linnaeus, 1758: Common
Dolphins

Delphinus delphis Linnaeus, 1758: Common
Dolphin
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Genus Tursiops Gervais, 1855: Bottlenose
Dolphins
Tursiops truncatus Montagu, 1821:
Bottlenose Dolphin
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Genus Lissodelphis Gloger, 1841: Right Whale
Dolphins
Lissodelphis borealis (Peale, 1848): Northern
Right Whale Dolphin
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Lissodelphis peroni (Lacépéde, 1804):
Southern Right Whale Dolphin (Peron’s
Dolphin)
Geographic Distribution
Biology
Genus Lagenorhynchus Gray, 1846: Shorthead
Dolphins
Lagenorhynchus (Lagenorhynchus) acutus
Gray, 1828: Atlantic White-sided Dolphin
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology

607

607
607
611
611
614
627
631

632

633
633
634
638
639
644
645

646

646
647
647
648
648

651

652
653
653

656

656
656
657
659
659

Lagenorhynchus (Lagenorhynchus) albirostris

Gray, 1846: White-beaked Dolphin
Diagnosis

Description

Geographic Distribution
Geographic Variation

Biology

Lagenorhynchus (Lagenorhynchus)

obliquidens Gill, 1865: Pacific White-sided
Dolphin

Diagnosis

Description

Geographic Distribution

Geographic Variation

Biology

Economic Importance

Lagenorhynchus (Peponocephala) electra

Gray, 1846: Broadsnout Dolphin
Genus Pseudorca Reinhardt, 1862:
False Killer Whales

Pseudorca crassidens Owen,

1846: False Killer Whale

Diagnosis

Description

Geographic Distribution
Geographic Variation
Biology

Ecanomic importance

Genus Orcinus Fitzinger, 1860: Killer Whales
Orcinus orca Linnaeus, 1758 : Killer Whale

Diagnosis

Description

Geographic Distribution
Geographic Variation
Biology

Economic Importance

Genus Grampus Gray, 1828: Risso’s Dolphins

Grampus griseus G. Cuvier, 1812: Risso’s
Dolphin

Diagnosis

Description

Geographic Distribution

ххИ

Geographic Variation
Biology
Genus Globicephala Lesson, 1828: Pilot Whales
Globicephala melaena Traill, 1809: Pilot
Whale
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Genus Feresa Gray, 1870: Pygmy Killer Whales
Feresa attenuata Gray, 1875: Pygmy Killer
Whale
Genus Phocoena G. Cuvier, 1817: Common
Porpoises
Phocoena phocoena Linnaeus, 1758:
Common Porpoise
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Genus Phocoenoides Andrews, 1911: Dall
Porpoises
Phocoenoides dalli True, 1885: Dall Propoise
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Genus Neophocaena Palmer, 1899: Black
Finless Porpoises
Neophocaena phocaenoides G. Cuvier, 1829:
Black Finless Porpoise
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology

698
699
701
702

702
702
704
706
708
VAY
718
719

721

722

72.2.
722,
725
726
729
735
737

738
738
738
740
741
744
748
750

750

750
750
751
sv
752

Family Monodontidae Gray, 1821 (Narwhals)
Genus Delphinapterus Lacépéde, 1804: Belugas
or White Whales

Delphinapterus leucas Pallas, 1776: Beluga or

White Whale
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance

Genus Monodon Linnaeus, 1758: Narwhals

(Narwhals or Unicorns)

Monodon monoceros Linnaeus, 1758:
Norwhal or Unicorn
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance
Superfamily Physeteroidea Gill, 1872
Family Physeteridae Gray, 1821 (Sperm Whales)
Subfamily Physeterinae Flower, 1864 (Sperm
Whales)

Genus Physeter Linnaeus, 1758: Sperm Whales
Physeter catodon Linnaeus, 1758: Sperm
Whale

Diagnosis

Description

Geographic Distribution

Geographic Variation

Biology

Economic Importance
Subfamily Kogiinae Gill, 1871 (Dwarf Sperm
Whales)

Genus Kogia Gray, 1846: Dwarf Sperm

Whales
Kogia breviceps Blainville, 1838: Dwarf
Sperm Whale

Diagnosis
Description

ххШ

755
755

IBY

150
757)
761
763
765
785
790

ТЭД

Е
792
792.
794
794
798
799
799
800

800
801

801
802
808
812
813
834
840

840

841

841
843

XXIV

Geographic Distribution and Biology
Kogia simus Owen, 1866: Owen’s Dwarf
Sperm Whale

Diagnosis

Description

Family Ziphiidae Gray, 1865 (Beaked Whales)
Genus Berardius Duvernoy, 1851: Pacific
Beaked Whales

Berardius bairdi Stejneger, 1883: Baird’s
Beaked Whale

Diagnosis

Description

Geographic Distribution

Geographic Variation

Biology

Economic Importance

Genus Mesoplodon Gervais, 1850: Beaked
Whales [Sword-tooth Dolphins]
Mesoplodon (Mesoplodon) stejnegeri True,
1885: Stejneger’s Beaked Whale

Diagnosis

Description

Geographic Distribution and Biology
Mesoplodon (Mesoplodon) bidens Sowerby,
1804: Sowerby’s Beaked Whale

Diagnosis

Description

Geographic Distribution

Geographic Variation

Biology

Genus Ziphius G. Cuvier, 1823: Goose-beak
[Cuvier’s Beaked] Whales
Ziphius cavirostris G. Cuvier, 1823: Cuvier’s
Beaked Whale

Diagnosis

Description

Geographic Distribution

Geographic Variation

Biology

Economic Importance

Genus Hyperoodon Lacépéde, 1804: Bottlenose
Whales

843
845

845
845
846
849

850

850
850
854
854
855
861
862

866

866
866
868
869

869
869
872
873
873
875

876

876
876
878
882
882
883
884

Hyperoodon ampullatus Forster, 1770:
Northern Bottlenose Whale
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Economic Importance

LITERATURE CITED
OTHER SOURCES

INDEX OF LATIN NAMES OF ANIMALS

957

989

hee
ie
и
ie (Na nahi
>. Lai |
& soy У by
т
| i ve
Gi > a к icv
a ae : т

a4

CLASSIFICATION OF CLASS
MAMMALIA

In this publication the old and widely used system of the major units of
the class, 1.е., orders, has been adopted. There is only one feature in it
which cannot be considered universal: pinnipeds are considered an inde-
pendent order and not a suborder of carnivores (Carnivora). However,
this approach, too, has many supporters. In the system of present-day
mammals, these two groups are separated naturally and fully, no less than
other orders. However, the close genetic relationship between pinnipeds
and land carnivores is striking and beyond doubt.

The system of orders adopted here is well founded, being based on
the morphology of present-day forms by M. Weber (1928) and of fossil
forms by G.G. Simpson (1945). The grouping of orders into taxa of a
much higher rank and the sequence of orders is after Simpson.

A tendency toward an extreme division of the orders has
recently developed. Thus it has been proposed that the order of
marsupials (Marsupialia) be divided into three (Polyprotodontoidea,
Caenolestoidea, and Diprotodontoidea) and even five (Didelphia,
Dasyuria, Peramelia, Caenolestia, and Phalangeria); insectivores
into four (Insectivora proper, Zalambdodonta, Macroscelidea, and
Tupaioidea); cetaceans into two (Odontoceti and Mysticeti); primates
into three (Lemuroidea, Simiae, and Tarsioidea); and artiodactyls into
two (Tylopoda and Artiodactyla proper). The number of orders has thus
increased from 18-19 to 30-31. None of these suggestions is yet well
founded.

Further, mammalogists dealing with extensive paleontological
material do not generally favor extreme division. An order is primarily
regarded as an integrating and not a differentiating concept. Otherwise,
there is the risk of a tendency, as in ornithological macrosystematics, for
the concept of an order to become essentially indistinguishable from
the concept of a family, and sometimes even simply substituted for
it. As a result, the scientific aspect of the system of vertebrates has
undoubtedly suffered, and is suffering even now, from the excessive

ХХУШ

division of the system which sometimes uses the term ‘fishlike vertebrates
and fish”. From the established viewpoint, Duplicidentata are regarded
as an independent order (Lagomorpha). Thus, we have here 19 instead
of 18 orders. Of these, 10 or 52.6% are represented in our fauna. One,
1.е., the sirenians (sea cow), is extinct.

The system of class adopted here is given below. The orders repre-
sented in the fauna of the USSR are marked with an asterisk. (V.H.)

CLASS MAMMALIA

Subclass PROTOTHERIA Order MONOTREMATA
Subclass THERIA
Infraclass METATHERIA Order MARSUPIALIA

Infraclass EUTHERIA

“Order INSECTITIVORA
Order DERMOPTERA
_| “Order CHIROPTERA
Order PRIMATES
Order EDENTATA
Order PHOLIDOTA

ee LAGOMORPHA

Cohort UNGUICULATA

Cohort GLIRES *Order RODENTIA

Cohort MUTICA “Order СЕТАСЕА
Superorder _| “Order CARVINORA
FERAE “Order PINNIPEDIA
Superorder

Canon PROTUNGULATA Order TUBULIDENTATA

FERUNGULATA Order PROBOSCIDEA

BS ies —| Order HYRACOIDEA

PAENUNGULATA | , Order SIRENIA
Superorder i |
MESAXONIA Order PERISSODACTYLA
Superorder

PARAXONIA Order ARTIODACTYLA

Key for Identifying Orders of Mammals [т the Soviet Union]

2).
РВ.
3(4).
4(3).
5 ( 6).
6 ( 5).
7 (10).
8 ( 9).
9( 8).
10 ( 7).
11 (14).
12 (13).
13 (12).

14 (11).

15 (16).

Hind limbs absent. Body fishlike with large bilobate caudal fluke
SCEMONIZOM alll yore soe heii. tare ae caer ee Mee ES CETACEA.
Hind limbs present. Body not fishlike; tail, if present, not in the
form of a bilobate fluke.

Forelimbs in the form of leathery wings...... CHIROPTERA.
Forelimbs of different structure.

Fore- and hind limbs very short, paddle-shaped, and in the form
of fins, 1.е., all digits right up to very tips enclosed in a common
SHAM О roc ate ay <a ue he ae akan PINNIPEDIA.
Fore- and hind limbs of different structure, not in the form of
fins.

Hooves on legs.

Only one hoof on each limb .......... PERISSODACTYLA.!
Two large hooves and two small ones set above them occur on
а creates ile ARTIODACTYLA.*

Hooves absent on legs (claws present).

Diastema occurs between large chisellike incisors and molars;
its length not less than length of entire row of molars on the
corresponding jaw. Canines absent.

MWOTNGCISOLS: ON UPPER JaW) Ps ose ene RODENTIA.
Four incisors on upper jaw; small blunt one occurs behind each
Olitwovlarge SharprONes 2. ase. este LAGOMORPHA.

Diastema between incisors and molars absent or much smaller
than length of molar row. Canines present.

Anterior portion of snout extends into well-developed small
proboscis. Anteriormost tooth on each jaw or only on upper
jaw much larger than adjacent tooth. ....... INSECTIVORA.

For description, see Vol. I.

The structure of the limbs differs in camels. Each limb ends in two broad calloused
pads and true hooves are absent, being replaced by two very broad claws. Camels are not
included in this key because they are domesticated animals (the wild camel is extinct) (for
_ description, see Vol. I).

If the tooth relation differs, the proboscis is always present; moreover, the forelimb

is extremely short, the wrist very broad with huge claws and set on edge—inner surface
backward (moles).

XXX

16 (15). Anterior portion of snout does not form proboscis. Anterior-
most tooth on each jaw not larger than adjacent tooth.......
CARNIVORA‘ (V.H.)

зоо eeiie)-evle (ele! ie. осо ооо соо

4 Skulls of the extinct Steller’s cow, а representative of the order of sea cows or Sirenia,
have been found on the coast of the Commander Islands. They are distinguished by the
absence of teeth.or even traces of them (alveoli) on either the upper or the lower jaw over
a length of about 60 cm (for description, see Vol. II, Part 1).

ORDER OF PINNIPEDS

Order Pinnipedia Illiger, 1811

iat

Whee
a

i
a
я

figs]
5

COHORT ОЕ CARNIVORES AND
HOOFED MAMMALS

COHORT Ferungulata Simpson, 1945

Superorder of Carnivores

Superorder FERAE Linnaeus, 1758

ORDER OF PINNIPEDS
Order PINNIPEDIA Illiger, 1811’

The order comprises mainly large mammals, morphologically well distin-
guished from the members of most of the other orders in their extreme
adaptations to an aquatic mode of life.

The body is somewhat spindle-shaped, streamlined, narrowed toward
both ends, with an extremely poorly developed or even undeveloped tail
(Fig. 1). Fore- and hind limbs are modified into oar-shaped paddles or
flippers, with only their distal portions projecting from the cylindrical
trunk and adapted for swimming. To a very small extent, these are used
for resting upon and for moving on a solid substrate, but this faculty
varies widely from family to family.

The fore- and hind limbs have five digits each. In some species (fam-
ily of true or earless seals, Phocidae) claws are well developed on the
fore flippers and, usually, on the hind flippers; in some species (all other
families) claws are absent on the fore flippers but present on the hind
flippers, though not in all species; in most species the claws are highly
reduced, with almost imperceptible rudiments (Fig. 2). The hand with

1 Рог reasons given (Vol. II, Part I, 1967, р. 53), Pinnipedia are regarded here as an
independent order separated from the order Carnivora although such an interpretation
is debatable. Weber (1928), Simpson (1945), Tenius and Hofer (1960), and some others,
mainly paleontologists, regard Pinnipedia only as a suborder of Carnivora. (K. Ch.)

12

13

Fig. 1. Body shape and dimensions of Pinnipedia (seals): A—lateral view;
B—dorsal view (K.K. Chapskii). 1—body length from anterior end of snout
(nostrils) to tip of tail in a straight line (Lcv); 2—same, length along the dorsal
surface (Lc); 3—total body length along the dorsal surface (with hind flippers,
Lp); 4—axillary girth; S—maximum girth, 6—length of fore flippers along outer
edge; 7—same, length along the inner edge from axillary fold; 8—tail length;
9—length of hind flippers; 10—girth of head around the ear openings.

Fig. 2. Shape of fore flippers and disposition and growth of claws on them
in the various families of Pinnipedia. A—eared seals (Otariidae); B—walruses
(Odobenidae); C—true seals (Phocidae) (figure by K.K. Chapskii).

long digits is not separated externally into individual rays; the digits are
covered with a web of skin, like a sheath, and set close together. The
foot, too, has a similar structure. The phalanges bearing the claws are
often slightly broadened and somewhat elongated with distinct gaps for
nail beds or without them (Otariidae).

The scaphoid (scaphoideum), lunate (lunatum) and _ central
(centrale) bones are fused in the carpus. The ulna and radius are

12

13

5

shortened but quite independent and not fused. The humerus is even
more shortened, but bears a highly developed deltoid crest; the for.
entepicondyloideum is seen in most cases but may be absent (even within
the same species). The clavicle is not developed.

The digits of the feet are highly elongated. Among earless seals (Pho-
cidae), however, the first and the fifth are particularly elongated (the fifth
digit is also considerably broadened). All the digits have a thick web of
skin joining them right up to the claws. In Phocidae, the elasticity of the
web facilitates powerful movement of the digits. When stretched, the web
forms a broad fan-shaped surface resembling the emarginate caudal fin of
a fish (Fig. 3). The end phalanges of the extreme digits of the feet are quite
broadened; in Phocidae, these have deep emarginate cavities for the claws.

The articular facet of the astragalus in true seals (Phocidae) is in
the form of a sharply bulged crest but slightly concave in walruses
(Odobenidae), and saddle-shaped in eared seals (Otariidae). The tibia
is relatively long; the fibula is normally developed, totally independent,
but fuses with the tibia in the proximal epiphysis in adults. The femur is
highly shortened, flattened, and broadened distally; the third trochanter
is absent; even the lesser trochanter is not developed in most cases.

The thoracic vertebrae usually number 15, the lumbar 5, and the

‚ sacral 4, while the caudal vary from 8 to 15. The vertebral column is

extremely flexible with highly developed intervertebral cartilages.

The skull differs in shape and size. In most cases, the upper wall of
the cranium is more or less distinctly flattened, while the sagittal crest
is poorly developed or absent. The cerebral portion is capacious, while
the facial portion is usually not longer than the cerebral but narrower
and disposed below it, though sometimes only slightly so (walruses).
The orbits are generally extremely broad and joined widely with the
temporal fossa; with rare exceptions, the zygomatic arches are greatly
shifted laterally while the interorbital space in most species is sharply
constricted. The bony palate is compact (neglecting the anterior and
posterior palatine fossae). The auditory bullae among many (Phocidae)
are highly swollen and rounded, but more flattened and complex among
others. The lachrymal bone is absent. The fossa pterygoidea is not devel-
oped. The alisphenoid canal is absent in true seals (Phocidae) but present
in others (Fig. 4). The ethmoturbinal bones are small and arranged in
five folds; the maxilloturbinal bones are usually highly developed and fill
most of the nasal cavity.

The mandibular condyle is semicylindrical with a fairly distinct con-
cavity (saddle-shaped) in the central part. The corresponding articular
fossa in the basal part of the skull is transversely truncate, quite extended,

Fig. 3. Structure of hind flippers: (top to bottom)—true seals (Phocidae), wal-
ruses (Odobenidae), and eared seals (Otariidae) (figure by N.N. Kondakov).

but of varying depth; however, neither its upper nor lower crests are so
highly developed and flexed as to reach the mandibular condyle.

The Pinnipedia are heterodont but the carnassial teeth characteristic
of predators (Carnivora) are not developed. The most common dental

formula is: 3 1 4 1
I- С -Р- М —-=34.
2 1 4 1 :
The number of teeth is less in some seals and walruses. The more

common permanent teeth formula among walruses is:
1 1 ыы

14

14

о

yesh a Mp EET

a TEE yy

3

Fig. 4. Disposition of alisphenoid canal (shown by an arrow) in the skull of
Steller’s sea lion Eumetopias jubatus (figure by N.N. Kondakov). 1—uncinate
process of pterygoid; 2—zygomatic; 3—palatine bones.

The incisors in the lower jaw are relatively small, sometimes very
weak; the lateral incisors in the upper jaw are highly developed as are
the canines. The canines, especially the upper ones, are quite large; they
are massive in walruses. The cheek teeth are less differentiated and usu-
ally flattened laterally. The structure of the crown is extremely diverse,
in most cases with many cusps, but the main cusp is usually raised much
above the others located behind and in front (or only behind). Mono-
cuspids are not uncommon. The premolars have one or two roots; the
true molar more often has two roots. Sometimes there is a diastema (or
its analogue) between the fourth premolar and the molar.

The stomach is simple with a poorly developed cecum. The brain is
microsmatic and relatively large. The hemispheres are highly furrowed
with convolutions and four suprasylvian fissures.

The hair coat is variously developed: very dense in fur seals and in the
newborn pups of almost all pagophilic species associated with ice. In an
overwhelming majority of the species, it consists of three categories: fur,
intermediary and guard hairs. In walruses, sea lions (genus Zalophus),
and true seals (family Phocidae) (except the newborns) the hair coat
is quite rough and relatively short, and the fur hairs very sparse with
practically no underfur. The hair coat is sparsest in walruses and bearded
seals (genus Erignathus). The tail is covered with hairs. The whiskers are
well developed and are most abundant in walruses; among seals, these
are most numerous in the bearded seal. The whiskers are longest in the
eared seals [Otariidae]. The color of the hair coat is quite diverse; it is
fairly monochromatic and dull in eared seals and walruses (Otarioidea)
and mostly spotted in true seals (Phocidae) (in some of them with bright
contrast). A fairly distinct color variation with age is characteristic of
many.

15

The skin glands are normally developed but there are no special
scent glands. Mammary glands occur on the ventral side of the body
with one or two pairs of mammary teats. The testes are concealed under
a subcutaneous adipose layer at the base of the hind flippers and are not
externally visible in true seals. Some kind of scrotum is seen in eared
seals (Otariidae), which is less developed in walruses. The copulatory
organ is concealed in a preputial pouch under the skin in front of the
testes; the preputial pore opens out ventrally between the navel and the
anal orifice. An os penis is present. The genital opening in the female
is disposed directly in front of the anus under a common skin fold. The
uterus is bipartite. A rudimentary os clitoridis is present. The placenta
is girdle-shaped (zonary) and deciduate.

Sexual dimorphism varies sharply. In polygamous species (eared
seals, Otariidae; elephant seals, Mirounga; walruses; and some others)
and also in hooded seals (Cystophora), the male is considerably larger
than the female. Some other features of dimorphism are also evident in
the male: processes of the nasal cavity in the form of a proboscis (trunk)
or a hoodlike swelling (elephant and hooded seals), features of skin
structure (tuberculate formations in the walrus), and some differences
in the thickness and structure of the hair coat (fur seal, Callorhinus; sea
lion, Zalophus). In many other species, differences are restricted to a
somewhat more massive skull, powerful dental apparatus, in particular
much larger canines, and some differences in color, size, and proportions
of the body and skull.

Age related changes are quite significant. Pups differ from growing
juveniles and the latter from adults in the structure and color of the hair
coat; external sexual differences become prominent with age. Seasonal
dimorphism is not developed. Molt occurs once, often with casting of
the cornified layer of the epidermis.

The range of size differences among the various species of Pinni-
pedia is not particularly large. The smallest of them (ringed seal, Phoca
hispida; Caspian seal, Ph. caspica; and females of the northern fur seal,
Callorhinus ursinus) is rarely longer than 170 cm in adulthood (from
tip of nose to end of tail dorsally, Lc) and usually weighs up to a hun-
dred kilograms. However, the minimum body size of adults of individual
populations is fairly low: the smallest of them, especially the Okhotsk
ringed seal, is only a little over 1 m long (Lc) and weighs barely 20 kg.
The largest of the Pinnipedia (elephant seals, Mirounga, and the male
walrus) measure (Lc) up to 600 and 400 cm long and weigh 4-5 and 1.5 °
tons respectively. The lower limit of weight is at least two-hundredths of
the upper limit, which itself is somewhat (by at least one-fifth) less than
the corresponding range for the order Carnivora.

9

In overall build, proportions and external appearance, biological
types, forms of adaptations, and other features, Pinnipedia also do not
exhibit as great a diversity as Carnivora. Adaptations to living in water
and for overcoming its resistance during fast swimming have left a gen-
eral imprint on the entire external appearance by evolving a streamlined
body. Evolution proceeded mainly in two directions in developing the
means of locomotion in water: 1) strengthening of the locomotor role of
the hind limbs and 2) transfer of the main function of propulsion to the
forelimbs. As a result, in some members of this group of mammals, 1.е.,
true or earless seals (Phocidae), the hind flippers became more developed
and adapted to serve as the main propeller in water; these flippers. are
turned backward and modified into paired caudal fins, similar to those of
a fish. These are also broadened and emarginated distally. In the other
group, eared seals (Otariidae), the fore flippers became the main source
[organ] of propulsion and evolved into massive crested lobes, almost
wholly devoid of a hair coat and claws. In this respect walruses occupy
an intermediate position since, besides the powerful fore flippers, they
also have hind flippers identical in structure with those of true seals,
which actively participate in forward motion during swimming. The body
curvature also plays a significant role in locomotion.

In conformity with the different types of locomotion in water, these
two main groups of pinnipeds are differently adapted to moving on land.
The hind flippers of true seals (Phocidae) do not bend forward and serve
as a support on a hard substrate. The fore flippers are weak and also
poorly adapted to movement on land. The animals rest on them and,
clutching the ground with the claws, haul up the rest of the trunk in
jerks and drag it along the ground. In this case their movement looks
like typical spasmodic heaves or resembles somewhat that of geometrids
[inch worms]: resting on the rear part of the body, they haul the front
part forward and then, shifting the center of gravity to it and quite often
clutching the substrate with the claws of the fore flippers, haul up the
rear. In some species, for example the Caspian seal, the fore flippers of
well-fed specimens are so small that they do not reach the ground. The
alternation of points of support is rapid and hence advancement is not
that slow but is short in duration—the animal soon becomes exhausted.
Eared seals can rest on the hind flippers (bending forward at the ankle
joint) as well as on the wrist of the fore flippers, and can shift from one
foot to the other, though quite awkwardly, and thus hop. They can negoti-
ate fairly steep rocky shores on which they climb with great agility in spite
of the fact that their limbs are apparently poorly adapted for this purpose.

The bulkiest species, elephant seals (Mirounga) and walruses, move
rather slowly on land. All the same, they exhibit extreme agility under

16

10

certain circumstances. Steller’s sea lion can dive into water from a great
height (Fig. 51). The animals swim in any position—on the back, side,
or belly, in a submerged state or with the head above the water surface
(or surfacing only periodically). Some are capable of jumping fairly high
above the water (fur seals) but even the very heavy seals can leap from
the water onto an ice floe in one jump.

Pinnipedia live in diverse climatic zones—arctic and antarctic, boreal
and austral, and even subtropical. Depending on the season, they live in
herds (often very large and dense) or in small isolated groups, singly or
in pairs (for a short duration). In the wider sense of the word, not even
one species can be regarded.as a resident, including even those which
are more confined to a given section or region. Somewhat developed
migrations, or at least local wanderings, are typical of almost all the
species of pinnipeds. Migrations in many species have a well-defined
character (fur seal, Greenland seal, hooded seal, walrus, etc.) marked
no less precisely than bird migrations, and extend over large distances
(hundreds or even thousands of kilometers).

Despite all their adaptations to life in water, pinnipeds do require
solid substrate on which they give birth, suckle the pups, molt, and in
most cases mate, and simply rest. Some of them use only ice floes for
this purpose (pagophilic? species and subspecies), while others select
beaches, mostly of islands (aegialoid [beach or shore loving] or pagopho-
bic> species and subspecies). As a rule, the more thermophilic species
use the land (eared seals, Otariidae; subtropical seals with 8 incisors,
Monachinae; and also some Phocinae with 10 incisors—true seals, and
the West European form of gray seal). The rest of the 10-incisored seals,
the hooded seal, and antarctic 8-incisored seals usually reproduce and
molt on ice. Some pagophilic species (ringed and Baikal seals, Caspian
seal, Greenland seal, and others) make round openings in the ice cover
for breathing and ventilation, and for crawling onto the ice. Some bur-
row holes in the ice for themselves and for concealing their pups under
the snow cover. There is no winter hibernation.

Polygamy is common to the aegialoid [littoral; pagophobic] pin-
nipeds. There is no strict monogamy since the males and females come
together only for a short while, invariably preceded by fairly severe fights
among the suitors. Males play no part in the care of the progeny.

Fertility is low: all pinnipeds usually deliver a single pup; twins are
extremely rare. The newborns are large, fully formed, covered with a thick
coat, with normally developed limbs and open eyes. Postnatal growth

2 Ice-loving, according to the terminology of М.А. Smirnov (1914, 1927, and 1936).
3 Land-loving, ice-fearing species.

18

т

is quite rapic in some (hooded seal, Greenland seal, and most other
pagophilic seals) and less so among others (Baikal seal, ringed seal). In
some others, growth is very slow (walrus, fur seal, sea lion).

The feeding habits of pinnipeds are not clearly differentiated, though
there are some mainly benthic feeders among them (walrus, Odobenus;
bearded seal, Erignathus); some live on very large plankton, mostly
fish and cephalopods (hooded seal; elephant seal; fur seals, Callorhinus
and Arctocephalus;, sea lion, Zalophus), and some live on planktonic
crustaceans (crabeater, Lobodon). Most survive on mixed food consisting
mostly of small and minute fishes, large plankton, some demersal
crustaceans, as well as cephalopods and other mollusks. The leopard seal,
HAydrurga, feeds also on large fish and warm-blooded animals, including
birds (penguins).

Among the sense organs, the most developed are those of hearing
and vision. The bony tympanic bullae are highly bulged in true seals (Pho-
cidae) but flattened in all others. The pinnae are either altogether absent
(true seals and walruses) or highly reduced (eared seals, Otariidae).

Pinnipeds are distributed in all the oceans except the Indian Ocean;
they are found in the southernmost fringe of the latter (generally not
above 30° S lat.) from the adjoining antarctic waters (Fig. 5).

An overwhelming majority of the species are confined to the regions
of cold and moderately cold waters in which the surface temperature does
not exceed 20°C at any time of the year (Davies, 1958b*). The exceptions
are primarily thermophilic seals of the genus Monachus, with isolated
distribution in three regions of the subtropical belt (one species in the
Mediterranean Sea, in the open ocean around Gibraltar and Western
Africa, and in the Black Sea; another in the Caribbean Sea‘; and a third
in the Hawaiian Islands). Many others can withstand relatively warm
water conditions: the southern population of the northern elephant seals
(Mirounga in California), the California, and Galapagos sea lions (Zalo-
phus), the South African Cape fur seal (Arctocephalus pusillus), the South
Australian and New Zealand eared seals of the genera Neophoca, Pho-
carctos, and Arctocephalus, some populations of the northern elephant
seal (Mirounga angustirostris), and others.

Pinnipeds are absent not only in the Indian Ocean but also in the
Malayan archipelago, southwestern Pacific Ocean south of 30° М lat. to

* Here and throughout the text an asterisk (*)-after a reference in the text indicates
that either the author is not listed in the “Literature Cited” or the author is listed but no
entry given for the publication date cited - General Editor.

4The Caribbean monk seal (Monachus tropicalis) is extinct; the few survivors of its
populations were finally exterminated in the first half of this century.

‘(irysdeyD УМУ) Bag yoRTG 94) рие элэцазииэц илэЧ 1 ш Aepunog WIsyVION

OOS = ОБ ae Om. Bat Oe EOS

110$ ЭЧ -elpodiuutg лор
ЕЕ LF Е. | | sey Ее

р =

oa
SNL

19

13

40° $ lat., and in its entire central part south and north of the equator
up to the 30th parallel. The only exception is the small area around the
Hawaiian Islands occupied by the relict small population of the Hawai-
ian monk seal, Monachus schauinslandi. Pinnipeds are totally absent in
the entire tropical portion of the Atlantic Ocean from 20° М lat. to
30°S lat. and some other pelagic parts of the world oceans, including
a large expanse of temperate latitudes of the Atlantic to north of the
equator (Figs. 5, 80).

At present, pinnipeds are most numerous in the boreal and arctic
regions of the Northern hemisphere and in the temperate zone of the
Southern, especially in the zones of confluence of cold and warm waters
(zone “front,” convergence).

Some 14 species inhabit the seas of the Arctic Ocean, northern and
subarctic parts of the Atlantic, and the northern parts of the Pacific
Ocean (mostly true seals with 10 incisors, i.e., Phocidae, walrus, and
some eared seals); just as many species inhabit the waters of the South-
ern hemisphere (mostly eared and 8-incisored true seals). The remaining
species are the above-mentioned subtropical monk seal and the inhabi-
tants of landlocked waters such as the Caspian Sea and Lake Baikal.

The species most adapted to living in icy conditions (especially the
ringed seal and to a lesser extent the bearded seal) penetrate into very
high latitudes, into the Atlantic sector of the Arctic, and reach north of
the 80th parallel and even the polar regions in eastern Severnaya Zemlya.

The zoogeographic range of pinnipeds is divided (Skleter,* 1897; and
others) into five regions: 1) Arctoatlantic, including the Arctic Ocean and
North Atlantic (endemic monotypic genera: hooded seals, Cystophora;
gray seals, Halichoerus, to which may be added the Greenland seal);
2) Arctopacific, covering the extensive area of the North Pacific (endemic
genera: northern fur seals, Callorhinus; sea lions, Eumetopias); 3) Mesat-
lantic (subtropical belt intersecting the Atlantic Ocean on both sides of
the Tropic of Cancer and including the Mediterranean and Black seas)
—area of distribution of the Caribbean and Mediterranean monk seals;
4) Mesopacific—similar to the small zone around the Hawaiian Islands
where the third member of the genus of monk seals, Monachus schauins-
landi, lives; and 5) Notopelagic—the zone encircling the Antarctic and
extending north roughly along the 20°C isotherm of February surface
temperature of the sea on the coasts of Chile and Peru and also covering
the coastal waters of South Africa, South Australia, and New Zealand
(endemic genera: monotypic genera of the Antarctic, 8-incisored true
seals—Phocidae; also the southern eared seals—Otariidae, especially the
genera Otaria, Neophoca, Phocarctos, and many other species of southern
fur seals of the genus Arctocephalus).

14

The general range of the order (Pinnipedia) did not undergo serious
changes in the historic past, though in the eighteenth, nineteenth, and -
twentieth centuries significant “gaps” have been noticed. Some popula-
tions of the various species of southern fur seals (Arctocephalus) were
totally exterminated and some of them, A. philippi (South America),
almost completely. The Caribbean monk seal, M. tropicalis, has also
become extinct in the present century while the Mediterranean species,
M. monachus, is facing extinction.

The populations of many species have shrunk sharply while those of
many others continue to shrink in the North and Arctoatlantic (common
seal, Phoca vitulina; walrus and, at places, the gray seal, Halichoerus
grypus; to a lesser extent populations of ringed seal, Ph. hispida, and
some others) as also in the northern part of the Pacific Ocean (sea
lion, Zalophus californianus,; walrus); the population of the bearded seal,
Erignathus barbatus, and of the ribbon seal, Histriophoca fasciata, has
decreased sharply as a result of unrestrained hunting.

Conservation measures and a rational system of utilization have
helped restore the populations. The population of the northern fur seal
(Callorhinus ursinus) and that of the following southern fur seals have
already reached high levels: Arctocephalus pusillus (in South Africa), A.
australis (at the coasts of Uruguay and Argentina), and the elephant seal
(Mirounga leonina) (especially on the island of South Georgia). Popu-
lations of the other southern fur seals and elephant seals as also the
northern fur seal (at the Pacific coast of North America), which faced
extinction, have begun to rise steadily.

The total world population of all the species of pinnipeds is esti-
mated at 16-20 million. The maximum numbers exceeding 1 million are
attributed to four species: 1) the ашагсис crabeater seal, Lobodon car-
cinophaga, probably numbering 4-5 million; 2) the ringed seal, Phoca
hispida, somewhat less numerous than the preceding species (1-2 mil-
lion); 3) the Greenland seal, Phoca groenlandica, has a total population
of 1.5-2 million; and 4) the northern fur seal, Callorhinus ursinus, has
a total population of 2-2.5 million. The census of the southern sea
lion, Otaria byronia, is estimated at 800,000, that of the southern ele-
phant seal at 600,000, and for the bearded seal (Erignathus barbatus),
hooded seal (Cystophora cristata), Weddell’s seal (Leptonychotes wed-
delli), South African fur seal (Arctocephalus pusillus), and Caspian seal
(Phoca caspica)—400,000 - 500,000 each. The population of most other
species is even less (Chapskii, 1966; and others).

Phylogenetically, the Pinnipedia are very close to the Carnivora from
whose more primitive ancestors they undoubtedly evolved comparatively
recently (in the geological sense of time), but evidently not later than in

20

15

the Oligocene. In fact, members of this order belonging to the Oligocene
have not yet been discovered. The oldest known fossils are from the
Lower and Middle Miocene. These were fully formed members of the
same families to which the present-day Pinnipedia belong. Thus, a mem-
ber of the family of eared seals could be recognized from the lower jaw
with teeth of the Lower Miocene seal, Allodesmus kernensis Kellogg. Sim-
ilarly, the Middle Miocene genera Leptophoca, True, Miophoca, Zapfe, or
the Upper Miocene genus Monotherium van Beneden belong to the fam-
ily of true seals (Phocidae). At that time (if not earlier), the present-day
subfamilies of Phocidae already existed.

The Miocene seals did not differ considerably from the present-day
species and morphologically did not stand very close to any of the land
or semiaquatic carnivores which could serve as the ancestral form for
the evolution of pinnipeds. All this offers a basis for presuming that the
initiation of the primary phyletic branches leading to the families of sea
lions and fur seals (Otariidae), the walrus (Odobenidae), and true seals
(Phocidae) should belong to a much earlier period than the Miocene.

Many cardinal differences in body structure between the eared
(Otariidae) and earless (Phocidae) seals greatly complicate establishing
a single root which could be regarded as the base for the evolution of
this group of mammals. Among these characteristics are: features of the
tarsal region (especially the structure of the ankle bone) and the resultant
differences in the ability of the animals to move on land, difference in
the general structure of flippers, numerous elements of craniological
dissimilarity (structure of tympanic region, mastoid, its association with
the paroccipital process, and presence or otherwise of the alisphenoid
canal).

Quite some time ago (Maivart, 1885*) attention was drawn to the
fact that one group of pinnipeds (eared seals and the walrus, Otari--
Oidea) was significantly close to bears in many craniological features,
while another group (true seals, Phocidae) reveals a similarity, though
less distinctly, with martens. From this arose the concept of diphyletic
origin (Kellogg, 1922; Howell, 1929,* 1930*; McLaren, 1960; Chapskii,
1963; King, 1964; and others).

In spite of some features complicating the adoption of this hypothe-
sis, especially that bears represent a much younger branch of carnivores
whose evolution is put in a much later period than Pinnipedia, the pos-
sible diphyletic origin of the latter is not excluded. The evolution of
true seals (Phocidae) from martens is quite convincingly demonstrated
by the find of a seallike otter—seal semantor, Semantor macrurus, in
the Pliocene formations of Western Siberia (Orlov, 1931*). This animal
possessed distinct transitional features of structure between the sea otter

21

16

and true seals. It was closer to the seal than the sea otter with respect to
the humerus (Kirpichnikov, 1955), though some other skeletal elements
have revealed, on the contrary, a very close genetic similarity with the sea
otter (K.K. Chapskii). The historical evolution of true seals (Phocidae)
evidently proceeded in a similar manner, but in a much earlier geologi-
cal age. The semantor was a somewhat incomplete branch formed in the
same direction, independent of the emerging seals, but under different
conditions. This could serve as an argument in favor of the validity of
different ways of formation of the different groups of Pinnipedia and,
hence the possibility of interpreting the latter not as a phylogenetically
valid order (or suborder) but as a composite group of animals arising
from different roots.°

The extremely scant paleontological material on true seals in general,
and especially the recent finds within the USSR, make it difficult to
estimate the exact number of not only species, but even of genera of
fossil Phocidae.

At present, 42 genera of Pinnipedia are known. Of these, 25 are
fossil (Simpson, 1945; McLaren, 1960; Mitchell, 1961,* 1968*) and 17
Recent. This proportion does not, however, indicate that Pinnipedia are
at the end of their evolutionary development. The Recent genera form a
fairly large figure of 40%, which is much higher than the corresponding
figure (28%) for the “contemporaneity” of land carnivores (Heptner er
al., 1967*).

The other indices from which the potentially developing state of the
order could be judged are seen from the areas of distribution, population,
and biological stability of the different species. The range of the common
seal (Phoca vitulina) is very extensive and is interrupted only by ecological
barriers; the range of the bearded seal (Erignathus barbatus), the ringed
seal (Ph. hispida), and the genus of southern fur seals (Arctocephalus)
is fairly large though not continuous; the range of the northern fur seal
(Callorhinus ursinus) and the Greenland seal (P. groenlandica) is also
fairly large.

The relative diversity of forms in spite of the narrow specialization
of the order as a whole is per se significant. The high population of many
species (see above), including even such relict species as the Caspian seal
existing under conditions of intense anthropogenic pressure, is an impor-
tant feature. In spite of their low fertility, pinnipeds can quite quickly

5 The origin of a group or two highly proximate families of the same order (suborder)
could hardly be regarded as polyphyly. This, at best, is paraphyly or a form of monophyly.
(V.H.)

17

restore their populations and thus largely withstand destructive hunt-
ing. They cannot, however, survive incessant killing. Thus, one species
(Caribbean monk seal, Monachus tropicalis) became totally extinct, while
the population of another (Mediterranean monk seal, M. monachus) is
vanishing right before our eyes; the population of yet another (Hawai-
ian monk seal, M. schauinslandi) is very small and only exists because
of protective measures. Some other populations, especially of antarctic
species, are in a pitiable state as a result of irrational killing. The popu-
lation of all species of pinnipeds is wholly dependent on their judicious
utilization and conservation.

The classification of the Pinnipedia is quite simple in its general fea-
tures and more or less generally accepted. The differences arise only in
the number and scope of systematic categories and the position. of the
entire group which is interpreted sometimes as an independent order
or suborder of carnivores (Carnivora), and sometimes even as a com-
posite group deserving a rank of several families (or subfamilies) in
the superfamily Canoidea of carnivores (Carnivora). Moreover, none of
the researchers inclined to such an interpretation (Frian, 1956*; Lien
and Waiens, 1956*) have been able to specifically accomplish such an
arrangement. The Pinnipedia, regarded as an independent order, are
divided into two superfamilies: Otarioidea and Phocoidea. The former
consists of two families: eared seals, Otariidae (Steller’s sea lion, sea
lions, and fur seals) and walruses, Odobenidae. The second superfam-
ily comprises only a single family—true seals, Phocidae.® Quite often
(N.A. Smirnov, 1929, 1935; and others), only two families are recog-
nized: Otariidae s. lato, which includes not only eared seals, but also
walruses, and Phocidae, true seals.

Otarioidea are characterized by the following features: the bullae
osseae are flattened and have complex angular outlines; the mastoid
forms a single extremely massive process in adults; the process descends
much below the tympanum part and fuses with the pr. paroccipitalis;
the alisphenoid canal is present. The astragalus resembles more that
of Carnivora than of members of Phocoidea. The hind limbs can bend
forward and are adapted to locomotion on land.

Phocoidea are characterized by the following features: the tympanic
bullae are fairly strongly bulged and rounded; the mastoid process is not
joined with the pr. paroccipitalis (when present), not bent downward,
and generally not well developed. The alisphenoid canal is absent. The
astragalus has a large process at the top (no similarity whatsoever with

6 The fourth family, Semantoridae (fossils of seallike otter—semantors; Yu.A. Orlov,
1931), sometimes placed in the Pinnipedia (Simpson, 1945), should not be included here.

22

18

that in land carnivores) and its articular surface is not saddle-shaped.
The hind limbs do not bend forward and do not participate at all in
locomotion on land.

There is no basis for assuming that new species not known so far will
be detected in the order.’ At the same time, the morphological features
of the subspecies of many true seals (Phocidae) should be reviewed. The
scope of some genera of this family should also be partly reviewed. How-
ever, there is hardly any need to revise the classical division of phocids
into subfamilies, as recommended by King (1966). In the family of eared
seals, Otariidae, there is evidently need for a more rational argument for
the independence of all the species of the genus Arctocephalus.

Thus the system of eared seals of the Southern hemisphere at the
level of genera and species cannot yet be regarded as conclusively estab-
lished. There is no single opinion even about the scope of the genus of
some northern seals of the subfamily Phocinae in view of the attempt
of some theriologists to give a broader interpretation to the concept
of genus. The antarctic 8-incisored seals can be regarded as conclu-
sively established monotypic genera. The morpho-ecological rationale of
changing the measurements of taxonomic differences prevailing among
seals of the subgenera of the widely interpreted genus Phoca (true seals)
is highly substantiated.

At present, 32 -34 present-day species are included in the order Pin-
nipedia, i.e., only some 1% of the total number of species of mammals
of the world. On average, a genus has 2.1 species. There are a maxi-
mum of 18-19 species in the family of true seals, Phocidae; eared seals,
Otariidae, have 12-13 while there is only one species in the family of
walruses, Odobenidae. The present-day pinniped fauna of the world is
represented almost evenly in the North Atlantic basin (10 species includ-
ing the Caspian seal), in the North Pacific Ocean (11-12 species) and
in the seas of the Southern hemisphere (13 species).

Two superfamilies, all the 3 families, and 8 genera, 1.е., 42% of the 19
present-day genera, are known in the seas and landlocked water bodies of
the USSR. However, the fauna of the USSR comprises only 13 species,®
1.е., about 39% of all the species known in the world fauna.

7 The unexpected recent “discovery” in the Pacific Ocean of a “new” species, described
under the name of the island seal Phoca insularis (Belkin, 1964), was actually one of the
forms of the pagophobic common seal, Phoca vitulina (see p. 314, and also Chapskii, 1967,
1969).

8 If the common seal (Phoca vitulina) and the subspecies are regarded as a single species;
see pp. 323 - 330.

23

19

The range of the order encompasses all the oceans of the USSR and
some of the largest inland water bodies (Caspian Sea, Lake Baikal, Lake
Ladoga).

The practical importance of pinnipeds is very high. Almost all the
species, but primarily those which periodically form massive herds, rep-
resent game animals of great economic value and are hunted mainly for
their fur and partly for their skin; the fat, meat, and some other body
parts are also used.

Historically, the hunting of pinnipeds has played a major role. Seal-
ing vessels combed the seas of both hemispheres in search of herds of
these animals, especially fur seals, and killed them mercilessly. This car-
nage continued for decades and ultimately led to the near total depletion
of stocks of the commercially valuable species. In the Southern hemi-
sphere, all the rookeries of the southern fur seal were almost wholly rav-
aged and the elephant seal population suffered. The latter faced extinc-
tion even at very low latitudes. The rookeries of the northern fur seal on
the Commander and Pribilov islands were subject to intense destruction.
Effective conservation measures alone prevented the recurrence of the
fate that befell many populations of the southern fur seal. The stocks
of walrus were severely depleted, especially in the Atlantic sector of the
Arctic, and also in other regions of its habitat.

In spite of its sordid history, sealing at sea has not lost its importance
to date. In some regions of the globe it continues to play an important
economic role as a supplier of raw material for the fur, hide, and other
industries. In some countries (such as Greenland), sealing is extremely
important and represents the only source of livelihood for northern
native populations. Hunting of pinnipeds is of great significance in the
life of the native coastal populations of Alaska and the Chukchi Penin-
sula, for the coastal villagers of the Soviet Arkhangel’sk region, and for
hunters of some other regions of our country. It provides Eskimos and
some coastal Chukchians meat for themselves and for their dogs, skin
for making footwear, harnesses, and other requirements, as well as fat,
etc. Moreover, it provides work for the people engaged at the collective
farms and in the state-owned sealing industry.

In the USSR and many foreign countries, seal skins are in demand
as highly valuable, durable, and fashionable furs. The most important
targets of sealing are the northern and southern fur seals, the Green-
land seal, the ringed seal, the Caspian seal, the hooded seal, etc. Fur
is generally obtained from fur seals and mainly young Greenland and
Caspian seals, hooded seals, etc., whjle the skin is obtained from large
animals. Blubber is mainly used commercially in the tanning industry,

24

20

for soap-making, etc., for making medicinal (“fish”) ой, and as an ingre-
dient in other products. Meat is used locally. Some pinnipeds (e.g., larga
seals) inflict much damage in fisheries by thriving on prime fish, espe-
cially salmon. The most important commercial species of seals are now
exploited almost everywhere on a rational basis, eliminating the danger
of their depletion. (K. Ch.)

1 (4).

2 (3).

3 (2).

4 (1).

1 (4).

2 (3).

Key to Families of Pinnipedia
Identification Based on External Features

Hind flippers bent at calcaneal joint with foot forward, serving
as body support on hard ground. Fore flippers longer than hind
ones, naked distally; claws on hands absent or poorly developed
and placed well behind margin.

Snout narrowed anteriorly and quite elongated; pinnae small.
Hands long, roughly triangular, wing-shaped. Claws absent (or
very small). Well-developed claws only on three middle digits
of hind flippers. Upper canines (in adults) not protruding from
СОЗ MOUS oe ogee yee yoke ops Eared seals, Otariidae (p. 58)
Snout very broad, anteriorly somewhat truncated and short. Traces
of pinnae not seen. Hands not highly elongated and not triangular;
small reduced claws seen on digits of hands and all digits of feet.
Upper canines notably protruding from closed mouth. .........
FUER A RAE APE BES RII oR OA Walruses, Odobenidae (p. 22)
Hind flippers not bent forward at calcaneal joint, always held
backward, and not serving as body support on solid ground. Fore
flippers not long, slightly shorter than hind ones, and fully cov-
ered with fur. Hand with well-developed claws at end of digits.
AUT LONI Coa eee eer ne Sa Barada True seals, Phocidae (p. 142)

Identification Based on Skull Features

Tympanic bullae relatively small, flattened from top downward,
with uneven rugose surface, and complex outline. Alisphenoid
canal (Fig. 4) present. Mastoid part of temporals with very mas-
sive protuberance directed downward. All or almost all cheek
teeth with single root (the last with two roots) and with simple
undivided crown.

Rostral portion [of skull] slightly narrower than cranial part in tem-
poral region. Ends of middle pair of upper incisors with transverse
angular notch (Fig. 6). Nasals shifted backward and projection of
frontals lodged in fork between their apices (Fig. 7). Supraorbital
processes well developed. Lower jaw teeth sharply differentiated

25

24

24

3 (2).

4 (1).

21

into incisors, canines, and cheek teeth (premolars and molars
similar). Upper canines similar to lower ones in size and structure.
payne ча рН. she ha Maes Eared seals, Otariidae (p. 58)
Rostral portion [of skull] hardly less wide than cranial portion
in temporal region. Ends of nasals posteriorly, at junction with
frontals, sharply incised transversely, not forming acute apices
(Fig. 7). Supraorbital processes not developed. Lower jaw teeth
similar to one another; upper canines large and may attain large
size, bearing no similarity to the lower ones, which do not differ
fromycheek teeth: ...‹........... Walruses, Odobenidae (p. 22)
Tympanic bullae relatively large, bulging, with almost smooth
semicircular surface and comparatively simple outline. Alisphe-
noid canal absent. Mastoid part of temporals without mas-
sive downwardly directed protuberance. Most cheek teeth with
two roots and with divided crown. ......................0 000

Fig. 6. Transverse notch in middle upper incisor of eared seals, Otariidae, and

the fur seal, Callorhinus ursinus (figure by К.К. Chapskii).

Fig. 7. Structure of nasals in pinnipeds of various families: A—eared seals (family
Otariidae); B—walruses (family Odobenidae); C—true seals (family Phocidae):

1—frontal, 2—maxillary, 3—premaxillary, 4—nasal bones.

22
25 Family of Walruses

Family ODOBENIDAE Allen, 1880

This is one of the largest groups of animals of the order Pinnipedia. The
trunk is massive; the head is rounded and, compared with the massive
body, appears small. Pinnae are absent. The whiskers are long, dense,

24 Fig. 8. Left zygomatic bone in pinnipeds: A—eared seal (fur seal, Callorhinus
ursinus); B—walrus (Odobenus rosmarus), C—bearded seal (EFrignathus
barbatus); D—common seal (Phoca vitulina) (figures by K.K. Chapskii):

1—anterior lower corner (or process).

25 Fig. 9. Skeleton of a seal (figure by N.S. Kondakov).

23

very thick, and directed downward. The neck is short but movable and
merges imperceptibly into a fairly clumsy trunk. The skin is thick, rough,
and forms numerous folds and wrinkles. The hair coat is sparse and
bristly; large portions of the body are naked in adults.

The fore flippers are slightly larger than the hind ones, terminating
in a frill of skin devoid of hair; the frill extends beyond the margin of
the digital phalanges. The first digit (inner) is the longest and the rest
shorter in the order of first to fifth. The hind flippers have a naked foot;
the fifth digit is the longest; the first is almost equal to it. The claws on
the flippers are weakly developed and disposed away from the margin
of the flippers. The hind flippers can bend under the trunk and assist in
movement on land (Fig. 10).

There is one pair of teats. The testes are disposed under a layer of
skin and fat (subintegumental); a scrotum is absent.

The skull is massive and broad with an extremely massive elevated
frontal portion, by which it differs from the skulls of all other Pinni-
pedia (Fig. 11). The skull bones are highly massive. The width of the
skull above the canines is almost equal to the width above the external
auditory meatus. A supraorbital process is absent. The hind margin of
the nasals forms an almost straight line. The orbits are relatively small,
their transverse width equal to 1/3 or 1/2 the width of the palate in the
line of molars. The bony palate is flexed inward like a boat, forming
at the back a very gentle arc turned forward with a bulge. The exter-
nal auditory meatuses are very small and their lower walls thickened.

Z Ye y
oe

Fig. 10. Fore and hind limbs of the walrus (figure by N.N. Kondakov).

27

24

Fig. 11. Skull of ап adult walrus, Odobenus rosmarus (figure by М.М. Kondakov).

The tympanic bullae are relatively small and flattened. The lower lat-
eral angle of the temporal bone bears a massive projection. This bony
mass exceeds the height of the tympanic bulla by.a few times. The ante-
rior section of the lower jaw is very massive and its two halves are
firmly fused. The dental formula of permanent teeth in most walruses
is:
2—2 1-1 1-1 3—3
ол. oS

The front incisors are small or altogether reduced, while the last
pair of incisors is indistinguishable from the molars in size and shape.
The upper canines are massive, up to 80 cm long, and directed vertically
downward. The molars are massive, with a single root, their cusps sloping
backward. In older animals the molars become worn and flat, and even
have a slightly concave surface.

The scapula is relatively long, without a perceptible arcuate notch
on the hind margin. Its crest is comparatively elongated, reaching the
hind margin. The os penis is very long, 508-512 mm, slightly S-shaped,
its posterior end broadened and terminating in a head; the anterior end
is obliquely truncated.

Paired air sacs are formed by a projection of the upper section of
the esophagus and the broad openings joined with it. There are no
closing valves. Each sac can hold up to 50 liters of water [air]. The

26

25

И
Ум Wi LENS

Wb

Fig. 12. Skull of a yearling walrus, Odobenus rosmarus (figure by N.N. Kondakov).

air-filled sacs inflate and, spreading under the skin of the neck, hold it
up; their ends lie between the scapulae (Sleptsov, 1940; Nikulin, 1941;
Fay, 1960*).

_ Walruses mainly inhabit the coastal shallow waters of arctic seas and
feed on benthic invertebrates. A large part of their life is associated with
ice floes.

Walruses are distributed only in the Northern hemisphere, in the
circumpolar region, with small interruptions. They inhabit the Bering,
Chukchi, East Siberian, Laptev, Kara, and Barents seas and the waters of
the Canadian archipelago in the northwestern part of the North Atlantic
Ocean.

Walruses constitute one of the three families of the order Pinni-
pedia, but the taxonomic position of the family has not yet been clearly
established. Some authors regard it as a subfamily of the family Otariidae
(Smirnov, 1935; Romer, 1939; and others), while many separate it into
an independent family. The latter view is more prevalent.

The relative proximity of the family of walruses to eared seals
(Otariidae) is, however, beyond doubt. This fact is particularly
emphasized by combining them into one superfamily, Otarioidea
Smirnov, and contrasting them in such a combined form with the
superfamily Phocoidea Smirnov, comprising only the family of true seals.

26

In origin, the walrus family is closely related to the family of eared
seals (Otariidae) and could be regarded as its derivative. Thus the ear-
liest and most primitive known form of the walrus family, Prorosmarus
alleni, from the Upper Miocene of the Atlantic coast of North America
bears some features of the skull structure and dentition characteristic
of Otariidae. In particular, the lower canines are preserved in this form
(Tenius and Gofer, 1960*).

Of the four genera of the family, only one is Recent. Apart from
Prorosmarus, two other genera, i.e., Trichechodon and Alachtherium, are
known from the Middle Pliocene and Pleistocene of Europe. The only
present-day genus, Odobenus, is known from the Pleistocene of North
America and Europe. Extinct as well as extant genera are known only
from the arctic seas of the European and American continents, i.e., the
present-day range of the family. The North Atlantic Ocean could perhaps
be regarded as the center of origin of this family.

The economic importance of this family is presently low, since the
walrus population has been greatly depleted in recent decades.

The family is represented by one genus, Odobenus Brisson, 1762,
with a single species, O. rosmarus Linnaeus, 1758, widely distributed in
the waters of the USSR. (V.A.)

Genus of Walruses
Genus Qdobenus Brisson, 1762

1762. Odobenus. Brisson. Regnum animale. Ed. 2, p. 30. Phoca rosmarus
) Linnaeus.
1766. Trichechus. Linnaeus. Syst. Nat., ed. XII, I, p. 49. Nec Linnaeus
1758 (pertains to manatee Trichechus manatus Linnaeus, 1758).
1772. Rosmarus. Brinnich. Zoologiae fundamenta, р. 34. Phoca ros-
marus Linnaeus, 1758. (V.H.)
See description of the family.

WALRUS

Odobenus rosmarus (Linnaeus, 1758)

1758. Phoca rosmarus. Linnaeus. Syst. nat. Ed. X, I, p. 38, North Atlantic
Ocean.

1811. Trichechus arcticus. Pallas. Zoogr. rosso-asiatica, I, p. 269, Novaya
Zemlya (“Frequens in Oceano arctico ... Copiosissimi in Insula
Navaja Zemla.” (V.H.)

29

27

1815. Trichechus divergens. Illiger. АБВ. Acad. Wiss. Berlin, 1804—II,
р: 68. 35 miles south of Ici Cape, Alaska (162° W long. and 70° М
lat.), Chukchi Sea. (V.H.)

1815. Trichechus obesus. Illiger. Ibid., p. 64, Nom. nud.

1831. Trichechus сооки. Fremery. Bijdrag. Nat. Vetensk, 6, р. 385. Ici
Cape zone in Alaska (Chukchi Sea, 70° М lat. and 163° 18’ W
long.). |

1922. Trichechus orientalis. Dybowski. Arch. Tow. Nauk. Lwow.., I, р. 351,
Nom. nud.

1940. Odobenus rosmarus laptevi. Chapskij. “Problemy Arktiki” (Prob-
lems of the Arctic), No. 6, p. 94. Laptev Sea. (V.H.)

Diagnosis
Only species of the genus.
Description

In general form, the walrus differs considerably from all other species
of Pinnipedia (Fig. 13). The body is large and massive, the skin thick,
covered with wrinkles and folds, and the limbs broad (Fig. 13). Movement
on land is slow and cumbersome and the animal utilizes all four limbs,
but in water it is quite agile and moves fairly fast. It crawls onto ice floes
with difficulty, using its tusks and fore flippers.

The head is relatively small with a massive snout that is blunt in
front. The eyes are small and shifted far back (Fig. 14). Long (up to
10-12 cm), numerous, very hard and thick whiskers occur on the front
part of the snout; they are directed downward and arranged in 13-14
rows. Each side of the snout bears 300-350 whiskers. The whiskers in
the middle part of the snout are usually very worn and measure hardly
0.5 to 1 cm.

Both tusks in the upper jaw are very long and directed vertically
downward. Instances are known of walruses with more than one pair of
tusks. A walrus caught around 1915 had two well-developed tusks on
each side, and all four were of almost normal length. Those on the right
side grew parallel to each other while those on the left were twisted
(Caldwell, 1964). Four skulls have been described with three tusks and
five with five tusks each. A skull was found in which there was a single
normally developed tusk on the left and five separate formations on the
right side consisting of fragments of small tusks of normal shape, two
dentine stubs 12 and 14 cm long rising from the surface of the gums,
and two dentine concretions 12 and 20 mm in diameter concealed in the
jawbone (Bel’kovich and Yablokov, 1960).

29 Fig. 14. Front view of a walrus head (figure by М.М. Kondakov).

29

The color of old walruses is a dirty olive on the back and rusty-brown
on the belly. This coloration is caused by the color of the hair, as also
the skin pigmentation, which is brownish. The tips of the flippers, devoid
of hair, are similar in coloration.

Sexual and age-related dimorphism are manifest in the body dimen-
sions, shape and size of tusks, and hair coat. Adult males are about 0.5 m
longer than females; the male skull is more massive and the tusks longer
and thicker than in the female. The tips of the tusks diverge sideways in
the male but in the female are somewhat proximate with a slightly spiral
curvature. The female, usually darker, attains maturity one to two years
earlier than the male. Pups of both sexes are identical in size, but the
female lags behind in growth during the period of sexual maturity and
ceases to grow altogether soon thereafter; cessation of growth sets in
later in the male. A clear example of sexual dimorphism is the presence
of large wartlike formations on the breast and shoulders of males, which
are absent in mature females.

The hair coat of young walruses is fairly dense and dark brown. It
becomes bristly, sparse, and brownish-yellow with age, with large bald
patches. Evidently the hair color undergoes no seasonal variations. (For
skull description, see under characteristics of the family; body and skull
sizes are given under “Geographic Variation”.) The weight of an adult
male can reach almost 1,500 kg and that of an adult female 800-900 kg.
The average weight of the visceral organs (Seven specimens) is: heart
6,167 g, lungs 14,062 g, liver 29,640 g, spleen 4,146 g, stomach 5,312 g,
intestine 33,640 р, kidneys 3,544 р, and pancreas 2,148 р. (V.A.)

Geographic Distribution

Arctic seas of the Atlantic and Pacific Oceans.
Geographic Range in the USSR (Reconstructed)

In the early twentieth century, walruses were probably regular inhab-
itants of the northern White Sea (Morzhovets Island). In the Bering
Sea during the nineteenth century, walruses bred regularly on Karaginsk
Island (59° М lat.) and in the 1880s were caught in thousands every
year. Walruses were reported on Cape Kronotskii (56° М lat.), Cape
Shipunskii (53° N lat.), and on the Commander Islands (Grebnitskii,
1902; Suvorov, 1914; Arsen’ev, 1928*; Ognev, 1935; N. Smirnov, 1935;
Nikulin, 1941) (Fig. 15).

Direct references are available to the breeding of walruses in the
northern part of the Sea of Okhotsk. The reports of a Yakutian army
commander in 1651 refer to the hunting possibilities of walruses in the

Зоиихо MOU SI зпем э41 YOY ит $10131 Dy) $9007 рэцозец pue э8ие1 рэ12пл11510991
ay) Jo Азерипод ay) зэюцэр aut] рноб -(aa,uasTy 7.) 31550] Э41 Ul $тириибол пиздоро ‘зпцем эчз JO чоппана ‘ст i) fe Te
; 7 et 06 OL

=F у, [ el В.
Sey RS
ay №. а |

ut \ Ps |
О
=} у

re)
д д
К {3
LB) eS “a
75% a
eo &
é is
Bet Fy
>,
к
$ ==) WZ а

к к К
re

31

Sea of Okhotsk. In the record for 1652, he reported sighting many
walruses on the beach of Cape Morzhov for two or more “versts”
[1 verst = 1.067 km] (Akinfov, 1848). These data pertain to the
northern part of the Sea of Okhotsk not far from the present Magadan.
(V.A.) The veracity of this report was confirmed by a reference to the
possibility of finding “fish teeth,” i.e., walrus tusks, at these places.
This is also supported by the find of tusks in the coastal rock mounds
along the northern coast of the Sea of Okhotsk, and the sighting of live
walruses. A young walrus was found around 1890 on Yamsk Island in the
northeastern part of the Sea of Okhotsk and a pair of tusks was found
around 1900 on the coastal dumps of Shelikhov Strait. Walrus tusks were
also found in Nogaev Bay (B.A. Zenkovich).

The present-day distribution of walruses in the USSR forms but a
small part of the range which prevailed in the past. Walruses inhabit
the waters of Franz Josef Land, Novaya Zemlya, Barents and Kara
seas, Severnaya Zemlya islands (more often on the eastern coast of
the archipelago), Vil’kitsk Strait, and in the shallow waters of the
Ob’-Yenisey. They inhabit the Laptev Sea (mostly its western part
close to the eastern Taimyr coast), Lena delta, landlocked waters of
the Novosibirsk archipelago (mostly its northwestern fringe), and are
encountered in the western part of the East Siberian Sea, mainly
in the region of Novosibirsk Islands and De Long Islands. Farther
east, they are found in the Chukchi Sea from De Long Strait in the
west to Wrangel Island in the north, Bering Strait and Anadyr Strait
to Cape Navarin (Chapskii, 1936, 1939, 1940, 1941, 1963; Tsalkin,
1937; Belopol’skii, 1939; L.N. Popov, 1939; Rutilevskii, 1939; Nikulin,
1941; Vinogradov, 1949; Zakharov, 1958; L.A. Popov, 1958, 1959,
1960).

Geographic Range outside the USSR (Reconstructed)

The southern boundary of walrus distribution in the Atlantic Ocean out-
side the waters of the USSR has also varied considerably. Judging from
the finds of walrus remains in excavations, this animal penetrated far
southward at one time. In the nineteenth century, its remains were found
in Denmark, England, France, and on the east coast of North America in
New Jersey, Virginia, and Carolina (Moor, 1952*). Instances are known
of the discovery of walrus remains in Maine and Massachusetts. More-
over, fragments of the skull and other bones were found in the Gulf of
Maine (G.M. Allen, 1930; Palmer, 1944). In the first half of the sev-
enteenth century, walrus hunting prevailed on Sable Island (44° N lat.),
close to the Canadian coast north of the Gulf of Maine (G.M. Allen,
1930) and on the Magdalen Islands (Mansfield, 1959).

33

32

In the middle of the nineteenth century, stray walruses were caught
on the Shetland and Orkney islands (Moor, 1952*). At the end of the
nineteenth century, walruses were perhaps permanent inhabitants on the
coast of Finmarken in northern Norway, found on the coasts of Scotland,
and in the Gulf of St. Lawrence. In general, at the end of the last century,
walruses were widely distributed in the northern part of the Atlantic
Ocean and in the Arctic Sea.

It has been assumed that in the eastern part of the Pacific Ocean,
walruses penetrated south of the Aleutian Islands, reaching Shumagin
Island and even the Alexander archipelago in the eastern part of the
Gulf of Alaska.

_ In the middle of this century, walruses were known to inhabit the
northern part of the Atlantic Ocean, being recorded in the waters of the
Canadian archipelago (Southampton and Devon islands, Fox Basin, and
Baffin Bay), the Labrador coast, Davis Strait, the west and east coasts
of Greenland, Spitsbergen, and were sometimes spotted near Iceland
(Fig. 16).

In the Pacific Ocean, walruses inhabit the eastern part of the Bering
Sea, from Bristol Bay in the south to the Bering Strait in the north (includ-
ing the St. Lawrence Islands, Nunivak, and St. Matthew), and extend along
the north coast of Alaska into the Chukchi Sea up to Cape Barrow.

The northern boundary of the range of walruses in the Barents and
Kara seas runs beyond 80° N lat. In April, 1957, a walrus was noticed in
the breeding grounds on Franz Josef Land north of 81° N lat. (Vaigachev,
1958). In the waters of the Pacific Basin, the area 72 to 74° N lat. can
be regarded as the regular northern boundary of the range of walruses,
but the animals can be found even more northward in favorable years,
depending on the situation of the ice floes. (V.A.)

Geographic Variation

Usually, three subspecies are recognized. All of them inhabit the waters
of the USSR.

1. Atlantic walrus (O. r. rosmarus (Linnaeus, 1758)) (syn. arcticus, ? obe-
sus).

This is the smallest form.

The maximum body length of males is 375 cm, of females 338 cm;
the corresponding averages are 345 and 293 cm respectively (Chapskii,
1963). The condylobasal length of skull in males is 256-379 mm @ =
369), in females 303-342 mm (х = 314); the maximum width of skull in
males is 268-291 mm, in females 222-257 mm (х = 234.2). The length
of male tusks along the curvature from the edge of the alveolus to the

33

$11550} зпем JO SPUlJ JO $215—ф ‘5э5ПИЕМ Jo $риц Аер-1и2$5924 Jo $213—$ Чоицхо MOU ‘эле зэ5пем
ЭША Ul $401891—2 ‘э8иел рэ12п11$1099эл1 Jo Алерипоа— т ‘(лэ.иэзгу “W'A) Smipuisos хпизаоро ‘зпцем ay} JO понпанзта ‘9т “314

С5

34

34

tip is 34-38 cm, in females 27-33 cm (Ognev, 1935). The tusk tenet in
one male was 52.5 cm (Chapskii, 1963).

In the USSR it inhabits the Barents and Kara seas; outside the
USSR, the North Atlantic Ocean to the Canadian archipelago in the
west, inclusive.

2. Laptev walrus (O. r. laptevi Chapsky, 1940).

Somewhat larger than the Atlantic form.

The maximum body length of males is 410 cm, of females 370 cm;
maximum length of the tusk is 65 cm in males, in females 58 cm. The
maximum circumference of the tusk in males is 21 cm, in females 14 cm
(L.A. Popov, 1960).

This form inhabits the Laptev Sea, the western part of the East
Siberian Sea, the Lena Delta, and the Novosibirsk archipelago. It is more
numerous near the coast of eastern Taimyr, where it is mainly confined
to the coastal region and the shallow waters; it is rare in the western
part of the East Siberian Sea in the region bordering the Laptev Sea.

Not found outside the USSR waters.

3. Pacific walrus (О. г. divergens (Illiger, 1815)) (syn. cookii, ? orientalis).

Largest form of the species.

The maximum body length of males is 450 cm (х = 336), of females
367 cm (х = 283). The skull is more massive than in other forms and
the frontal section considerably broader. The condylobasal length of the
skull in males is 383-428 mm (x = 396.8), in females 315-357 mm (x =
332); maximum width of the skull in males is 290-333 mm (х = 309),
in females 219-265 mm (x = 245). The length of the tusk in males is
46-80 cm, in females 40-60 cm; its width in males is 66-84 mm, in
females 41-53 mm (Ognev, 1935; Nikulin, 1941; Freiman, 1941).

It inhabits the Bering, Chukchi, and eastern part of the East Siberian
seas. Some contacts between the Pacific, Laptev, and East Siberian Sea
walruses are possible.

Outside the USSR, it is found in American waters of the Bering and
Chukchi seas. (V.A.)

Biology

Population. As a result of prolonged hunting, the walrus population has
declined steeply throughout its range.

The population of the Atlantic walrus has suffered the most and
only a few stray herds are now known. The walruses of the Kara
Sea suffered the highest destruction and those remaining now do not
exceed a few thousand. Small groups are confined to Franz Josef Land

35

34

35

and Spitsbergen. The population in the eastern part of the Canadian
archipelago is split into a few groups. In the northern part of Hudson
Bay and in the region of Southampton and Cox, some 3,000 are known
(Mansfield, 1959). Population figures for other regions are not available.

The population of the Laptev Sea walruses has been less affected by
exploitation, as it has always been relatively small. Nevertheless, hunting
has had its impact here as well, and the present population barely exceeds
5,000 - 6,000.

The Pacific walrus populations are best protected. Aerovisual
estimates, aerial photographs, and studies of coastal breeding grounds
and rookeries on the icy coasts revealed a summer population of
30,000 - 35,000 in the USSR. According to American and Canadian
authorities, some 15,000 walruses inhabit the US waters in summer. The
total population thus is 50,000 (Fay, 1957; Fedoseev, 1962; Krylov, 1968;
Gol’tsev, 1968).

Habitat. An outstanding feature of the habitat is the relatively shal-
low water and abundance of benthic mollusks and partly of crustaceans.
Drifting ice is common in these sections and, in the winter months, wal-
ruses inhabit only the ice (Fig. 18). In the absence of ice in summer,
walruses form coastal rookeries mainly on sandy or pebbly shoals on the
coasts of the continent or islands.

Food. There is no information on the winter food of walruses, nor
on seasonal food variations. Our knowledge is limited to a list of animals
retrieved from the stomach of walruses in the summer months.

Fig. 17. A group of walruses оп an ice floe. Chukchi Sea (photograph by
V.M. Bel’kovich).

36

Fig. 18. Adult females and young walruses (left). Chukchi Sea (photograph by
V.I. Krylov).

In the first two years juveniles survive on the mother’s milk and take
to independent feeding only in the third year. By then, the tusks have
grown sufficiently large to enable the walrus to independently scrape
food from the bottom of the sea. The main food items of adults comprise
bottom-dwelling invertebrates (Table 1).

Throughout its extensive range, the walrus primarily feeds on various
mollusks, which in the species composition of its food occupy first place.
These are followed by crustaceans, of almost equal dietary importance.
All the other food items, i.e., worms, echinoderms, ascidians, and fish,
can be regarded as secondary. In the stomach of an Atlantic walrus, over
a hundred polar cod were found (Chapskii, 1936); fish remnants were few
in the stomach of Pacific walruses (V.I. Krylov); only remnants resem-
bling otoliths were found in the stomach of walruses in the Canadian
archipelago (Mansfield, 1958). In addition to the food items listed, the
stomach of one walrus (site of find not mentioned) contained Tridacna
(Moor, 1952*). Remnants of pinnipeds and even whales were found in
the stomach of some walruses in all the regions studied. |

The walrus seeks its food from the sea floor. It is assumed that it digs
the bottom with its tusks, selects mollusks, breaks the shells, and eats
the molluskan bodies. However, there is a suggestion (Mansfield, 1959)
that the walrus bites only the soft, protruding portion of the mollusk,
based on the fact that shells are very rare even in the stomach of those
walruses caught immediately after feeding.

и)

эершиэ1Я эерип8еа
Ч epodesaq
snomuofy09
шпциолто шт 5740120 ‘ds ssyonq
0128102 vipuojag sndjog ‘ds suvsojuonds
(ати И)
зиетрюзу эерчоол1, ‘ds м08и042042]25 ppivs snpv30a10g
pupunony snuoposkiy) uakosy snansodnyq ysl
зилэрочщоя шпи12тЯ 1190$ vaylopisayy 511912018 Виши"
5ища2М 0]]2и5]0$ suesoejsnic suLIopoulyoYy
p8ajkyouo2 5щапио auvIspy (syeaq) spinbs трио хптари4
15405 2и9РИ DAIDIN в[п91э40 SULIOM,
snuniysa sniniyoq vsonxayf vasvky J порто) snindvug еродол1$е9) эериешиео
5торпо) snindoug DSNDID DION snaniyoq еродАээ1эа (вэщлор152И!) еродо$]
SULIOM, sniua] DININNT ИНО ‘ds oydig УРАН
5порир]изо48 5711020]
эерпАюааН 5244425 тире ‘ds vouvoN UOSUDIIOLI IS
snjupuvd 02402]02 DIUOID $1229е1$1215) ‘ds wnuizong suesoejsnic
aepisnyl 0240442А4ц vrpjoy 021240 DADIDOS ‘ds и21224 DOUDN
эерыешшео) 51102404 эт 5 viunidan ‘dds шифр) unIpsvgy
40] и08и0420122М psosni DADIDOS шпи12тЯ 51102404 2и05У DINIAD DADIDOS
57117124002 5РАН 9211240 DADIDOS vasoqiaddy v1pjoy 021240 IADIDOS wnulzong
ojido 52122200) pivouny vi И piwouny ОИ DA
$41229Е15110) sysny[oy| ЗПП УПО $$

Sg ae ese ee р АЕ ВЕСИ ЗИ ИВА ЕР a

(З©6Т ‘Pleysuey) (9¢61 ‘Wwasdeyo)
(AOA ГА {TPO ‘UHNOIN {8261 ‘A2,UasTV “V'A) (6561 рие 8661 ‘лодоа ‘v'T) osejodiyore ueIpeue) Bag еле
зпем ole snijem bas лэ14е] snajem эциеп\у

pec AO ie Е

SUO{SII пошел UT 5эзащелм JO poo `Т э14вт,

9€

38

РЕВВА Е Е
ystj Jo
$43010 ‘eaoeIplosy
вэрюлаЧ19 юн
‘ds wntpiuoky],
‘ds pupuimonD
зилэрошоЯ
eyaeyoAjog
snjupnos snjndvug
ОМ
еродо$]
эершэдши7] эериАюдан

(лоИтЯ ТА ‘ТУбТ ‘ЧЧахим {8261 ‘лэмэзгу УД) (6561 рие 856т ‘лодоа `У“Т) (SS61 ‘реузие у) (9Ебт ‘иузаечо)
зпием oloeg эпем bas лэ34е] osejadiyose иетреце) B9S LILY
зпем эциепу

рэпиио) Т 71481

37

37

39

The walrus gathers food where it can easily reach, mainly at depths
of 30 to 50 m (Nikulin, 1941). The depth to which the walrus dives while
feeding can be as much as 180 m (Mansfield, 1959).

Some walruses prey on seals and, occasionally, on birds. Seals usu-
ally avoid the area inhabited by a walrus-predator, but will return there
if the predator is killed (V.A. Arsen’ev, 1935). Two categories of preda-
tory walruses are known: the first group feeds on uncommon foods occa-
sionally when invertebrates are not available for some reason, while the
second category comprises regular predators. The latter have long, thin,
sharp-tipped tusks; these are lone males (Fay, 1955*).

Home range. There are no separate sections for individual walruses
or even for groups of them. In the absence of ice in summer, walruses
form coastal rookeries; however, it is still not clear whether the same
animals return to the same sections. At the rookeries in the Laptev Sea,
the same walruses evidently return to their own sections (L.A. Popov,
1958).

Daily activity and behavior. No systematic diurnal activity has been
established among walruses.

The Atlantic walruses of all ages and sexes gather on the beaches

_in August and September since there are almost no ice floes in the

USSR waters in the regions of their summer habitat. There they remain,
sometimes for 1.5 months, often going into the sea to feed. The animals
driven away from the shore by stormy waves return to the beach as soon
as the weather clears.

Fig. 19. A male walrus with a broken tusk. Chukchi Peninsula (photograph by
V.I. Krylov).

39

37

40

The Laptev walruses also form mixed coastal rookeries in which
females with pups (Fig. 20) and juveniles live along with the large
males. They lie in a definite sequence—females with pups close to
the water, juveniles and the remaining adults farther away. They rest
crowded together compactly in various postures, but mostly on the side.
Yearlings and juveniles sometimes lie atop the adults. Fights are quite
common between large bulls, who put their powerful tusks to good use.
At all times, some animals are moving toward the water while some are
returning to the rookery; thus the rookery is never quiescent (L.A. Popov,
1958). In September and October, the coastal herds disperse and the
coast remains empty until the next season.

The Pacific walruses spend much of their time on ice floes. Two types
of colonies are distinguishable. One consists exclusively of adult males
and the other of females with pups, among which a small number of
males are sometimes observed. In calm weather walruses sleep soundly.
The resting animal does not even deign to look at a ship approaching the
ice floe and, when disturbed, takes to water rather reluctantly. Walrus
herds have often been observed resting on ice floes within 100 т of
freshly killed animals. They did not react even to gunshots, the clatter
of operating winches on a ship, nor to any other noise. Yet, in general,
their hearing is better than their sight.

Fig. 20. Suckling walrus pup on an ice floe. Chukchi Sea (photograph by
V.I. Krylov).

41

Exclusively male haunts are usually small but mixed herds run into
several hundreds. The rookeries are mostly organized along the edges of
drifting ice floes. The animals rest on permanent floes, whether smooth
or hummocky. Frightened animals literally dive into the water, but soon
surface, gather in herds, and begin groaning loudly. The herding instinct
is strong among walruses (Dunbar, 1955). An injured animal is helped
by others to float on the water surface. Pups that tire of swimming, often
climb atop their mothers or any other walrus.

In autumn the Pacific walruses form coastal rookeries in some other
regions as well (Fig. 21); these may be permanent or temporary. The
latter can be classified as regular or occasional.

Only two permanent rookeries are preserved at present: at Rudder
and Meechken in the Gulf of Anadyr. Only males, mainly 7 to 11 years
of age, gather here (Fig. 21). Juveniles (mainly 4-6 years old) and older
males come to the beach in small numbers. Females of all ages remain
confined to ice floes during autumn. A few thousand walruses gather in
each rookery, lie packed close together, often in two tiers. The more
mobile young walruses, lying closer to the water, leave the shore more

38

Fig. 21. Coastal rookery of male walruses. Chukchi Peninsula (photograph by
P.G. Nikulin).

40

39

42

often than the older males resting farther away; then the older walruses
become less densely crowded and rest for longer intervals and more
peacefully. Permanent rookeries are formed at the end of August and
function until the end of September -October when, with the appear-
ance of ice floes, the walruses depart for them. During the period of
walrus abundance on the Chukchi Peninsula there were many permanent
rookeries, but now almost all of them are deserted.

Temporary rookeries serve as resting sites for walruses migrating
from the Chukchi Sea to the Bering Sea. Such rookeries are visited
by animals of both sexes and of all ages, including females with pups.
The period of their formation and the population of animals depend
on the ice conditions each year. Sometimes temporary rookeries contain
1,000 or more walruses, but animals may leave within 2-3 to 7-10 days.
Regular temporary rookeries are formed every year at the same places.
Some 15 such were counted in recent years on Arakamchechen Island,
in the region of villages Dezhnev and Ue’len, on Capes Intsov and
Serdtse-Kamen’, on Idlidlya and Kolyuchin islands, at village Vankarem
on Karpkarpka island, and a few on Wrangel and other islands.

Finally, occasional rookeries are formed in the warm, less snowy
years at the most unexpected places (sometimes even in villages). Wal-
ruses, tired of long sojourns in the water, form such rookeries, which

Fig. 22. Marking of a walrus. Chukchi Peninsula (photograph by V.I. Krylov).

40

Fig. 23. Marked male walrus. Chukchi Peninsula (photograph by V.I. Krylov).

are extremely short in duration. Having rested on the coast, the animals
quickly return to the sea, continuing their migration.

On the coasts of the Chukchi and Bering seas, over 30 permanent
or temporary rookeries have been recorded in various years. At present,
15 have completely vanished, with no walrus recorded in them in recent
decades (Gol'’tsev, 1968).

Walruses have often been observed sleeping on the water. Having
filled its air sacs, the animal assumes a vertical posture with its head and
the blown-up sacs of the neck jutting out of the water, and sleeps in that
position (Nikulin, 1941).

Seasonal migrations and transgressions. Walruses perform regular
seasonal migrations (Fig. 24) but the details of such migrations are not
very clear.

The Atlantic walruses inhabiting our waters spend the winter-spring
months in the southeastern part of the Barents Sea. In October, in the
Kara Strait region, coastal rookeries are formed, but with formation of

9 In 1963, 500 Pacific walruses were marked for the first time in the Rudder rookery
using a specially designed tag shaped like a large button with a base diameter of 3 cm and
stem length of 5 cm. The tag was attached to a long pole and then jabbed into the skin
of the walrus (Krylov, 1965). The animals permitted the markers to approach within the
requisite distance (Krylov, 1965). No tags have been returned thus far (1967).

41

41

44

Fig. 24. Migrations of the Pacific walrus, Odobenus rosmarus divergens (V.A. Arsen’ev).

the ice cover, the animals return to the ice. In June, as the ice floes begin
to drift, the walruses usually abandon this region and enter the Kara Sea
by two routes: through the Kara Strait or by encircling Novaya Zemlya
from the north.

The animals passing through the Kara Strait in the second half of
July and in August inhabit the ice along the western coast of Yamal
Peninsula from Belyi Island to Sharapov Spit. The easterly winds of
August drive the ice away from here and the walruses migrate with it. By
early October these animals are seen on the southern coasts of Novaya
Zemlya close to the Kara Inlet, where they form coastal rookeries that
function until formation of the ice cover.

Another group of walruses, together with the ice floes, migrates to
the northern extremity of Novaya Zemlya and spends July-August on
ice floes in the coastal waters. By September, as the drifting ice moves
away from the Novaya Zemlya coasts, the animals begin to congregate
in the immediate proximity of the coasts, mostly on the northeastern
extremity of the island, and to form beach rookeries in a section from
the Oransk Islands to Cape Sporyi Navolok. By October, ice has usually

42

45

begun to form afresh on the northern coasts of Novaya Zemlya and the
walruses abandon the coastal waters and move onto the drifting ice to
reach the Barents Sea. They move along the western coasts of Novaya
Zemlya and by the end of the month have reached their winter habitat
in the southeastern part of the Barents Sea (Chapskii, 1936).

The migrations of the Laptev walruses have not been studied. In
August-September they form coastal rookeries, mainly on the south-
western strip of the sea, on Faddei, Andrei, Preobrazhen’e, Begichev,
the Peschan Islands, and in Pronchishcheva Bay. In the autumn, as the
ice floes appear, the walruses move onto them and migrate northward,
spending the winter in ponds of open water in the ice and in cracks fairly
close to the coastal rookeries. In August, 1951, some 400 walruses were
sighted at 87° N lat. and 140° E long. (Uspenskii, 1958), which perhaps
could be regarded as having strayed beyond the range.

The Pacific walruses spend the winter in the shallow waters of the
southeastern part of the Bering Sea, reaching the area of compact ice
floes in Bristol Bay. In March-April, they begin moving northward on
the ice floes, approaching the coasts of Chukchi Peninsula in the Provi-
deniya Bay-Cape Chaplin region. In May, walruses are seen even in the -
Bering Strait, and by June have emerged, together with the floes, into
the Chukchi Sea, reaching Wrangel Island. During July, however, walrus
herds continue to move through the Bering Strait and spread out in their
summer grounds, from the eastern part of the East Siberian Sea to Point
Barrow in Alaska, only in August.

In mid-October, when intense formation of new ice occurs and polar
ice starts to drift southward, the walruses move to the Chukchi coasts.
They swim mostly through clear waters for a distance of 50-100 km from
the coasts to the Bering Strait and, entering the Bering Sea, proceed to
their winter grounds. Large walrus herds swimming to the Bering Strait,
even among compact ice floes, have been sighted in October, evidently
struggling to escape the fast-freezing sea (P.G. Nikulin).

Some walruses remain in the Gulf of Anadyr in summer on the ice
floes of Cape Bering and Krest Bay until the ice has completely cleared,
after which they remain in clear water. In July-August, coastal rookeries
are formed in the Gulf of Anadyr (Rudder) and, at this time, some wal-
Tuses move in groups or singly in the clear water along the Chukchi
coasts eastward, forming new rookeries in the Bering Strait (Arakam-
chechen) and even at its confluence with the Chukchi Sea (Inchoun).
They remain there until the end of September or October, after which
the animals move onto the ice. Rookeries are also found on Wrangel
and Herald islands (Belopol’skii, 1939; Nikulin, 1941; Kleinenberg et al.,
1964).

43

46

Walrus finds outside their usual range have been reported for both
the Atlantic and Pacific populations. Almost every year single walruses
are sighted in summer and autumn in the inlet of the White Sea and
in Mezensk Bay. In 1956, a walrus was killed on the eastern Murman
coast (Bel’kovich and Khuzin, 1960). Walruses have been sighted on the
coasts of Norway, mainly in the northern areas. A walrus was found in
1902 and 1903 in G’esver Fjord, in 1904 close to Trondheim, in 1917
around Kirkenes, and in 1931 in Grosbaken Fjord. Two walruses were
sighted in 1942 at Finmarken, and in 1953 one was found lying on the
beach at Makkaur lighthouse (Lund, 1954).

An interesting journey of a walrus along the Norwegian coasts
has been described. In October, 1926, in the southern part of Norway
(Haugesund), a male supposedly from the coasts of Scotland was sighted.
The same male was seen on November 11 on the coast of Holland, at
the end of November on the northern coast of Denmark (near Skaagen),
and finally on January 9, 1927, was killed in Bokhuslan region in Sweden.
An even more amazing journey was performed by another male in 1954
(assuming that it was the same animal throughout). In early January, a
male was noticed at the northernmost tip of Norway in Bakkeby region
and then sighted on Trena Island on February 3, having traveled 600 km
in 26 days. From there, the walrus took to the coast on March 15
and traveled another 700 km south into Batalden region, from where
it entered Swedish waters. It soon returned northward and, on March
27, beached around Sul (about 62° М lat.), then traveled still northward.
In mid-April, it was sighted on Helligwer Islands (67° М lat.), again near
Trena, and then at Lofoten. In mid-June, the male was seen at 69° N lat.
at Sandesundver (Lund, 1954).

The following journeys of Pacific walruses are known. In 1931, a
herd was sighted in Korf Bay (60° N lat.) in Kamchatka, in the summer
of 1935 about 500 of them inhabited Natalii Gulf (61° N lat.), in 1939
they were sighted on Karaginsk Island (59° N lat.) and on Cape Paklan
(59° 40’ N lat.), and in August of the same year they surfaced on the
coast of Verkhoturov Island (59° N lat.) where two of them were killed.
In July, 1969, 20-25 walruses were sighted. Some walrus skulls were
found on the Commander Islands while in the summer of 1969 three
carcasses were found. One of them, a female 3 m long with 60 cm long
tusks, found on June 17, was torn to shreds by polar foxes (some 40 of
them gathered), which lends credence to this animal having been killed
on the coast (Chugunkov, 1970) (Fig. 25).

In May, 1940, close to Yamsk Islands (northeastern part of the
Sea of Okhotsk), not far from a rookery of fur seals, a young female
was killed (without embryo, length of tusks 20 cm) (Moiseev, 1951).

43

47

Fig. 25. The carcass of a female walrus found on the coast of Mednyi Island. А
polar fox is seen on the carcass (photograph by D.I. Chugunkov, 1969).

Possibly, this was one of the pups which hunters often took onboard
ship in the Chukchi Sea. However, in the summer of 1966, on Yamsk
Islands, four adults were again seen (G.A. Fedoseev). In April, 1954, a
large female was killed on Kad’yak Island in the Gulf of Alaska (Fay,
1957).

Reproduction. Information on walrus biology is predominantly col-
lected in the summer months during the hunting season and hence data
on breeding are extremely scant. Data on the Pacific walrus are somewhat
more comprehensive.

The growth of follicles in the ovaries commences in the first half
of April and mature follicles are seen in early May; such follicles are
present in gestating females as well. In the first half of June, the growth
of follicles ceases. In mid-May, ruptured follicles are replaced by corpora
lutea, indicating the fertilization of egg cells.

Mature spermatozoa form in males in early April; by the first ten
days of June spermatogenesis has ceased. The maximum quantity of
mature sperm is observed from mid-April to May end. Thus the mat-
ing of walruses occurs during May and to a lesser extent in early June.
By this time, some females are already with embryos in the early stages
of growth (Fig. 26).

44

44

48

Fig. 26. Embryo of the Pacific walrus, Odobenus rosmarus divergens. Bering Sea
(photograph by V.I. Krylov).

Pups are born in about the same period as mating—from April end
to May end. Individual instances of birth are known in early June as well.
The period of whelping extends for a month or more. Hence gestation
extends for almost 12 months. Whelping duration can be ascertained
from the differences in the sizes of embryos found at a given time. Thus
the length of embryos investigated on July 30 varied from 13 to 21 cm,
on August 30 from 27 to 37 cm, on September 14 from 34 to 47 cm, and
on September 23 from 43 to 53 cm.

The embryos of Pacific walruses measured in the same period were
5-11 cm longer than those of Atlantic walruses and 4-7 cm longer than
those of the Laptev Sea. This suggests either a very early whelping in
the Pacific walrus, or that the much larger size of this subspecies is
determined even during embryonal growth (Krylov, 1966, 1969).

The reproduction tempo of walruses is the slowest among all the
species of pinnipeds and the whelping pattern is highly complex. Among
the Pacific walrus females (285 studied) mothers were seen suckling pups
born in that year and again gestating; suckling yearlings and gestating;
and some gestating but not lactating; i.e., females at different stages
of reproduction were encountered. The following relationships have
been established between groups of females with different reproduction
rhythms (Krylov, 1968):

49

Females, whelping annually 4.5%
Females, whelping once in two years 12.3%
Females, whelping once in three years 42.2%
Females, whelping once in four years 41.0%

The reproduction rhythm of Laptev walruses is similar to this pattern
(L.A. Popov, 1960).

Young females have a more frequent reproduction rhythm com-
pared to older ones. Most females with an annual whelping cycle fall
in the age group 6 to 11 years; those with a three-year cycle from 12
to 18 years; and those whelping once in four years or more are over
15 years of age (Krylov, 1968). The annual population growth is 8%
since the average of pups per mature female is 0.35 (Mansfield, 1959).

45 According to other data, the annual population growth is 11.2% (Krylov,
1968).

' The prevailing view regarding polygamy among walruses (Allen,
1880; Nikulin, 1941; Freiman, 1941) has not been confirmed by recent
investigations. During the period of reproduction walruses do not form
harems but live in family groups of three to six animals comprising the
male, female, and pups of different ages (Tikhomirov, 1964c; Krylov,
1968). Ice is not a suitable substratum for organizing a harem, whether
of walruses or other pinnipeds (М. Smirnov, 1937*).

Growth, development, and molt. Reliable information on the growth
of walruses (and other pinnipeds) became available only after the devel-
opment of a method for determining the age of each animal. Age is deter-
mined from the annual depositions (rings) in the dentine of the teeth
(Fig. 27). This method has been verified in many species of marine and
land animals and is generally accepted (Tikhomirov and Klevezal’, 1964*).

Differences in the tempo of growth of males and females are percep-
tible already in the juvenile and persist throughout their lives (Table 2).

Pups of both sexes grow very fast. The tempo of growth slows down

46 at two years of age, evidently due to a changeover from suckling to inde-
pendent feeding. Again, a reduced tempo is seen in females of 7-9 years
and males of 8-9 years. This lag coincides with the period of sexual
maturity, which in females begins at 6-8 years with the peak in the
7th year, and in males at 7-9 years with the peak in the 8th year. This
is followed by a fairly uniform increase in body length in females up to
11-14 years and in males up to 17-20 years. Very slow growth continues
up to 20 years in females and up to 23 years in males; thereafter body
growth ceases. Females older than 25 years enter a climacteric period
and old males also evince no interest in mating. Among females whose

45

50

Fig. 27. Annual layers on a polished section of а cheek tooth of the Pacific walrus,
Odobenus rosmarus divergens (photograph by V.I. Krylov).

age was determined, the oldest was 30 years, and among males 43 years
(Krylov, 1966, 1967, 1968).

The body growth curve of walruses of the Canadian archipelago is
very similar. Here, by the beginning of the third year, the body length
of walruses averages 2 m and weight 340 kg; the average length of adult
females is 2.6 m and weight about 505 kg; the corresponding values for
males are 3 m and 750 kg. The maximum weight of females is 725 kg
and of males 1,270 kg. Females cease to grow in the 15th and males in
the 20th year (Mansfield, 1959).

Simultaneous with increasing body length the tusks grow in length
and thickness. The newborn pup has no teeth and the canines begin to
cut a few months later. By the end of the first year, the canine mea-
sures 2.6 cm (average of five measurements; V.I. Krylov) or 2-9 cm
(Mansfield, 1959). Slowing down of the tempo of tusk growth is com-
mensurate with that of body length. Tusks grow throughout the animal’s
life but detecting their growth in adults is impossible because they wear
down constantly, thus decreasing in length. The tips of male tusks diverge
sideways, while those of females are slightly proximate. This difference is
distinctly visible in the frontal view. Female tusks are thinner and some-
what crescent-shaped, while those of the male are stronger and almost

Straight.

The sex ratio in newborns is close to 1:1. Suckling extends for about
two years but the stomach of pups older than a year often contains
mollusks along with milk; the stomach of those older than two years

51

45 Table 2. Change of body length with age in the pacific walrus (V.I. Krylov)
Age Male Female
No. of Mean No. of Mean
animals length animals length
Newborn 4 138 8 129
1-4 months 3 158 5 146
Year-old 5 198 11 185
2 years 4 238 5) 218
3 years 6 247 3 238
4 years й 260 7 258
5 years 11 281 11 268
6 years 13 292 6 276
7 years 24 297 И) 276
8 уеагз 25 306 6 278
9 years 30 306 11 286
10 уеагз 49 322 13 290
11 years 32 337 8 307
12 years 45 338 18 308
13 years 24 338 8 298
14 years 30 352 17 307.
15 years 35 354 25 299
16 years 43 351 18 308
17 years 34 358 22 304
18 years 35 358 15 312
19 уеагз 31 362 8 315
20 years 29 370 10 311
21 уеагз 19 371 8 312
22 years 12 370 4 328
23 years 17 375 7 326
24 years 14 371 8 321
25 years 14 369 4 321
26 years 13 372 4 321
27 years 4 367
28 years 12 370
29 years 7 371
30 years 12 363
31-32 years 6 370
33 years 5 372
34-38 years 7 374

sometimes contains milk. Females under parturition every year suckle
two pups simultaneously—the newborn and the yearling.

The first molt occurs soon after birth. In the first few days the body
of the pup is covered rather densely with grayish-brown hairs, which are
gradually shed and become sparse 1-2 months later. Adults have large
bald patches and the hairs are small and sparse in other places; however,

52

they molt every year. The period and duration of той have not been
established but evidently molt extends over a few months. In July and
August, most of the walruses studied in the Bering and Chukchi seas had
hairs that could easily be pulled out, while some animals had already
grown strong new hairs by early July (Nikulin, 1941). In the Kara Sea
in July to October, hairs of “various stiffness” were observed (Chapskii,
1936).

Enemies, diseases, parasites, mortality, and competitors. The enemies
of walruses are killer whales at sea and polar bears on ice (rarely also in
coastal rookeries). The killer whale more often attacks the young, smaller
animals but can be dangerous even to a large animal. In 1936, in the Gulf
of Anadyr, whales attacked a herd of walruses under observation. Some
15 killer whales encircled a group of 60-70 walruses. Two whales broke
into the center of the herd, split it into groups of 10-12 animals, and
totally destroyed one such group. Meanwhile, the rest of the walruses in
the compact group rapidly swam to the coast. The stomach of a killer
whale caught on August 11, 1936 in the Gulf of Anadyr revealed the
skin, blubber, and pieces of flesh of a walrus (Zenkovich, 1938). Only a
large male can keep the polar bear at bay; however, this predator is not
a significant threat to walrus herds.

The only species of lice known among the ectoparasites of walruses,

47 Antarctophthirius Boh., is found in large numbers in the whiskers, in the
skin folds on the sides, in the hind flippers, around the anal opening, etc.
Up to 10 lice per cm? of skin were counted here and there (Ass, 1934).

Eleven species of endoparasites are known. The trematode
Ohdneriella rossica Skrjabin, parasitizing the hepatic ducts of the liver,
has been reported only in the walrus. Orthosplanchus fraterculus Odhner,
a parasite of the gall bladder, has been found in the walrus, the bearded
seal, and the sea otter. The cestode Diphyllobothrium cordatum Leuckart,
which inhabits the small intestine of the walrus, has also been found in
the bearded, common, and Greenland seals; Diphyllobothrium latum has
been reported in many species of pinnipeds and whales, in land carni-
vores, and in man. Diphyllobothrium romeri Zschokke has been found in
the small intestine of only the walrus. The nematode Anisakis (Anisakis)
rosmari Baylis is found only in the walrus stomach. Contracaecum oscu-
latum osculatum Mosgovoy and Ryjikov localizes in the stomach and
small intestine and is found in many species of pinnipeds of the North- —
ern and Southern hemispheres. Terranova (Terranova) decipiens Krabbe,
parasitizing the stomach and intestine, has a wide range of hosts among
pinnipeds and cetaceans. Three species of acanthocephalans parasitize
the intestine. Corynosoma strumosum Rudolphi has been found in seven
species of pinnipeds and two species of cetaceans; Corynosoma semerme

48

53

Forssell in three species of seals, in guinea pigs, and in many birds. The
third species is Corynosoma valdum van Cleave.

Ten of the 11 known species of helminths of walruses have been
reported in the Atlantic walrus, but only five in the Pacific walrus, per-
haps due to the thoroughness with which helminths were studied in the
former (Margolis, 1954; Delyamure, 1955).

The diseases and natural mortality of walruses have not been studied.
It has been suggested that mortality may reach 18-20% of the annual
population increment (Chapskii, 1936), which is rather doubtful. Prob-
ably, the mortality of pups during the first two years of life is extremely
insignificant because of the two-year lactation cycle and the absence of
a large number of enemies.

Adult mortality is mainly the result of hunting by man. Mortality,
mainly of females and pups, occurs in coastal rookeries due to suffocation
by much larger animals. On St. Lawrence Island, in 1936, a large herd
of walruses was chased to the coast by killer whales and more than 20
of them were crushed as a consequence. In July, 1949, 54 dead walruses
(mostly females) were found on Punuk Island 5 km from St. Lawrence
Island (Schiller, 1954).

In November, 1951, many walrus carcasses were found on the coast
of St. Lawrence Island. All of them were greatly decomposed, their heads
severed, and the limbs hanging from the skin. More than 50 carcasses lay
on the beach and many more were afloat in the sea. Based on age and
sex composition (as far as could be determined from such remnants),
a whole herd had perished. The investigator felt that the animals had
been killed by a massive wave (possibly caused by an underwater explo-
sion) close to the Siberian coast, where they sank, then floated after
decomposition, and were brought by winds and currents to the coast
of St. Lawrence Island. The deaths were evidently not due to infection,
as the Eskimos fed this meat to their sledge dogs and used it as bait
for foxes (Schiller, 1954). The cause of mortality of this herd remains
unestablished.

The hearded seal, which also feeds on benthic invertebrates,
competes with the walrus for food to some extent, as mollusks and
crustaceans are of great importance as food for this seal (Pikharev, 1941).
There are no animals which compete with the walrus in habitat selection
on the coast or on ice.

Population dynamics. The walrus population throughout its range
has greatly decreased, mainly as a result of hunting. At present, in the
Soviet Union, the USA, Canada, and Norway, hunting has been banned.
The inhabitants of the Chukchi Peninsula and Alaska are permitted to
hunt walruses, however, to meet their personal requirements.

49

54

Information is not available on the population dynamics of wal-
ruses of the Atlantic Ocean and Laptev Sea. The population dynam-
ics of the Pacific walrus is as follows: 1850-1860, 200,000; 1860 - 1880,
150,000; 1880 - 1910, 80,000; 1910-1950, 60,000; and 1950 - 1956, 45,000
(Fay, 1957). Some attempts have been made in recent years to deter-
mine the population by different methods: aerial photography, aerial
observations, observations from sealing vessels and in coastal rookeries.
The total walrus population in the waters of the USSR was roughly esti-
mated at 30,000. Considering that approximately 20,000 walruses were
counted in American waters during the summer months, the entire wal-
rus population may be estimated at 50,000 (Zenkovich, 1938; Collins,
1940; Buckley, 1958; Fedoseev, 1962; Gol’tsev, 1968; Krylov, 1968).

Field characteristics. The walrus is a large animal with a huge thickset
body, relatively small head, and broad blunt snout, with two huge tusks
directed downward, which are absent in all other pinnipeds. These tusks
are not visible in one-to-two-month-old pups, or they are so small as to
be indistinguishable. The skin lies in large wrinkles and folds, and the
hairs are sparse and coarse in adults.

Walruses form large herds on ice or beaches, on which they are
densely packed (Fig. 28). The animals swimming in water present a char-
acteristic picture of a round head with long tusks. (V.A.)

Economic Importance

Walrus hunting is not important to the overall economy of the country
but is significant in the economy of some regions. In the Chukchi Penin-
sula, the walrus is one of the main sources of raw material for the local
inhabitants. Products of walrus hunting have served as food for the peo-
ple and sledge dogs, and material for building canoes and even houses
(‘уагапраз”) until quite recently. Shoes, dresses, harnesses for dogs, etc.
are made from this raw material.

Walrus hunting is done by two methods—by boats (now banned)
and on the coast. Sealing at sea was done from specially designed small
wooden schooners. As the schooner approached a breeding site on the
ice, motorized whaleboats were lowered from the sides and hunting
commenced. Occasionally, walruses were shot directly from the schooner.
Walruses were killed using firearms, for which reason crews of the
hunting vessels always included some sharp shooters. The skin with the
blubber was removed from the animals killed on the ice and the animals
then hauled in parts onboard the schooner.

Sometimes walruses swimming in water are killed (this practice has
been .banned recently). In this method, a manual harpoon is hurled,

48

55

Fig. 28. А coastal rookery of walruses in the Gulf of Anadyr (photograph by
A.V. Yablokov).

piercing the body of the priorly injured or killed animal. The harpoon
is connected by a short line to floats (inflated seal skin or various types
of artificial floats) which prevent the carcass from sinking. Hunting in
water has invariably been wasteful because no less than 40% of the killed
animals sink and are lost (Р.С. Nikulin and V.I. Krylov). According to
Canadian investigators, such losses have been no less than one-third of
the kill (Mansfield, 1959).

When hunting from a ship, the hide with a layer of blubber is
prepared and salted in the hold and brought to the processing plant
on the coast. The carcass is sometimes brought to the coast and given
to the locals but quite often simply left behind. Only recently has the
bringing of the carcass to the coast been made compulsory. In all meth-
ods of hunting the tusks are invariably removed and used in the making
of various articles.

Local hunters use rifles in hunting walruses in their breeding sites
on ice floes. The sea is surveyed constantly from high ground and when
walruses drifting on floes are sighted, the hunters enter the sea. Hunting
is done by a team of 7-10 men from motorized whaleboats or indige-
nous canoes (now rare) fitted with outboard motors. The whaleboat
approaches the floes as closely as possible and then the hunters open

56

50

Fig. 29. “Warts” on the skin of an adult male. Chukchi Peninsula (photograph
by A.V. Yablokov).

fire. The body of killed animals is cut into large pieces, which are stacked
in the whaleboat and brought to the coast. Wounded walruses are har-
pooned and then shot.

Hunting in coastal rookeries is of little significance. It is resorted to
only by the locals for whom some areas are reserved. Killing is done using
special pikes mounted on long poles. No sound is permitted. Hunters
follow a strict sequence of killing, conforming to traditional practices
geared to conservation. Only some and not all the walruses are killed. It
is believed that a colony ceases to exist if all the walruses present in it
are killed. Campfires and other types of contamination of the rookeries
are prohibited as such scare the animals. The tradition prohibiting the
killing of all animals is based on the assumption that the same walruses
return to the same region year after year.

The number of coastal rookeries in the Chukchi Peninsula and their
populations have decreased considerably. If at the beginning of the 1930s
there were more than ten permanent rookeries, only two are known at
present. Thus walrus hunting in the coastal rookeries has lost its earlier
importance. At present, walrus hunting is done in summer when herds |

3)//

50 Table 3. Walrus hunting in the Chukchi Peninsula (Krylov, 1968)
Year Total killed — Of which
Used Losses (40%)

1932 5,180 3,750 1,480
1935 9,730 6,950 2,780
1938 11,570 8,264 3,306
1941 5,043 3,602 1,441
1944 4,654 3,324 1,330
1947 4,410 3,150 1,260
1950 5,642 4,030 1,612
1953 3,815 2.725 1,090
1956 8,140 5,814 2,326
1959 4,456 3,183 1273
1961 3,602 2,573 1,029

51 Fig. 30. Skeleton of an eared seal (Steller’s sea lion) (figure by N.N. Kondakov).

migrate on ice floes close to the coasts or in regions of their summer
habitat on ice floes.

50 During this same period, in Alaskan waters an average of 2,200 to
2,600 walruses were killed, of which some 1,300 were used (30-50%
loss). Thus about 10,000 Pacific walruses were killed annually. Attempts

51

52

58

to shoot the much larger animals led to the preponderance of males in
the catch which disturbed the natural sex ratio (Fay, 1957; Scott and
Kenyon et al., 1959; Krylov, 1968).

After the banning of state hunting, the catch of walruses in waters of
the Chukchi Peninsula reduced: an average of 2,000 animals were hunted
here until an annual limit was imposed, initially at 1,000 and later raised
to 1,500.

The slow tempo of reproduction determines the extremely slow
restoration of depleted herds of walruses. In order to conserve
their stocks, measures aimed at conservation of animals and hunting
regulation are necessary. Primarily, restriction on hunting is necessary
for each individual herd, so that the killing will not exceed the annual
increase in population and, consequently, not deplete the stocks.

The prospects of exploiting the Pacific populations may be based on
the following premises: total population 50,000; sexually mature, 70%
of the herd; at a 1:1 sex ratio, mature females form 35% of the total
population or 17,500. About 8.5% of the females do not take part in
reproduction, i.e., only 16,000 females are productive. The pups born
per annum number 5,600. Keeping in view not only the need for popu-
lation conservation, but also the need to reinforce it steadily, the annual
catch in Chukchi and Alaskan waters should not exceed half the annual
population increment or 6% of the population, i.e., about 3,000. Further,
measures should be taken to cut down the irretrievable hunting losses.
(V.A.)

SUPERFAMILY OF EARED SEALS
Superfamily OTARIOIDEA Smirnov, 1908

Family of Eared Seals
Family OTARIIDAE Gill, 1866

Males are stocky and large animals while females are of moderate size
with a lighter build. The head is elongated, narrowed exteriorly, and
proportional to the body size. Small pinnae are present. The neck is
long and movable.

The limbs are very long, the fore flippers usually are not shorter than
the hind ones and terminate in skin-cartilaginous tips; their undersurface
is bare. Claws on the fore flippers are either absent or rudimentary. The
hind flippers are capable of bending forward under the trunk and help
in movement on land. The claws on the three middle digits of the hind
flippers are small but well developed and disposed far from the outer
margin. Claws are absent on the two extreme digits.

52

59

Color varies from black to straw-yellow in the various age and sex
groups. The guard hairs on the neck of adult males are long and form a
fairly perceptible short “mane”. The adipose layer is insignificant.

The testes are in the scrotum.

There are four teats.

The width of the skull above the canines is 1/2 - 1/3 that over the ear
openings. The nasals are separated posteriorly by an acute projection
of the frontals (Fig. 31). The supraorbital processes are well developed.
The bony palate is relatively flat. The structure of the tympanic bul-
lae is complex, flattened, and relatively small. Articular, mastoid, and
paroccipital processes are fused into an extremely massive projection,
jutting far downward and sideways beyond the margin of the tympanic
bulla.

The teeth are well differentiated into incisors, canines, and molars.
The upper and lower canines are roughly identical in size, with the upper
ones normally developed (not transformed into large tusks), and the
molars conical. The distinct milk teeth are well developed and are shed
a few weeks after birth. The dental formula is:

3 1 4 2—1
I 5 С т’ Р д М хе 30 are

The scapula is stunted but broad, the humerus and often the ulna
is shortened, but the elements of the hand are extremely elongated and
form a large flipper.

Sexual dimorphism is very pronounced, while age-related dimor-
phism is insignificant.

The family is characterized by distinct polygamy and large harems are
formed in the summer rookeries. The animals survive mainly on certain

Fig. 31. Nasal portion of skuli. A—Steller’s sea lion, Ewmetopias jubatus;
B—northern fur seal, Callorhinus ursinus (figure by N.N. Kondakov). 1—nasal
bone; 2—premaxillary bone.

54

60

types of fish and cephalopods. Eared seals undertake regular seasonal
migrations.

These seals inhabit warm and temperate waters of the Northern and
Southern hemispheres, entering cold seas only in the summer months.
In the Northern hemisphere they inhabit only the Pacific Ocean and the
surrounding seas; in the Southern hemisphere they live along the Pacific
coast of South America and then along the coasts of Australia and New
Zealand. They are found in the Atlantic Ocean along the coasts of South
Africa and South America and also on many subantarctic islands. They
are not found in the northern half of the Atlantic Ocean (Fig. 32).

The systematics of the family have not been properly worked out.
Eared seals have sometimes been grouped into one family with the wal-
ruses and assigned the rank of a subfamily; at other times the two groups
have been regarded as independent families. The majority of authors
attribute a common origin to them. |

This fact and the similarity of several vital morphological features
justify combining the two families into a single superfamily, Otarioidea
Smirnov, contrasting them jointly with the family of true seals, Phoci-
dae, considered under the superfamily Phocoidea Smirnov. In general,
however, within the order the family is regarded, quite justifiably, as the
least specialized (primitive); some authors have compared their origin
with that of the family of bears (Ursidae) of the order Carnivora.

The family comprises 13 genera, of which seven are extinct and six
extant. Members of four of the extinct genera (Allodesmus, Neotherium,
Desmatophoca, and Pithanotaria) first appeared in the Lower Miocene,
the genus Dusignatus in the Upper Miocene, and the other two (Ponto-
lis and Pliopedia) in the Middle and Upper Pliocene. All the fossils of
Otariidae have been found on the Pacific coast of North America and
for this reason this area may be considered the center of origin of the
family (Simpson, 1945; Scheffer, 1958; Mitchell, 1966).

Among the extant genera, Arctocephalus is known from the Pliocene
of South America and the Pleistocene of New Zealand, i.e., within the
distribution range. Two other extant genera (Zalophus and Eumetopias)
are known from the Pleistocene, also within their contemporary range.

The extant fauna includes 6 genera with 12 species: the genus
of southern sea Поп, Опа, with one species, О. byronia [= О.
flavescence]; the genus of Steller’s sea lion, Eumetopias, with one species,
Е. jubatus; the genus of Californian sea lion, Zalophus, with one-species,
Z. californianus (with three subspecies); the genus of Tasmanian sea
lion, Neophoca, with two species, № cinerea (Australia) and №. hookeri
[= Phocarctos hookeri| (New Zealand); the genus of southern fur
seals, Arctocephalus, with six species: A. pusillus (South Africa), forstert

Fig. 32. Range of the family of eared seals, Otariidae (V.A. Arsen’ev).

62

(New Zealand), doriferus (Australia), gazella (Kerguelen), australis
(South America; includes three subspecies), and philippii (possibly
two subspecies) [most authors recognize three additional species of
Arctocephalus: townsendi (Guadalupe Is.), tropicalis (subantarctic) and
galapagoensis (Galapago Is.); and the genus of northern fur seals,
Callorhinus, with one species, C. ursinus (with three subspecies).

Some authors (Scheffer, 1958; and others) are inclined to divide the
extant family into two subfamilies: Otariinae Boetticher with the gen-
era Otaria, Eumetopias, Zalophus, and Neophoca, and a more specialized
Arctocephalinae Boetticher with the genera Arctocephalus and Callorhi-
nus. The second group includes species having fur with underfur (fur
seals), while those of the first group have no fur.

Of the six contemporary genera, two (Callorhinus and Eumetopias)
inhabit the northern half of the Pacific Ocean, three (Arctocephalus,
Otaria, and Neophoca) the Southern hemisphere, mostly the southern
parts of all the three oceans including subantarctic waters [Arctocephalus
townsendi inhabit islands off Mexico in the North Pacific], while the last
genus, Zalophus, is found in both the Southern and Northern hemispheres.

Two genera are found in the fauna of the USSR: Steller’s sea lion,
Eumetopias Gill, and the northern fur seal, Callorhinus Gray. These ani-
mals live mainly in the coastal belt of the Far Eastern seas and in the
northern part of the Pacific Ocean.

The economic importance of the various genera varies. The genus of
northern fur seals (and, to a much lesser extent, the genus of southern fur
seals) is of immense economic value for its extremely valuable fur. The
rest of the genera of the family are not exploited and have no economic
significance. (V.A.)

Key to Species of Eared Seals (Otartidae) [of the USSR]

Identification Based on External Features

1 (2). Hand of fore flipper, both on dorsal and ventral surfaces, without
hair (totally naked). Narrow tips of lateral (I and V) digits of
hind flippers not broader than the three middle ones (II-IV) and
Slightly shorter. Hair coat on trunk with dense, soft, and silky
underfur, better developed in young animals. Length of whiskers
in adult males 30-38 cm, in females 20-25 cm.

Body length of adult males from tip of nose to tail end in a
straight line (Lcv)!° 135-200 cm, of females 110-150 cm. Color

10 Not along the dorsal curvature

2 (1).

3 (4).

4 (3).

п

2 (1).

63

of adult males dark brown, young males and females silvery-gray,
adult females darker with gray streaks. ......................-

позу POTS! Northern fur seal, Callorhinus ursinus (р. 98)
Hand of fore flippers not entirely naked dorsally, metacarpal
region covered with hair. Narrow tips of extreme (I and V) digits
of hind flippers slightly longer and broader than the middle ones
(П-ГУ). Hair coat on trunk either lacking underfur or latter
scanty.

Body length of adult males more than 240 cm (up to 340). Head
massive, with broad and high snout; forehead of males not steeply
raised. Whiskers thick and long (up to 50 cm long and 2 mm
in diameter). Color of adult males varies from straw-yellow (in
summer) to brown (in winter); females yellowish-brown.........
ОНА Steller’s sea Поп, Eumetopias jubatus (р. 68)
Body length of adult males less than 240 cm, of adult females
less than 185 cm. Head with pointed, longish snout, in males with
steeply raised forehead. Whiskers very thin and short. Color of
trunk dark, cinnamon-brown to sepia. .....................---

инж Californian sea Поп, Zalophus californianus (р. 92)

Identification Based on Skull Features

Distance [diastema] between fourth upper premolar and first
molar (IV and V) teeth behind canine considerable, approximately
twice distance between each adjoining pair of premolars. In males,
gap equals 5-8% of condylobasal length of skull. Supraorbital
process massive, fairly squarish, its upper outer edge bent slightly
acurately outward. Sagittal crest in adults high and long; length of
rostral part of skull measured from anterior end of premaxillae to
anterior margin of orbit longer than distance from anterior margin
of orbit to upper posterior zygomatic process. Condylobasal length
of skull in adult males 360-404 mm, in females 290-327 mm.
AS giver). ао topes Steller’s sea lion, Eumetopias jubatus (p. 68)
Distance between last upper premolar and first molar (ТУ and
V) teeth behind canine not more than between remaining teeth
in row, constituting not more than 4% of condylobasal length
of skull (males). Supraorbital process triangular with tip turned
backward. Arcuate flexure of outer side not seen. Sagittal crest
well developed, undeveloped, or extremely small. Length of ros-
tral part of skull same or greater than the distance from anterior
margin of orbit to apex of upper posterior zygomatic process. Skull
size small.

56

64

3 (4). Length of rostral part of skull measured from anterior end of
premaxillae to anterior margin of orbit more than that from ante-
rior margin of orbit to apex of upper posterior zygomatic process.
Nasal processes of premaxillae at level of anterior part of nasals
not broadened and gradually taper toward upper end. Sagittal crest
in adult males well developed, long and high. Condylobasal length
of skull in males up to 330 mm, usually up to 300 mm, in females
пр 25

о: > Californian sea Поп Zalophus californianus (р. 92)

4 (3). Length of rostral part of skull from anterior end of premaxillae to
anterior margin of orbit equal to distance from anterior margin of
orbit to apex of upper posterior zygomatic process. Nasal process
of premaxillae at level of anterior part of nasals considerably
broadened. Sagittal crest absent or very poorly developed.
Condylobasal length of skull in males 220-260 mm, in females
180- 200, MM. т: лоне Gene heed: pest eee: wae ee

Coe ae ae Northern fur seal, Callorhinus ursinus (p. 98). (V.H.)

Subfamily of Sea Lions
Subfamily OTARIINAE Boetticher, 1934

Genus of Steller’s Sea Lion

Genus Eumetopias Gill, 1866

1866. Eumetopias. Gill. Proc. Essex. Inst. 5, p. 7. Arctocephalus monte-

riensis Gray = Phoca jubata Schreber, 1776. (V.H.)
This is the largest member of the family. Males are massive (body length
over 3.5 m) and females half as large. The pinnae are small. The fore
flippers are covered with fur beyond the metacarpus. The margins of the
flippers are represented by a thick, compact coriaceous edge, which is
totally bare. All digits of the fore flippers are devoid of claws; the tips
of the digits are in the form of round disks (rudiments of claws). On
the hind flippers, the two outer digits are without claws (with thickened
horny disks); there are three middle digits with well-developed claws
(Fig. 33).

The hair coat consists mainly of guard hair; the underfur is either
absent or extremely scanty.

The skull is large, in adult males massive, with large crests; the latter
are absent in females. Length of the skull in males exceeds 350 mm, in
females up to 300 mm. Anterior portion of the skull at the level of the
canines is relatively broad. Suborbital apertures are relatively large. The
bony auditory tympana [tympanic bullae] are small and flattened. The

65

56 Fig. 33. Fore and hind flippers of Steller’s sea Поп (figure Бу М.М. Kondakov).

premaxillae gradually narrow toward the apex. The posterior section of
the palatines is almost straight or semiarcuate. That part of the jaw
bearing molars is relatively massive (Figs. 35, 36, 37).

Transverse notches are present on the masticatory surface of the
medial upper incisors into which the cusps of the lower incisors fit. The
lateral incisors are almost as large as the canines, which are thick and
massive. The premolars and molars have simple roots and cusps.

» iit
“А их | у: К д‘ “it
И hia!

57 Fig. 34. Nostril of Steller’s sea lion, Eumetopias jubatus (figure by N.N. Kondakov).

Si

57

66

The genus Eumetopias occupies within the subfamily of sea lions
(Otariinae) a position at the commencement ofa series of specializations.
This is a genus neighboring Опа (southern sea lion), which commences
the series of eared seals and thus stands at the base of the series of
pinnipeds in general. Being a fully specialized typical form of the order
in all respects, this genus of sea lions still bears on its skull the features
of land carnivores. This is the most primitive of pinnipeds of the USSR
fauna and one of the most primitive in the world fauna.

Steller’s sea lion is distributed in the warm [cold] and temperate
waters of the Northern hemisphere and inhabits the coastal waters of
the Pacific Ocean along the Asian and American continents roughly from
33° to 65° N lat. In the west it is found from the coasts of the Korean
Peninsula in the south to the northern coasts of the Sea of Okhotsk,
eastern coasts of Kamchatka, the Commander Islands, and the Bering
Strait. On the eastern coast of the ocean, these sea lions are found from
Pribilov Islands, St. Matthew, Nunivak, Aleutian, and other islands in
the north to the Californian coasts in the south.

The genus consists of a single species, Steller’s sea lion, Eumetopias
jubatus (Schreber, 1776).

pt 7 №

i

= й ges

>
>

Nie

vad
Zama

Fig. 35. Skull of adult male Steller’s sea lion, Eumetopias jubatus [dorsal view]
(figure by N.N. Kondakov).

67

57 Fig. 36. Skull of adult female Steller’s sea lion, Еитеор!ах jubatus [dorsal view]
(figure by N.N. Kondakov).

57 Fig. 37. Skull of young Steller’s sea lion, Eurnetopias jubatus [dorsal view] (figure
by N.N. Kondakov).

58 The economic importance of sea lions is insignificant. In spite of
their relative abundance, their hunting is not well organized and only
a few are caught in the USSR as well as in other places in the North
Pacific Ocean. (V.A.)

STELLER’S SEA LION
Eumetopias jubatus (Schreber, 1776)

1766. Phoca jubata. Schreber. Die Sagetiere, 3, p. 300, Table 83B. Bering

| Island.

1811. Phoca leonina. Pallas. Zoogr. Rosso-Asiat., 1, p. 104. Non Lin-
naeus, 1758. Japan, Kuril’sk islands, Kamchatka. ;

1828. Otaria stelleri. Lesson. Dict. class. Н.М. 13, р. 420. Bering Island.
(V.H.)

Diagnosis

Only species of the genus.
Description

The males are large, massive, and heavily built; the females are usually
a meter shorter than the males and more slender in appearance. The
snout is broad and blunt with a slightly upturned nose. The whiskers are
long (up to 30 cm in females, and up to 60 cm in males) and thick (up
to 2 mm), on average 69 to 71 [?] cm in length. The neck is long and
movable, thick in males, and relatively thin in females. The thick neck
of the males is the result of skin folds and long guard hairs on the nape,
which perform a protective function during fights with other males.

The hair coat consists mostly of guard hairs. The underfur is sparse,
of poor quality, and almost totally absent inold animals.

The color of the hair coat varies depending on the sex and age of
the animal and the season. Newborn pups possess a soft pelage. The
upper part of the body is dark brown or sandy, gradually turning brown
toward the sides; the color is a monochromatic dark chestnut-brown on
the belly. There are no color differences between males and females
at this stage. The color turns perceptibly lighter after the first molt.
Immature juveniles are a light brown. Adult females and males are almost
identically colored, the back creamy, the belly dark umber with a creamy
tinge, more intense in males. The nape of males is a dark purple but in
females creamy. The color of the belly is noticeably darker than that of
the back. The winter fur is darker than the summer fur. In winter old
males are predominantly chocolate or brown, almost black on the belly.
The color gradually becomes lighter in winter; in the summer months
(before molt) the upper part of the body is straw-yellow (Nikulin, 1937;
AS. Perlov).

For the skull description, see the description of the genus. The
condylobasal length of the skull in males is 367-404 mm (x = 389.7), in

Plate I.

Steiler’s sea lion, Eumetopias jubatus Schreber. In the center are an adult male and female.

Top—movement of the animals on land and diving into water; bottom—postures of mating

and playful behavior, and a group of these animals on a rock in the sea (Kamchatka, Cape
Shipunsk, 1973) (figures by V.M. Smirin).

59

69

females 308 - 320 mm (x = 314.7); zygomatic width in males 211-261 mm
(х = 239.8), in females 176-185 mm (х = 180.9); maximum width of
the skull in males 198-238 mm (х = 223.7), in females 154-164 mm
(x = 160.2) (Ognev, 1935; Nikulin, 1937; Chapskii, 1963).

The average body length of males is 320-330 cm, of females about
230 cm. Males weigh 700-800 kg (sometimes over 1,000 kg), females up
to 320 kg. The average weight of internal organs (for 12 specimens) is:
heart 3,233 g, lungs 12,439 g, liver 18,829 g, spleen 526 g, stomach 7,687 g,
intestine 21,473 g, kidneys 1,667 g, pancreas 1,420 g, and mesenteries
5,708 g (A.S. Sokolov et al.). (V.A.)

Taxonomy
See under the characteristics of the genus.
Geographic Distribution

The Steller’s sea lion mostly inhabits the coastal belt of the North Pacific
Ocean where its distribution is very extensive. The range of the species
has undergone no significant change.

Geographic Range in the USSR
Stray animals and small groups are seen throughout along the coasts of

the Sea of Japan, eastern coast of Sakhalin, on Shantar and Kuril islands,
northeastern coasts of the Sea of Okhotsk, eastern coast of Kamchatka,

and Koryak Land. Rookeries are formed every year on [ony Island in the

central part of the Sea of Okhotsk, on Ol’sk and Yamsk islands at the
inlet into Shelikhov Gulf, on Kunashir Islands, Iturup, Urup, Simushir,
Russhua, Onekotan, Srednev Hills, and Kamennye Lovushki, and on
some other islands of the Kuril range; in the Bering Sea at some points
on the eastern coast of Kamchatka (on Shipunsk, Kozlov, and Navarin
capes), on the Commander, Karaginsk, and Verkhoturov islands. Some
rookeries function throughout the year, while some are inhabited only
in summer. In the northern part of the Bering Sea animals reach up to
the Gulf of Anadyr and the Bering Strait (Fig. 38).

Steller’s sea lion has been reported from neither the northwestern
part of the Sea of Okhotsk (from Tauisk Bay to Ayan Bay) nor the upper
reaches of Shelikhov Gulf.

Geographic Range outside the USSR (Fig. 39)
This species is found on the western as well as eastern coasts of the

Pacific Ocean, on the coast of North Korea, and in the Pacific waters of
Japan (Hokkaido Island_and the northernmost part of Honshu Island).

70

59 Fig. 38. Range of Steller’s sea lion, Eumetopias jubatus, in the USSR (V.A. Arsen’ev).

On the Japanese islands, within 35 -37° М lat., rookeries of the Steller’s
sea lion were recorded in the past (Ognev, 1935) but no longer exist
today. In the waters of the Korean Peninsula and Japan the Steller’s sea
lion was observed only in the winter months.

In the eastern part of the Pacific Ocean, Steller’s sea lions are dis-
tributed from southern California (roughly 33° N lat.) along the entire
coasts of the USA and Canada up to the Gulf of Alaska and the Aleutian
Islands, and in the eastern part of the Bering Sea from Bristol Bay to
the Bering Strait. Permanent rookeries are found on many islands of
the Aleutian range (Attu, Kiska, Amchitka, etc.), on the islands of the
Gulf of Alaska, on the Pribilov Islands and on nearby Vancouver Island,

=.

ed
} fj
А
к $. - > \\
р а yt
= 4 09

re x
"6 A <P. . $ “(65
- D OS к о
oe os ver 28“. . a А
Ly Е

Ost 091 Ost C7 06 09 oc 0 or

62

WZ

and also near the coasts of California. Seasonal rookeries are known
on the Bering Sea islands (St. Matthew and Nunivak). In many regions
the distribution is seasonal (Scheffer, 1958; King, 1964; Nishiwaki, 1966,
1966a).

The ranges of Steller’s sea lion and the fur seal lie close by; quite
often, these two species form coastal rookeries at one and the same
places. The ranges of Steller’s sea lion and the walrus almost never over-
lap. (V.A.)

Geographic Variation
Not established.
Biology

Population. Only a tentative estimate of the total population of Steller’s
sea lion is possible because the animals present in the coastal rookeries
are accounted for but not those swimming in the water. In the northern
part of the Pacific Ocean over 160 permanent and temporary rookeries
with widely varying populations have been recorded (Fig. 40).

In the USSR waters the maximum number of Steller’s sea lions
occurs on the Kuril Islands, where 29 permanent and four temporary

Fig. 40. Rookey of Steller’s sea lion. Mednyi Island, July, 1969 (photograph by
S.V. Marakov).

63

73

Fig. 41. Male and female Steller’s sea lion. Mednyi Island, July, 1972 (photograph
by S.V. Marakov).

rookeries are known (Urup and Iturup islands, Srednev Hills, etc.)
(AS. Perlov). The total number of Steller’s sea lions on the Kuril Islands
was determined roughly at 20,000 (Klumov, 1957; Belkin, 1966). On Iony
Island in the central part of the Sea of Okhotsk, 3,000- 4,000 Steller’s
sea lions arrive in the summer months. Comparatively large rookeries
with a total population of 4,000-5,000 are known in the northeastern
part of the Sea of Okhotsk (Yamsk, Ol’sk Islands, etc. at the entrance
to Shelikhov Gulf). Small rookeries are known on the coast of northern
Sakhalin, but the population of Steller’s sea lion there hardly exceeds
1,000. Some ten rookeries of different sizes with a total population of
8,000 - 10,000 animals are known on the eastern coast of Kamchatka. Оп
the Commander Islands, Steller’s sea lion is found quite extensively and
its population varies from 4,000-5,000 in summer to 8,000- 10,000 in
winter (Muzhchinkin, 1964; Nesterov, 1964). Thus the total number of
Steller’s sea lions in the coastal rookeries of the USSR is 40,000 - 45,000.

The maximum number of Steller’s sea lions inhabit the Aleutian
Islands. Here 98 rookeries are known on many islands in different parts of
the range, with a total population of 100,000. The population of Steller’s
sea lion on the Pribilov Islands is around 1,500- 1,600. A large rookery

74

63 Fig. 42. Head of a male Steller’s sea lion. Mednyi Island (photograph by
F.G. Chelnokov).

on Morzhov Island in Bristol Bay contained 4,000-5,000 animals. The
total number of Steller’s sea lions on the islands in the Gulf of Alaska
62 is 75,000-76,000, on the coast of British Columbia 11,000 -12,000, and
on the coasts of Oregon and Washington states 1,000-1,500. Finally,
an additional 6,000 live on the coasts of California (Kenyon and Rice,
1961). Thus the population of Steller’s sea lion beyond Soviet waters is
estimated at 198,000 - 200,000, with a total of 240,000 - 250,000.

‚ Habitat. The coastal rookeries of Steller’s sea lion are of
two types—harems in which reproducing females, males, and pups
concentrate, and rookeries of idle animals (bachelors) which do not
take part in reproduction. The rookeries are located mostly on almost
inaccessible, uninhabited islands or rocky capes. Smooth and level rocky
areas are essential for harems and for females to whelp. Large thickets
of sea kale and algae grow near most harems. The bachelors’ colonies
are Situated in less convenient sites or along the fringes of harems, most
often on rock piles, cliffs, and reefs (Fig. 43). There is no level ground
in these rookeries. In many cases, deep waters adjoin these rookeries so
that the animals can dive into the sea directly from the cliffs, which are
sometimes quite high. The areas of rookeries vary widely, from a few

64

64

Fig. 43. Habitat of Steller’s sea lion. Mednyi Island (photograph by S.V. Marakov).

stray rocks to a large rocky area, depending on the population in the
herd, which may range from several tens to thousands.

Food. Fish and cephalopods of no less than 20 species are the two
basic food groups of the adult Steller’s sea lion. Some geographic vari-
ation has been noticed in the food of these animals. In the Califor-
nian waters, their stomachs revealed flounder, halibut, and bass; on
the Oregon coast (USA): bass, goby, and lumpfish; on the coast of
British Columbia: herring, bass, cod, skate, shark, salmon, octopus, squid,
bivalves, and once a crab; in the Gulf of Alaska: lamprey, salmon, smelt,
sand eel, bass, greenling, goby, shrimp, crab, isopods, squid, and octo-
pus; on the Aleutian islands and in the Bering Sea: pollock, capelin,
sand eel, flounder, goby, herring, halibut, greenling, salmon, and mol-
lusks; on the Commander Islands: smooth lumpsucker, cod, greenling,
salmon, flounder, and rarely octopus; and on the Kuril Islands: sand
eel, greenling, bass, pollock, navaga, goby, flounder, salmon, octopus,
squid, and three species of crustaceans (Pike, 1958; Mathisen, Baade
and Lopp, 1962; Fiscus and Baines, 1966). Very often stones, sand,
gravel, and occasionally algae are found in the stomach of Steller’s sea
lions.

The commercial species of fish are not usually the mainstay but, with
the development of the fishing industry in the Bering Sea, Steller’s sea
lions have begun to live successfully on human effort. The animals have `

65

76

developed а conditioned reflex to the working of fishing trawlers. Оп
hearing the noise of a trawl being hauled, the animals in the vicinity
quickly gather around the boat, dive into the trawl and feed on the
herring. Some animals attempting to get at the fish, damage the net from
the outside and spill the catch. The animals have begun to regularly visit
the trawlers and sometimes in a single trawl as many as 15 Steller’s sea
lions have been found (Tikhomirov, 1964).

The feeding grounds of Steller’s sea lions are sometimes 50-70 miles
(80-112 km) from the coast. In these regions the depth is usually 200 m,
but large groups usually feed close to the coasts; only small groups or
even stray animals venture far into the open sea. At places of high fish
concentrations, the sea lions gather in large herds. Where there are no
fish shoals, the sea lions hunt for them alone or in small groups of two
to five animals (Fiscus and Baines, 1966).

Steller’s sea lions very rarely feed on warm-blooded animals. How-
ever, in 1966, close to Yamsk Islands in the Sea of Okhotsk (59° М lat.),
the stomach of a large male contained the remains of a young ringed seal
weighing 6.8 kg. A Steller’s sea lion was caught on an ice floe. In 1959,
the stomach of six of the nine sea lions caught contained the remains of
seals consumed by them, including adult seals (Tikhomirov, 1959, 1964).
It is quite likely that seals are an incidental, rather than a regular food
item for the sea lion.

Steller’s sea lion females suckle their pups until they are a year
old. It was noticed on the Kuril Islands time and again (more than
ten instances) that a female suckled both a newborn and a yearling.
The stomach of year-olds often contained milk (up to a liter) though
sometimes other foods (beaks of squids and even fish) were found along
with milk (Belkin, 1966a). In the Gulf of Alaska, milk was found in the
stomach of five-year-olds (of the 17 examined) (Mathisen, Baade, and
Lopp, 1962). 1!

11 А suckling pup held in captivity did not touch food for the first 13 days and had to be
force-fed. It later began voluntary feeding. It was given daily 250 to 300 g of rice or beans,
300 to 400 g of meat products, 600 to 900 g of fish, and 30 to 50 g of seal blubber. The
food was mixed with boiled water to make up a volume of 5.5 to 7 liters and fed in four
or five portions (of 1.5 liters each). The ratio was simplified later. In addition to resting at
night, the pup slept in the day after feeding, especially in sunny weather. It soon became
quite tame, moved about the camp, entered the tent, and demanded food. It was released
in the sea but soon returned voluntarily to the camp. After a month it was brought to
Petropavlovsk where it silently accompanied its master through the streets (in spite of the
amused crowds and packs of dogs), climbed stairs to a second floor apartment and laid
down at the door. A few days later, an unfortunate mishap killed it (Kuleshov, 1950).

77

Home range. In harems on the beach, a male is surrounded by 5 to
20 females. Each harem has its own distinct area, its size (5 to 20 m? or
more) depending on the strength of the harem, total area of the rookery,
free zones, animal population, strength and aggressiveness of the male
as the head of the harem, and other features. Harems are usually estab-
lished on comparatively level ground, sometimes even at a height of 10
to 15 m above sea level.

The animals not involved in harem activities stay nearby or on iso-
lated rocks or hill spurs, gathering sometimes at a height of several tens
of meters (Fig. 44). From such a height, the huge young males can easily
and freely dive headlong into the sea. With the dispersal of the harem,
the animals no longer live in age-related groups.

Three types of coastal rookeries have been noticed on the Kuril
Islands: harems, herds of maturing young males (bachelors), and juve-
niles (one- or two-year-olds) (Belkin, 1966).

Daily activity and behavior. On Unimak Island (close to the tip of the
Alaskan Peninsula) Steller’s sea lions living outside the harems gather
early in the morning every day in summer into compact groups of a few
hundreds or thousands and set out for the feeding grounds. Here they
break into groups of a few tens each and feed throughout the day. Before
nightfall they reunite into large groups and return to the rookery (Fiscus
and Baines, 1966). The average number of animals seen on the beach in
the morning hours (ten observations) was 45 and in the evening (seven
observations) 70 (Kenyon and Rice, 1961).

In spring the mature large males (bulls) are the first to appear on the
beaches, where they stake out areas for setting up their harems (Fig. 45).
Claims lead to severe fights and competition. Next to arrive are the
mature females, most of whom are gestating. Harem formation com-
mences, again with fights among rivals. Sometimes after entering the
rookery the female undergoes parturition and the mother shields the
pup for a few days. Later the females go out to sea for feeding and
return to the coast from time to time to suckle the pups. The harem
bulls sometimes abandon their harems and go out to sea; their places
are immediately taken by other bulls. Whether the bulls that go out to
sea return to the harem Ваз not been ascertained. The young of both
sexes, not interested in harem activities, make regular visits to the water.
The harem bulls keep the harmless bulls and young males at bay by chal-
lenging them at the first sign of encroachment (Fig. 46). By June end the
harem bulls become more tolerant and pay no attention to other bulls
in the first half of July, when the harem has disbanded.

In the rookery, the animals are restless and groan incessantly; their
groaning can be heard for miles. The adult male groan is a deep-drawn

78

66 Fig. 44. Steller’s sea lion, Eummetopias jubatus оп a cliff (photograph by S.V. Marakov).

bass, somewhat like the distant siren of a ship, while that of a female
is shrill and penetrating, somewhat like the mooing of a cow. The pup’s
yell is quite shrill and booming, like the bleat of a sheep (Nikulin, 1937).
In the rookeries (mainly bachelor zones) traffic from the coast to the sea
and back is quite regular. This movement intensifies correlated with the
disbanding of the harems. In stormy weather bulls outside the harems
prefer to be at sea, while those on the beach try to gather on cliffs far
away from water; in a prolonged storm, having spent a few hours on the
beach, they, too, take to the sea but remain close to the coast.

Sometimes, Steller’s sea lions are noticed on ice floes where they
remain quite at peace. Animals swimming in the sea are mostly engaged
in seeking food.

Seasonal migrations and transgressions. Steller’s sea lions obviously
do not undertake distant migrations, as they are relatively well-settled

67

79

Fig. 45. Large male Steller’s sea lion (photograph by S.V. Marakov).

animals. In many regions their coastal rookeries function throughout
the year (South Kuril, Commander, and Aleutian islands); however, the
animal population in these rookeries fluctuates seasonally.

Nevertheless, seasonal rookeries are established in the northern
parts of the range in summer, but are wholly abandoned in winter. In
the Sea of Okhotsk (lony and Yamsk islands) seasonal rookeries are
occupied by animals of various ages and sex groups, harems are formed,
and pups born. On the Bering Sea islands (Nunivak, St. Lawrence, and
St. Matthew) bachelors form exclusive rookeries; no females have ever
been sighted in them. In these cases the Steller’s sea lions perform regular
seasonal migrations, evidently for relatively short durations, although
their wintering sites have not been confirmed. The main reason the
animals abandon these islands is the appearance of dense floating ice
floes.

68

67

80

Fig. 46. Rookery of bachelor Steller’s sea lion, Eummetopias jubatus; at the center
is a bull northern fur seal, Callorhinus ursinus. Mednyi Island, June, 1969
(photograph by S.V. Marakov).

On the Commander Islands the sea lion population in winter is con-
siderably higher than in summer. Here mainly young males are seen
although sometimes groups resembling harems occur but without fights
among males as in harems. In 1966 - 1967, a gradual increase in the pop-
ulation of males and adult females was recorded on the Commander
Islands, while in 1968, the gradual appearance of adults in the rookeries
on Mednyi Island was recorded. The formation of genuine harems has
not yet been recorded but observations over decades have confirmed a
few cases of newborn pups. Evidently the change in population of sea
lions on the Commander Islands is associated with their migration from
the rookeries on the eastern coast of Kamchatka (Chugunkov, 1968;
G.F. Chelnokov).

Thus, though Steller’s sea lions do not undertake regular seasonal
migrations, there are regular local wanderings around all the rookeries.

Only a few relatively long transgressions of this sea lion are known.
From time to time, stray animals appear in the northern part of the
Bering Strait on the Diomide Islands (approximately 66° М lat.). The

68

81

Fig. 47. Steller’s sea lions in water. Mednyi Island (photograph by Е.С. Chelnokov).

reported case of a sea lion on Herschell Island in the Chukchi Sea
(69° N lat.) has not been properly verified (Kenyon and Rice, 1961).
Reproduction. Females begin to arrive at the rookeries from May end
to early June, roughly two to three weeks after the males have arrived.
Sometimes, immediately upon arrival, or after a few days, the females
undergo parturition, each giving birth to a single pup. After a few days
the female is mated by a bull. In between birth and mating the bull does

not permit the female to leave his harem and only.after fertilization does

she begin going out to sea, returning periodically to suckle her pup.
The arrival of females at the rookey is not simultaneous and hence
the periods of birth and mating are somewhat protracted: roughly from
the end of May to early June to the end of June. On Chernabur Island
in the Gulf of Alaska, the first births were recorded on May 24 and
the last on June 27 (Mathisen, Baade, and Lopp, 1959*). Since mating
occurs a few days after parturition, gestation obviously extends for about
a year and parturition is perhaps an annual affair. In spite of the harems
being disposed extremely far away from each other along the latitude and
longitude, the periods of reproduction at various places are quite close.
Whelping on the Kamchatka coast occurs in June (Sleptsov, 1950), over
much of June on the Kuril Islands (Belkin, 1966), from May 23 through
June 27 on the Aleutian Islands and in the Gulf of Alaska (Scheffer, 1945;

69

70

82

Mathisen, 1959), and from May end through June end on the coast of
British Columbia (Pike and Maxwell, 1958).

Growth, development, and molt. In Steller’s sea lion the period of
latent development of embryos extends 2.5-3 months and fetal growth
takes place for 9-10 months. Before birth the embryos grow very large.
A female 246 cm in length contained an embryo 94 cm long weighing
12 kg on May 14; an embryo 105 cm long weighing 15 kg was found on
May 19 in a female 234 cm in length, another female, 230 cm in length,
contained an embryo 100 cm long weighing 16 kg on June 2 (Sleptsov,
1950).

The length of newborns at 100-120 cm is roughly half the body
length of the mother and they weigh 17-20 kg. For the first few hours .
the pup is altogether helpless but soon begins to move independently
all over the rookery. In the first 7-10 days these movements are highly
restricted but 25-30 days after birth (Fig. 50) pups move about freely
in water and, by early August, easily cross the channels between cliffs
(Belkin, 1966).

Pups grow very fast and size differences between the sexes are dis-
cernible almost from birth. On the eastern coast of Kamchatka, on June
4 and 6, measurements of 73 pups 10- 15 days old were taken. Males vari-
ed in length from 104-121 cm and females 101-116 cm. A male 107 cm
long weighed 24.8 kg, while a female 104 cm long weighed 27.6 kg [sic]
(Sleptsov, 1950). On Iony Island, on July 23-25, 1933, the body length
of 1-1.5-month-old males varied from 125-138 cm and their weight
from 27-44 kg; the corresponding values for females were 112-132 cm
and 24-36 kg (Nikulin, 1937). The subsequent growth of pups was not
studied.

The average length of yearlings from Alaska was 175 cm (Mathisen,
1959) and their further growth is given in Tables 4 and 5.

These sea lions cease to grow perhaps at the age of 15-20 years. Of
the seven sexually mature males caught on Iony Island, the smallest was
228 cm long and the largest 353 cm, while the corresponding values for
adult females were 170 and 263 cm. A male 345 cm long weighed 768 kg
and a female 263 cm long weighed.208 kg (Nikulin, 1937).

The earlier established period of the onset of maturity for females of
the Sea of Okhotsk at two to three years (Nikulin, 1937; Sleptsov, 1950) is
rather doubtful. The youngest reproducing female (on Chernabur Island),
with a length of 245 cm, was nine years old; the 265 cm long oldest female
was 22 years of age. The youngest mating male was five years old and
the oldest (320 cm) was 19 years old (Mathisen et al., 1959*). The life
span of the Steller’s sea lion has not been established.

83

69 Table 4. Change in body length (cm) with age in male Steller’s sea lions on Vancouver
Island (Fiscus, 1961)

Age No. of animals Average Age No. of animals Average

studied length studied length
1 year 1 178 11 years 3 295
2 years = = 12 years 4 299
3 years 2 208 13 years — —
4 years — ae 14 years — =
5 years 1 272 15 years 2) 317

7 years 1 262 16 years 2 325.
8 years 2 310 17 years — —
9 years 4 288 18 years — =
10 years 3 301 19 years 1 317

70 Table 5. Change in body length (cm) with age in female Steller’s sea lions in British
Columbia (Spalding, 1964)

Age No. of animals Average Range of
studied length length
Newborn 9 100 90-105
3 months 5 120 110-130
9 months 1 160
1 year 23 164 130-202
2 years 3 180 163-189
3 years 3 207 190-221
4 уеагз 5) 213 198-233
5 уеагз 5 223 196-244
6 years 6 227 220 - 232
7 years 7 230 210-242
8 уеагз 5 235 228-245
9 years 7 232 206 - 254
10 years И) 235 221-246
11 years 8 232 216-254
12 years 6 240 226 - 254
13 years 1 230 216-244
14 years 5 236 204-249
15 years 6 244 234-253
16 years 6 243 236 - 246
17 years 2 243 236 - 249
18 years 4 239 218-257
19 years 3 248 245 -251
20 years 1 239
20+ years 4

241 218-254

a

—

84

The newborns are covered with dark brown soft hair. After molting
of the juvenile hair, the fur becomes somewhat lighter; however, the
light brown color persists in the juveniles. Adults sport a rust-colored
coat while the very old animals have straw-yellow fur.

Periods of annual molt have not been established but are believed
to extend from August through November (Nikulin, 1937).

Enemies, diseases, parasites, mortality, and competitors. At sea, killer
whales are the main enemies of sea lions, mostly attacking the young.
Some rookeries are threatened by the brown bear, which is dangerous
only for the young. Diseases and natural mortality have not been studied.
Many pups perish during storms, when waves lash the beach and carry
away the newborns incapable of swimming. During fights among harem
bulls the tiny pups get crushed and die.

The ecto- and endoparasites of the sea lion are diverse. Two species
of mites have been found in the nasopharynx: Orthohalarachne attenuata
Newell, 1947 and Orthohalarachne fluctus Newell. Lice, Proechinophthirus
fluctus (Ferris) and Antarctophthirus microchir (Trouessart and Newman),
parasitize the skin.

Of the 13 species of helminths found in the Steller’s sea lion, two
were cestodes, seven nematodes, and four acanthocephalans. Trematodes
were not detected. The cestodes, Anophryocephalus ochotensis Delamure
and Krotov (known only in Steller’s sea lion) and Diplogonoporus fas-
ciatus (Krabbe), parasitize the intestine. The nematodes, Anisakis triden-
tata Kreiss (not known in other pinnipeds but found in many species of
whales) and Anisakis similis (Baird), localize in the stomach. The stom-
ach and small intestine of this sea lion are parasitized by Contracaecum
osculatum osculatum (Rudolphi) (known from many other species of
pinnipeds in the Northern and Southern hemispheres). Terranova decip-
iens (Krabbe) is widely prevalent in pinnipeds and whales. Parafilaroides
nanus Dougherty and Herman and Parafilaroides proficus Dougherty and
Herman, parasitizing the lungs, are found only in this sea lion. Uncinaria
lucasi Stiles has been found in the intestines of an underyearling.

Among the acanthocephalans, Bolbosoma bobrovi Krotov and Dela-
mure parasitizes the small intestine of the sea lion and the Kuril fur
seal. Corynosoma villosum van Cleave (widely distributed in marine and
land mammals), Corynosoma strumosum (Rudolphi), and Corynosoma
ventronudum A. Skrjabin have been found in the intestines of the sea
lion.

The mite Orthohalarachne diminuata Newell was detected in the tra-
chea and bronchi of the Steller’s sea lion (Ass, 1934; Margolis, 1954;
Delyamure, 1955; A. Skrjabin, 1948).

85

71

Fig. 48. Young Steller’s sea lion with a fur seal. Mednyi Island (photograph
by S.V. Marakov).

Sea lions compete with fur seals, which are ecologically proximate to
them during residence in the coastal rookeries (Fig. 48). Otherwise, these
two species are quite isolated from each other. In the coastal rookeries,
fur seals and sea lions compete in territorial fights for harems. These are
most distinctly seen in the small rookeries of the Kuril Islands, where
harems of sea lions and fur seals are formed at the same sites. Compe-
tition is seen less elsewhere. The breeding period of the sea lion com-
mences somewhat earlier than that of the fur seal. When the harem
periods of the two species coincide, the sea lions (being larger and
more powerful) drive away the fur seals from the adjoining rookeries
and prevent the formation or spread of fur seal herds. However, the
harem period of sea lions is relatively short and their harems begin to
disband usually by the end of June. At this time, the pups of the sea
lions begin moving freely in water, some adults move to other sites, and
the number of Steller’s sea lions in the rookeries considerably declines.
Among the fur seals, however, harem activity is at its zenith by this
time and they are no longer threatened by the sea lions as in the sec-
ond half of June. Thus they are able to freely enlarge their harems and
supplement the number of fur seals taking part in reproduction. The

73

12

86

sea lions entering the rookeries of the fur seals unceremoniously dis-
turb their harem regime with utter contempt for the aggressive fur seal
bulls. However, the sea lions themselves do not exhibit much aggression
and simply suppress the fur seals by their mere physical bulk (Belkin,
1966a).

A few thousand Steller’s sea lions, with young ones predominant
among them, inhabit the Commander Islands in summer. However, some
old males and adult females are also found there. Sometimes even small
harems are formed. In winter and spring the sea lions occupy fur seal
rookeries but have almost completely vacated them by the time the fur
seals are in heat. The two species coexist quite peacefully on the Com-
mander Islands, with no aggressive competition (Fig. 49). On the Pri-
bilov Islands (Alaska), compared with the millions of seals, the sea lion
population is so small that the former completely dominate them and
thus the latter exert no influence whatsoever on life in fur seal rook-
eries.

Such then are the historic relations between the two species of eared
seals in the areas of their cohabitation. They coexist relatively well during

Fig. 49. Steller’s sea lions in a rookery of fur seals. Mednyi Island (photograph
by S.V. Marakov).

74

87

the most important period of their annual life cycle, i.e., in the period
of reproduction.

Steller’s sea lions and fur seals share many items of food, but their
competition in this field is not at all clear (ее under “Enemies, Diseases,
Parasites, Mortality, and Competitors” in the section “Fur Seals”).

Population dynamics. In some parts of the range, significant changes
in Steller’s sea lion populations have been noticed in recent decades,
essentially as a result of human intervention. Mention should be made
first of one of the largest rookeries on [ony Island in the Sea of Okhotsk,
rookeries on Yamsk Islands (northeastern part of the Sea of Okhotsk
at the entrance to Shelikhov Gulf), and some on the eastern coast of
Kamchatka. At the same time, on the Kuril Islands where sea lions are
not hunted, there has been a steady increase in the number of rook-
eries and population of animals in them. Over the rest of the exten-
sive range, population fluctuations, mainly due to natural factors, are
insignificant.

At present, it is the complex of natural factors that exclusively affects
the population of sea lions in our [Northern] hemisphere and they are
practically unaffected by human intervention (there is almost no hunting
of sea lions).

Field characteristics. Steller’s sea lion, one of the largest members of
the order Pinnipedia, is larger than other species of the family of eared
seals. Its canines are comparatively massive and of the same size in the
upper and lower jaws; there are no tusks as in walruses. The pinnae are
small but distinct (Fig. 50).

In the period of residence on the beaches, when the natural color of
the dry animals can be seen, adults are of different shades of rust; by the
end of summer, the hair coat on the neck, shoulders, and back becomes
straw-yellow, on the flanks light rust, and on the belly rusty. Females are
somewhat darker than males. The bulls groan in a deep bass, reminiscent
of a ship’s siren, while the groan of adult females and young ones sounds
like the mooing of cows, and the voice of an underyearling like the bleat
of a sheep. The noise in the rookery can be heard miles away.

Sea lions mainly rest on individual rocks and cliffs (excluding
harems), gathering quite often along cliff projections at a great height.
The animals dive easily into water from precipitous coasts, from heights
of 10 m or more (Fig. 51).

Economic Importance

There has been no state hunting for Steller’s sea lion for over 20 years.
Local hunters use only some rookeries accessible to them and so some

72

88

Fig. 50. Steller’s sea lion pup. Kuril Islands, July, 1962 (photograph by С.М. Kosygin).

herds have not been exploited at all. In view of this, the economic impor-

_ tance of Steller’s sea lion is negligible.

This sea lion is hunted by different methods, including the use of
rifles. Butchering a large sea lion shot among huge boulders is a difficult
task, requiring many hours of hard work.

The raw material obtained from Steller’s sea lion, like that from
other marine animals, can be used quite completely. The skin of the
sea lion serves as a good raw material for making leather goods. Local
hunters greatly value articles made from the skin of the sea lion. The
meat of a young sea lion has an excellent flavor while the meat of other
age groups is satisfactory as food for fur-bearing animals. Locals use it
as food for their sledge dogs. The oil melted from the blubber is useful
in tanneries and a few other industries. The endocrine glands serve as
excellent raw material for making hormonal endocrinal preparations, and
vitamin A is produced from the liver fat.

Steller’s sea lions are few in number and their reproduction tempo is
slow; hence the utilization of their stocks should be planned rationally.
The formation of compact coastal rookeries facilitates rational exploita-
tion of the sea lion, primarily by separating the animals into age and sex
groups. The young males (bachelors) which are in excess for a polyga-
mous mode of life, should be the main target followed, perhaps, by some
adult males. (V.A.)

73

75

89

Fig. 51. Steller’s sea lion diving from a cliff. Mednyi Island (photograph by
F.G. Chelnokov).

Genus of Northern [or California] Sea Lions!2
Genus Zalophus Gill, 1866

1866. Zalophus. Gill. Proc. Essex. Inst., Salem, Communications, 5, p. 6.
Otaria gilliespiit MacBain = Опа californiana Lesson.

The body size is moderate, smaller than Steller’s sea lion (Eume-
topias) but larger than the fur seal (Callorhinus).

The hind flippers are shorter than the fore flippers, their length less
than a quarter of the body length; the outer digits (I and У) on the hind
flippers are somewhat longer and broader than the middle ones (II-IV).
The upper surface of the hand is naked only terminally; the basal half of

12 Tn view of the unusual presence of these sea lions among the Soviet fauna (see below), a
brief description cf the genus as well as the species is given, which suffices for identification.

The morphology is mainly taken from V.B. Scheffer (1958), M. Nishiwaki and F. Nagasaki

(1960), К.К. Chapskii (1963), J. King (1964), etc. (УН.)

90

it is covered with hairs. The undersurface of the hand is wholly naked.
Dense silky underfur is either altogether absent or very sparse. The color
is dark, monochromatic.

The skull is relatively narrow and elongated. Its rostral part, mea-
sured from the anterior end of the premaxillae to the anterior margin
of the orbits, is more than the distance from the anterior margin of the
orbits to the upper posterior process of the zygomatic bone. The profile
of the nostrils resembles a hollow inclined line (not forming a sharp pro-
jection). The supraorbital processes are triangular with a pointed apex
turned backward. The nasal processes of the premaxillae are not enlarged
in the region of contact with the‘anterior parts of the nasal bones, and
narrow uniformly toward the upper end. The sagittal crest in males is
very high and long, commencing in the interorbital region.

The dental formula is:

3 1 4 2—1

I 5? С т Р a? M ae 34.
The wide gap [diastema] between the last upper premolar and the first
molar (teeth IV and V behind the canine) is lacking, the distance between
them being the same as between the premolars, which is not more than
4% of the condylobasal length of the skull (males). The lateral incisors
are massive, tusklike, and the crowns of the cheek teeth are conical with
undeveloped or faint lateral cusps. The first true upper molar (M’) has
one root.

_ These sea lions represent a form which is relatively close to Steller’s
sea lion (Eumetopias). Further relations bring it close to the Australian
sea lion (Neophoca) and the southern sea lion (Otaria). They are usually
grouped together in the subfamily Otariinae, rather than in the group of
fur seals comprising southern (Arctocephalus) and northern (Callorhinus)
fur seals forming the subfamily Arctocephalinae. Thus this sea lion has
very little in common with our fur seal.

The California sea lion genus is ancient; its phyletic links can be
traced in the Miocene.

Its range extends into the North Pacific Ocean between the equa-
tor and 49° М lat., with three isolated populations (Fig. 52): 1) along
the American coast (California to British Columbia); 2) the Galapagos
Islands; and 3) the Sea of Japan and the Pacific coast of the Japariese
islands.

The genus consists of only one species, the California sea lion, Zalo-
phus californianus Lesson, 1828.

This species appears (appeared ?) incidentally in the USSR waters
in the southern part of the Far East. (V.H.)

"(1эи34эН `9’А) чолэцой{ 1е puly 94) зэеотрш инечуес Jo dry и1э15эм 110$ эЦ1
18 10Q зпиримоД]2 `7 ‘цоц Bas JO зэюэ4$ цеплодтео 24) рие 571/40107 uo] eas еплоуе) эц JO зпиэ3 aU} Jo эЗиеч 26 “314 OL

77

78

92

CALIFORNIA SEA LION
Zalophus californianus (Lesson, 1828)

1828. Otaria californiana. Lesson. Diction. class. Hist. Nat., 13, p. 420.
California.

1858. Otaria gilliespii. MacBain. Proc. Edinb. В. Phys. Soc., 1, р. 422.
California.

1866. Otaria japonica. Peters. Monatsschr. K. Preuss. Akad. Wiss.,
Berlin, p. 669. Sea of Japan. (V.H.)

Diagnosis
Monotypical species of the genus.
Description

This sea lion differs from fur seals and especially from Steller’s sea lion
in its lighter and more slender general build; thin, elongated, and flexi-
ble body; and very long and movable neck. The anterior part of its body
is not as massive as in the other aforementioned species. The head of
the males has a sharp snout and a sharply raised forehead, as a result
of the development of the sagittal crest (Fig. 53). Females have a gen-
tler upper line of profile, and the head, on the whole, appears slender
and elongated. The whiskers on the upper lip are long and directed
downward.

The appearance of the female has much in common with the female
fur seal but apart from some large overall dimensions, differs in the
shortened and more hairy hind flippers, and elongated head with a
broader snout.

The color of the California sea lion is very dark, varying from dark
brown of different shades to sepia. An animal with wet fur appears almost
black. With age, the head becomes somewhat lighter in color.

The skull (see characteristics of the genus) is elongated, with a
long rostrum, elongated nasals, and relatively close-set zygomatic arches
(Fig. 54). The interorbital constriction is elongated; the cranium is rela-
tively small and slightly bulged.

The body length of adult males (in a straight line from tip of nose
to end of tail, Lcv) is 215-230 cm, of females 160-185 cm; weight of
males 230-315 kg, of females up to 100 kg. The condylobasal length
of the skull of adult males measures up to 330 mm, of females up to
251 mm. (V.H.)

77

Fig. 53. California sea lion, Zalophus californianus (figure by М.М: Kondakov,
after the American form of the species).

Geographic Distribution and Geographic Variation

The distribution of the California sea lion (see under characteristics of
the genus) is considerable because its range is divided into three sec-
tions located far from each other. This is one of the rarest cases of
an interrupted range of species among mammals. While the distance
between the west American (Californian) and Galapagos ranges is about
2,000 km, that between the Californian and Japanese ranges is sepa-
rated by the whole of the Pacific Ocean (along 40° М lat.), ie., about
8,000 km. Nevertheless, the species identity of the three populations is
indubitable.

The following populations are regarded as special subspecies: 1) Z.
c. californianus (Lesson, 1828)—Pacific coast of North America from the
southern tip of the Californian Peninsula (Las Tres Marias Island, about
21°30’ М lat.) to the southern part of British Columbia at 49° М lat.

77

94

Fig. 54. Skull of the California sea lion, Zalophus californianus (figure by
N.N. Kondakov).

(reconstructed range); 2) Z. c. wollebaecki (Sivertsen, 1953)—Galapagos
Islands; and 3) Z. c. japonicus (Peters, 1866).

The Japanese sea lion is distributed on the eastern coasts of the
middle part of the Korean Peninsula, on the western coasts of Honshu
Island, along its eastern coasts, on the southern coasts of Hokkaido,
and in Sangar Strait (Fig. 55). The range further includes the entire
southern part of the Sea of Japan and that part of the Pacific Ocean
adjoining Honshu, usually not more than a few kilometers from the
coast (Nishiwaki and Nagasaki, 1960). Apparently the range was more
extensive in the past.

The California sea lion is usually not included in our fauna (Smirnov,
1908; Ognev, 1935; Bobrinskii, Kuznetsov, and Kuzyakin, 1965); how-
ever, there is one reference to its possible appearance in our waters
(Chapskii, 1963). Reports of its presence in our waters should be con-
sidered reliable. The reference is not to permanent presence or regular
sightings, but to irregular transgression of individuals. Such is the ref-
erence (Kuroda, 1938, under the name Eumetopias gillespii, cited from
Ellerman and Morrison-Scott, 1966*) to the sighting of this species on

78

79

95

Fig. 55. Range of the Japanese sea lion, Zalophus californianus japonicus (from
Nishiwaki and Nagasaki, 1960).

the Kuril Islands. One sea lion was killed at the end of March or early
April, 1949, on the rocks in the eastern extremity of the small Moneron
Island in the Sea of Japan, slightly west of the southwestern extremity
of Sakhalin, by the keeper of the lighthouse, D. Barabash. The animal
was alone. Its skin, very badly damaged, was examined in July of that
year by the zoologists of Sakhalin Institute, Academy of Sciences, USSR,
A.I. Gizenko and V.G. Voronov, who immediately recognized it as that
of a sea lion. “The general color of the skin is brown, slightly cinnamonic,
and a dark band 3-5 cm wide runs from the tip of the nose to the sin-
ciput [forehead]. The orbital apertures in the skin are surrounded by
pale bands 3-4 cm wide; there is a flesh-colored spot 3-4 cm in diam-
eter below each of these apertures with a slight singeing. Long hairs,
characteristic of Steller’s sea lion, are lacking on the nape” (from the
diary of А.Т. Gizenko, entry of July 10, 1949; letter dated June 9, 1973;
V.G. Voronov). One live animal, possibly of this species, was sighted in
the northern part of Shiashkotan Island (northern Kurils) in November,
1970, and in 1967, a carcass torn by wolves was found on the shoals in
the northern part of Kambal’ny Bay, west of the southern tip of Kam-
chatka (V.G. Voronov). The last instances pertain to the far northern
latitudes and are highly dubious but the appearance of the Californian
sea lion on Moneron is quite possible. As far as we are aware, there is
no museum material confirming the presence of the Californian sea lion
in Our waters.

80

96

There is good justification to assume the presence of this species in
our waters, as the northern boundary of the zone of its permanent habi-
tation or appearance falls on the latitude of Vladivostok and conditions,
at times, facilitate its northward penetration. Such conditions can arise
during intense warm currents, with which this species is evidently asso-
ciated in the Sea of Japan. Thus instances are known in Vladivostok
of the appearance of sea snakes (Pelamys platurus) and turtles (Der-
mochelys coriacea and Caretta caretta), which normally inhabit far more
southern horizons than sea lions and fish. Drifting carcasses have also
been seen. (V.H.)

Biology

The population of the Japanese sea lion has evidently always been low
but has declined sharply in this century. In the early 1950s, its popu-
lation was estimated to be 200-500 and, in 1958, roughly 200 (Schef-
fer, 1958). According to some recent information (Nishiwaki, 1972*),
they had possibly vanished altogether by the early 1970s. In any case,
“the absence of sightings... in the last 30 years suggests the disappear-
ance of this species, though it is possible that some individuals may
have survived on the eastern coast of the Korean Peninsula” (Nishiwaki,
1974).

Data are not available on the biology of the Japanese form. In the
breeding period the animals are confined to the rocky coasts, mainly of
islands, and form harems. The rest of the time is spent mostly at sea.
They feed on fish and cephalopods. (V.H.)

Subfamily of Fur Seals
Subfamily ARCTOCEPHALINAE Boetticher, 1934

Genus of Northern Fur Seals

Genus Callorhinus Gray, 1859

1859. Callorhinus. Gray. Proc. Zool. Soc. London, p. 359. Phoca ursina
Linnaeus.

1866. Arctocephalus. Gill. Proc. Essex. Inst. 5, p. 11, Nec Geoffray Saint-
Hilaire et Cuvier, 1826 (A. pusilus Schreber, 1776).

1892. Callotaria. Palmer. Proc. Biol. Soc. Washington, 7, p. 126. Substi-
tute for Callorhinus Gray. (V.H.)

The males are large and heavily built, while the females are much
smaller and more elegant. The pinnae are small but longer than in the
much larger Steller’s sea lion, narrow, and somewhat pointed.

97

Claws are absent оп all five digits of the fore flippers and replaced by
small horny disks. The outer digits of the hind flippers on which likewise
no claws are present, are only slightly shorter than the inner ones with
well-developed long sharp claws.

The hair coat consists of guard hair and dense soft underfur, better
developed in the young. The color of the hair coat depends on the age
and sex (see below under “Description’’).

The skull is moderate in size, its length in adult males being less than
300 mm (equal to that of the female Steller’s sea lion). Crests are not
prominent. The anterior part of the skull is of moderate width. The cra-
nial capsule is spacious and relatively high. The bony auditory tympana
[tympanic bullae] are very small and, flattened.

The lateral incisors on the upper jaw are slightly longer than the
medial ones. The canines are relatively large and sharp, and the sizes
of the upper and lower ones are almost identical; the upper canines do
not grow into tusks. The molars and premolars have conical crowns and
simple single roots. The skull of the male is larger, more massive, and
with more prominent crests than that of the female.

These polygamous animals gather in large, dense coastal rookeries
in summer. They undertake seasonal migrations.

The genus Callorhinus is one of the highly specialized genera of the
family and is usually regarded (Scheffer, 1958) as the culminating link in
the chain of genera of eared seals.

These are inhabitants of temperate and cold waters of the North
Pacific Ocean where they are encountered on the western and eastern
coasts. In the western part, they are seen from the Sea of Japan (up
to 36-37° М lat.) and the coastal belt on the eastern coast of Japan
(36 - 38° М lat.) to the central part of the western coast of the Bering Sea
(Olyutorsk Bay, 60° N lat.), inhabiting the waters of the Sea of Japan, the
southeastern coast of Sakhalin, the Kuril Islands, near the eastern coast
of Kamchatka, and the Commander Islands. In the eastern part, they are
seen from the coasts of California (33 -35° М lat.), along the entire coast
of the USA and Canada to Bristol Bay, and the eastern coasts of the
Bering Sea (64-65° N lat.).

Callorhinus is one of the six genera of the family of eared seals. This
genus consists of only one species, the northern fur seal, Callorhinus
ursinus (L.), 1758, which inhabits the waters of the USSR and is also
found outside these areas.

The economic importance of the northern fur seal is quite substan-
tial. These fur seals provide highly valuable, high-quality, and extremely
durable fur, and the cost of these furs is quite high as the number of
animals caught is comparatively small. (V.A.)

81

98

NORTHERN FUR SEAL

Callorhinus ursinus (Linnaeus, 1758)

1758. Phoca ursina. Linnaeus. Syst. Nat., Ed. X, 1, p. 37. Bering Island.

1792. Siren cynocephala. Walbaum. P. Artedi genera piscium ... p. 360.
155° W long. and 53° N lat. to south of Kad’yak Island (after Schef-
fer, 1958. У.Н.).

1811. Phoca nigra. Pallas. Zoogr. rosso-asiat., 1, p. 107. “Dal’nie Kuril’-
skie о-уа” [Remote Kuril Islands] (“Ex ulterioribus insulis curili-
cis”). The description is evidently of a young (black) animal.

1828. Otaria kracheninnikovi. Lesson. Dict. class. H.N., 13, p. 420. Substi-
tute for Ursus marinus G. Steller (1751) = Phoca ursina Linnaeus,
1758. Bering Island.

1835. Phoca mimica. Tilessius. Oken’s Isis, p. 715. Terpeniya Bay,
Sakhalin.

1866. Arctocephalus californianus. Gray. Catal. of Seals and Whales, Brit.
Mus., p. 51. Monterey. California.

1897. Callorhinus alascanus. lordan et Clarck. Fur Seals and Fur-seal
Islands, p. 45. Pribilov Islands.

1898. Callorhinus curilensis. lordan et Clarck. Ibid., р. 45. Tyulenii Island,
Sakhalin. (V.H.)'*

Diagnosis
Only species of the genus.
Description

The head is comparatively small, the snout short, pointed, proportional
to the width, and the nose not upturned.

The fore and hind flippers are very long, the latter measuring 30%
of the body length (Fig. 56).

The guard hairs in the hair coat are longer than the fur hairs. The
underfur is more developed in males up to 3 - 4 years and in females up to
5 -6 years, being sparse in much older animals. Thickening of the skin and
long hairs on the neck of adult males are distinctly visible features from

13 Also known in the Russian language as “Morskoi kot,” “kotik,” or “kot”. Black fur
seals are the newborn ones before the first molt; gray fur seals—2 to 3 months after birth;
bachelors—young males aged 2 to 5 years; idle (maturing) bulls—males aged 6 to 7 years;
bulls—fully mature males older than 7 years; cows—females of all ages.

14 Synonymy based on the assumption of the systematic identity of all “herds” of the fur

seal and data on mixed populations (see “Geographic Variation’). (V.H.)

8

8

1

N

99

=

' on VOPR ING
МА iting

Fig. 56. Fore and hind flippers of the fur seal, Callorhinus ursinus (figure by
N.N. Kondakov).

the age of five years. The whiskers in both sexes are directed downward
and long: in males 30-38 cm, in females 20-25 cm.

Adult males are monochromatic yellowish-brown, gray or brownish-
black, with coloration determined by the color of the guard hair. The
anterior part of the snout (around the lips and nose) is grayish-yellow,
while the flippers are black. The underfur is not dense and is rusty-brown.
The coloration of young males is quite diverse, with a predominance of
yellowish-brown and silvery tones. The lower part of the body is lighter in
color than the upper. Females are cinnamon-brown, rusty-grayish-yellow
around the nose and lips, with dark brown head, nape, back, throat, and
upper part of the breast and a very light belly. The flippers are brownish-
black. The color of young males resembles that of females. The winter
and summer hair coats are identical.

The color of the whiskers varies with age. In males and females up to
three years of age inclusively, the whiskers are dark, almost black, while
in those 4-5 years old, they are dark or light, with some animals sporting
dark- and light-colored whiskers. In five-year-old males yellowish-white
whiskers predominate. In both sexes of fur seals older than five years,
the whiskers are wholly monochromatic, yellowish-white.

The premaxillae are sharply narrowed centrally and highly broadened
preapically. The posterior margin of the palatines usually has an acute
angle projecting forward.The jaw section bearing molars is comparatively
weak (Figs. 57, 58, 59). (For a detailed description of the skull, see above
under description of the genus.)

100

>. 5

wise
Fe,

: .: ee

82

82 — НБ. 58. Skull of adult female fur seal, Callorhinus ursinus (figure by М.М. Kondakov).

Age-related Average Size (cm) of Male
Fur Seals of Tyulen’ Island

Age in Years Size

1 about 95 - 100

2 from 90 to 112
3 from 103 to 123
4 from 119 to 136
5 from 130 to 150
6 (maturing bulls) about 175
Adult bulls about 200

Females 4-5 years old do not exceed 115 - 120 cm, while adults reach
130 cm or slightly more.

The condylobasal length of the skull in males is 222-241 mm & =
232.3), in females 182-200 mm ( = 189); width at the zygoma in males
135 - 142 mm (х = 138.7), in females 111-116 mm (х = 114.3); maximum
width of skull in males 118-131 mm (х = 124.3), in females 94-98 mm
(x = 96); length of upper row of teeth in males 65-73 mm (x = 67.7),
in females 50-60 mm ( = 54).

83

101

Average Weight (kg) of Male
Fur Seals of Tyulen’ Island

Age in Years Weight
15-20
about 20
about 29
35-36
about 55

Ak WN

The weight of a bull ‘can reach 250 kg.

Average Weight (kg) of Female
Fur Seals of Tyulen’ Island

Age in Years Weight

2 25
(one specimen)
25.6
29.1
30.7
35.5
33.0
41.0
31

OONAUN HS W

(one specimen)
Older than 10 years 24-45

The weight of large females can ‘reach 60 kg or more.

The weights (in g) of the internal organs recorded for a nine-year-old
female (body length 126 cm, weight 45 kg) were: heart 260, lungs 570,
liver 1,450, spleen 51, kidneys 220, pancreas 250, and mesenteries 290.
The weights (in g) of the internal organs of an eight-year-old male (body
length 174 cm, weight 122 kg) were: heart 700, lungs 1,410, liver 4,700,
spleen 330, kidneys 528, pancreas 320, and mesenteries 1,540 (Ognev,
1935; Chapskii, 1963; Dorofeev, 1964; A. Sokolov et al., 1969). (V.A.)

Taxonomy
See characteristics of the genus.
Geographic Distribution

This fur seal is predominantly seen in the coastal waters on the western
and eastern coasts of the North Pacific Ocean, including the Sea of Japan,

83

102

Fig. 59. Head of a bull and а female fur seal, Bering Istand, July, 1960 (photo-
graph by S.V. Marakov).

the Bering Sea and part of the Sea of Okhotsk. In view of long seasonal
migrations the range varies in the course of a year. These fur seals spend
the winter months in water in the southern parts of their range and the
summer months in the coastal rookeries in the northern parts. The range
has not changed significantly from the historic past.

Geographic Range in the USSR

In the summer months the main rookeries of the fur seal are located
on Tyulen’ Island in the southeastern extremity of Sakhalin in Terpeniya
Bay and on the Bering and Mednyi Islands of the Commander group.
Small rookeries are seen on two groups of islands (Kamennye Lovushki
and Skali Sredneva) of the Kuril range. Individual or small groups of
fur seals may be encountered at many points on the coast of the Sea of
Japan, on the eastern coast of Sakhalin, on some islands of the Kuril

range (Urup, Itirup, etc.), on the eastern coast of Kamchatka right up

to Olyutorsk Gulf (60° N lat.), and sometimes even farther north up to
the Gulf of Anadyr (Fig. 60).

Fur seals winter outside the USSR waters but some stray animals
can be seen close to our coasts. Fur seals wintering in the Sea of Japan

84

84

103

are sometimes seen on Primor’e coasts in the Sea of Japan (at some
stray points) and in the waters of the southern Kuril Islands; a few tens
of fur seals are often seen in winter in Olyutorsk Gulf and clese to the
southeastern extremity of Mednyi Island (Commander Islands).

Range outside the USSR (Fig. 61)

One more (the largest) summer rookey of the fur seal is located on the
Pribilov Islands (St. Paul and St. George) in Bristol Bay (southeastern
part of the Bering Sea). Fur seals move for feeding to the north of
these islands (100 miles or more) in summer when large herds can be

250 500 750 1000 km

Fig. 60. Range of the northern fur seal, Callorhinus ursinus, in the USSR. “+”
indicates the sites of summer coastal rookeries (V.A. Arsen’ev).

104

‘IOWUINS UI [eas Inj
24} JO SaLIayoo!I [e\seoo—Z [eas Inj эЧ1 JO spuly 1иэ2э- т ‘(лэ.иэзгу УД) Snuisin snuTYsO]]DD Чеэ$ IN} ay) JO э8иеы ‘19 ‘84

< 000% 000 000Z 0001 © 0001
(2
ц
РА
р
a Ч
os

$8

86

105

seen there. Some move even farther northward, quite often reaching the
southern part of the Bering Strait. Simultaneously, they are seen along
the entire coast of Canada and that of the USA as far as California.

In the summer of 1965, on the beach of St. Miguel Island in Cali-
fornia, some 10 adult fur seals were sighted. In the next two years, 15 to
20 adults and some pups were seen there, and by 1968 the population
had increased to about 100 (including newborns). The females included
five tagged animals, four from the Pribilov Islands, and one from the
Commander Islands with a 1960 tag. The animals lay close to a rookery
of California sea lions and some elephant seals (Peterson, Boeuf, and
Delong, 1968). This rookery was inhabited in subsequent years also.

Three regions of large winter concentrations of the fur seals are
known: the Sea of Japan, the Pacific waters of Japan, and the coastal
regions of California. In the Sea of Japan, fur seals are found mostly in
the western (Korean Bay) and central parts of the sea in the large Yam-
ato shallow waters (38- 40° М lat. and 133 - 136° Е long.). The southern
boundary of their extent passes along 36-37° М lat., but they are not
found south of it. In the Pacific Ocean wintering herds were seen from
the eastern coast of Hokkaido 36-38° N lat. on the coasts of Honshu

-150 miles or more away from the coast. In the Californian region the

southern boundary of the range passes through 33 -34° N lat. but toward
the north the wintering area reaches 40° N lat. Some groups could be
found in winter over much of the course of the migration route, along
the coasts of Washington and Oregon states, in the waters of British
Columbia, in the Gulf of Alaska, and even close to the Pribilov Islands
in the Bering Sea. (V.A.)

Geographic Variation

The typical variation of fur seals cannot be fully explained by conven-
tional systems. It has long been customary to distinguish three subspecies,
which were generally isolated on the basis of their breeding grounds, win-
tering sites, and migratory routes from summer rookeries to wintering
waters (allopatric). While the differences of form are slight and poorly
defined, there is nevertheless some exchange of individuals between all
the three subspecies, though to an insignificant extent. With increase
in population density (overcrowding), this exchange has also increased
from year to year.

For these reasons, some authors (Scheffer, 1958; and others) refrain
from recognizing distinct morphological geographic variation in this
species. This point of view is perhaps justified; however, the geographic
isolation of the three groups of fur seals is quite real and this compels

87

106

us to recognize three independent populations ог stocks of the species.
A thorough analysis of the prevailing taxonomic relationships between
these populations is therefore necessary.

1. Commander fur seal, С. ursinus ursinus (Linnaeus, 1758) (syn.
kracheninnikovi).

The head and neck are relatively more elongated than in the other
forms. On the fore and hind flippers, 2-3 underdeveloped claws are vis-
ible. The general background color of the young of both sexes is muddy,
while the underfur is a rusty-brown.

Summer rookeries are on the Commander Islands and winter sites
are in the Sea of Japan and on the islands tending to the Japanese waters
of the Pacific Ocean in the south up to 36°N lat., opposite Honshu
(Hondo) Island.

2. Kuril fur seal, C. ursinus niger (Pallas, 1811) (Syn. mimica curilensis).
The head is broader than that of the Commander fur seal. On
the fore flippers 2-3 underdeveloped claws are visible. The underfur
is whitish.
Summer rookeries are on Tyulen’ Island in Terpeniya Bay (Sakhalin)
and winter sites in the Sea of Japan and the Pacific Ocean east of Japan.

3. Pribilov Alaskan fur seal, C. ursinus cynocephalus (Wallbaum, 1792)
(syn. californianus, alascanus).

The head is broadest, compared to the other forms massive, and
the neck is thick. Claws are absent on the fore flippers. The general
background color of the young fur seal is cinnamonic.

Summer rookeries are seen on the Pribilov Islands in Bristol Bay
of the Bering Sea and winter sites in the Pacific waters on the southern
coast of the Aleutian archipelago, along the mainland coast of North
America, and south up to California (San Diego, around 30° N lat.).

Large-scale tagging of pups was undertaken in the last two decades
in all the breeding islands of the fur seal. In some years over 50,000 pups
were tagged and these provided voluminous factual information on dis-
tribution and migration (Fig. 63). The recovered tags showed that the
fur seals of all three populations intermix in the winter areas and also in
the coastal rookeries. The extent of mixing of the different populations
varies. The population of Tyulen’ Island is the most isolated; here the
annual recovery of tags exceeded 1,000. Among the fur seals with tags
caught here, those with Tyulen’ Island tags accounted in some years for
up to 97-99%; those with Commander Island tags 0.6-0.9%; and those
with Pribilov Island tags 0.3- 1.3%. At the same time, in the rookeries

86

87

107

Fig. 62. Nostril area of the fur seal, Callorhinus ursinus (figure by М.М. Kondakov).

on Tyulen’ Island females with Pribilov tags were seen every year. These
gave birth there and suckled the pups. The age of some of them, as
determined from the tag number, exceeded 10 years.

The fur seals of Tyulen’ Island are comparatively isolated even in
the winter areas in the Sea of Japan. In.1963 - 1965, of 49 tagged animals
caught, 42 were of the Tyulen’ Island seals, four of the Commander, and
three of the Pribilov Islands (Arsen’ev, 1964).

Fur seals from Tyulen’ Island mix to a large extent with those of
other populations while wintering in the Pacific waters of Japan. Here,
in 1961-1965, of 566 tagged animals recovered, 376 (66.8%) were of
Tyulen’ origin, 121 (21.6%) of Commander origin, and 66 (11.6%) of
Pribilov origin. Three had tags of the Kuril Islands.

Mixing of the fur seals on the Commander Islands is far greater than
on Tyulen’. Among the tagged fur seals, about 10% had Pribilov tags and
less than 1% Tyulen’ tags. It is interesting to note that on Mednyi Island,

USSR
этинро”

dv: M7914

а AOLONIHSY™

НЕ: =H 1142 =

40N

Fig. 63. Tags for marking the fur seal: Russian tags on top and American tags below.

108

more fur seals with Pribilov tags were recovered than those with tags of
the adjoining Bering Island.

The population of the Pribilov is very poorly supplemented with fur
seals from the western part of the Pacific Ocean. Among 5,000 - 6,000
tagged fur seals, not more than 20-40 were “aliens”. There is no doubt
that the Pribilov population, at least ten times larger than any other
population, is of great importance. =

The recovery of tagged fur seals on the Kuril Islands showed that the
stock here is supplemented equally by all the populations of fur seals.
(V.A. and V.H.)

Biology

Population. The population of fur seals is determined in the summer
rookeries on the beaches, where a good proportion of each population
gathers. The population of the Commander Islands is about 200,000,
Tyulen’ Island 160,000 - 170,000, and the restored population of the Kuril
Islands is roughly 15,000. The largest population, close to 2 million, is
the Alaskan one on the Pribilov Islands. The total population of fur seals
in the northern parts of the Pacific Ocean is thus roughly 2.5 million.

Habitat. In winter and spring, for approximately six months, the fur
seals live scattered in the open sea and form no sizable herds. In summer
they form large coastal herds on the same islands every year.

Rookeries are of different types: in some cases these are purely sandy
or pebbled beaches at the foot of a cliff (Tyulen’ Island and the main
harem оп Mednyi Island); others are rocky platforms with scattered large
boulders (northern rookery on Bering Island); some other rookeries are
wholly covered by massive rocks with no level surface. Fur seals need
a typical protective belt, most often in the form of underwater reefs
or boulders projecting above the water which serve as good breakwa-
ters and protect the rookery from storm waves. In the quiet zone thus
formed pups learn to swim. Sometimes huge shallow waters in front
of the rookery serve as protection. Many rookeries are characterized
by massive thickets of seaweeds (Voloshinov, 1889; Grebnitskii, 1902;
Suvorov, 1912; Tikhenko, 1914).

Food. Newborns grow up on the mother’s milk for the first four
months of life in the coastal rookeries (up to October or November)
(Fig. 68). With the changeover to a marine mode of life, they begin to
feed on the same food as consumed by adults.

The stomachs of fur seals caught in the coastal rookeries were usually
empty. Only some of them (under favorable conditions) revealed food
remnants, generally in the form of small bits of fish or cephalopods. This

109

information is not adequate for characterizing the food of fur seals in
the coastal rookeries (Kenyon, 1956). Therefore, the information on the
food of fur seals in the different parts of the range characterizes their
food when living at sea.

Among the fur seals caught in the sea throughout the range of the
species, many stomachs were empty. In the western part of the Pacific
Ocean the number of empty stomachs varied from 22 to 84% (average
45%) of those examined, and in the eastern part 31 to 63% (average
45%).

In the western part of the Pacific Ocean, the fur seals feed on 21
species of fish and 10 species of cephalopods (in some cases fish or
cephalopods were identified only to the level of genus or family). The
number of species of animals consumed by the fur seals in the eastern
part of the Pacific Ocean was considerably higher. Here 48 species of
fish and six of cephalopods were identified as food of the fur seals.

In the western part of the Pacific Ocean in the region of the Com-
mander Islands, sand eel and greenling serve as the main food, and in
the Sea of Okhotsk (southeastern part), Alaska pollock, greenling, and
some species of squids. In the Pacific waters of Japan most stomachs con-
tained lantern-fish although two species of squids were quite common.
In the western part of the Sea of Japan (Korean Bay), Alaska pollock
is the almost exclusive food (in some, 99% of it) while in the central
part of the sea (Yamato coast) the squid Gonatus magister serves as the
predominant food (Table 6).

In the eastern part of the Pacific Ocean, in the Californian region,
the most important food items are hake followed by anchovy, saury, and
two species of squids; along the Oregon and Washington coasts, hake and
sand eel, and to a small extent herring, capelin, sea perch, and squid; and
on the Alaskan Peninsula sand eel and capelin foilowed by herring and
squid. Prime fishes comprise the main food items only in some regions
and the prevailing view about the damage caused by fur seals to Alaskan
fisheries is clearly an exaggeration. Seasonal changes in food have not
been adequately studied (Taylor et al., 1955; Scheffer, 1950a*; Kenyon,
1956; Wilke and Kenyon, 1957; Arsen’ev and Fedorov, 1964; Pike, 1964,
1966a*) (Table 7).

The average daily food requirements of fur seals in Pacific waters is
1,700 g and in the Sea of Japan 2,500 g, depending on the calorific value
of food (anchovy, mackerel, and squid in the former case and Alaskan
pollock in the latter) (Panina, 1966). The maximum weight of the stom-
ach contents of large males is 10 kg (17 kg in one case), of adult females
2.5 to 3 kg.

89

110

Table 6. Food of the fur seal in the western part of the Pacific Ocean (Arsen’ev and
Fedorov, 1964)
Pacific waters of Sea of Japan Sea of Okhotsk Bering Sea, region
Japan of Commander Islands
1 2 3 4
Fishes Fishes Fishes Fishes
Headlight fish, Pacific salmon, Japanese anchovy, Pacific salmon,
Diaphus sp. Oncorhynchus sp. Engraulis japo- Oncorhynchus sp.
nicus
Gissu, Ptero- Humpback salmon, Pacific salmon, ‚ Humpback salmon,

thrissus gissu

Sardine, Sardinops
melanosticta

Japanese anchovy,
Engraulis
japonicus

Pacific salmon,

Oncorhynchus sp.

Humpback salmon,
Oncorhynchus
gorbuscha

Lantern fishes,
Myctophidae

Lantern fish,
Scopelarchus
linguidens

Lantern fish,
Myctophum сай-
forniense

Anchovy,
Notoscopelus
elongatus

Pacific saury,
Cololabis
5ата

Oncorhynchus
gorbuscha

Alaska pollock,
Theragra chalco-
gramma

Atka mackerel,
Pleurogrammus
monopterygius

Asian greenling,
Pleurogrammus
azonus

Sand fishes, Tricho-
dontidae

Cephalopoda

Squids

Watasenia scintillans

Gonatus fabricit
Gonatus magister

Ommatostrephes sloani-
pacificus

Oncorhynchus sp.

Humpback salmon,
Oncorhynchus
gorbuscha

Pacific herring,
Clupea harengus
pallasi

Lantern fishes,
Myctophidae

Pacific saury,
Cololabis saira

Alaska pollock,
Theragra chalco-
gramma

Greenling, Pleuro-
grammus sp.

Asian greenling,
Pleurogrammus
azonus

Cephalopoda

Squids
Gonatus sp.

Oncorhynchus
gorbuscha

Alaska pollock,
Theragra chalco-
gramma

Char, Salvelinus sp.

Atka mackerel,
Pleurogrammus
monopterygius

Pacific sand lawce,
Ammodytes hexap-
terus

Smooth lumpsucker,
Aptocyclus ventri-
cosus

Greenland halibut,
Reinhardtius hippo-
glossoides

Cephalopoda

Squids
Gonatus fabricii

Gonatus magister

Contd.

111

1 2 3 4
Lantern fish, Gonatus fabricti
Electrona sp. Gonatus magister
Barracuda, Gonatopsis
Sphyraena sp. borealis
Pacific cod, Ommatostrephes
Gadus macroce- sloani-pacificus

phalus
Alaska pollock,
Theragra chalco-
gramma
Pacific mackerel,
Scomber japonicus
Japanese horse
mackerel, Trachurus
japonicus
Prometheus fish,
Promethichys pro-
metheus
Mackerel, Pneumato-
phorus japonicus
Cephalopoda
Octopoda
Polypus vulgaris
Squids
Loligo bleekeri
Onychoteuthis banksii
Moroteuthis bonnbergii
Watasenia scintillans
Gonatus fabricii
Gonatus magister
Chiroteuthis veranyi
Ommatostrephes
sloani-pacificus

The fur seals held in an oceanarium at Enosima (Japan) were fed
mainly squids. The animals were given as much as they could consume,
which averaged 3 to 7 kg. Seals of 2-5 years of age consumed 3 -4 kg and
those of 6-9 years 5 -7 kg. Experiments with two and three feeds showed

91

112

Table 7. Food of the fur seal in the eastern part of the Pacific Ocean (Arsen’ev and

Fedorov, 1964)

California

Oregon, Washington, and

British Columbia

Gulf of Alaska and
Bering Sea

Fishes
Lampetra tridentata
Squalus acanthias
Alosa sapidissima
Clupea harengus pallasi
Engraulis mordax
Oncorhynchus sp.
Hypomesus pretiosus
Thaleichthys pacificus
Tactostoma macropus
Magnisudes baryosoma
Myctophidae
Tarletonbiania crenularis
Cololabis saira
Merlucius productus
Syngnathus californiensis
Trachipterus trachipterus

_Trachurus symmetricus

Brama rayi

Medialuna californiensis
Scomber japonicus
Sebastodes sp.

Sebastodes jordani
Anaplopoma fimbria
Atherinopsis californiensis
Citharidithus sp.
Liopsetta exilis
Polichthyus notatus

Cephalopoda

Tremoctopus sp.
Loligo opalescens
Onychoteuthis sp.
Onychoteuthis banksit
Abraliopsis sp.
Gonatus fabricii
Gonatus magister
Gonatopsis borealis
Dosidicus gigas
Moroteuthis robusta

Fishes
Lampetra tridentata
Hydrolagus collici
Alosa sapidissima
Clupea harengus pallasi
Engraulis mordax
Oncorhynchus sp.
Oncorhynchus gorbuscha
Oncorhynchus kisutch
Oncorhynchus nerka
Oncorhynchus tshawytscha
Salmo gairdneri
Hypomesus pretiosus
Mallotus villosus
Thaleichthys pacificus
Scopelosaurus
Myctophidae
Tarletobiania crenularis
Cololabis saira
Gadus macrocephalus
Merlucius productus
Microgadus proximus
Theragra chalcogramma
Gasterosteus aculeatus
Trachypterus trachypterus
Trachurus symmetricus
Brama rayi
Sebastodes sp.
Sebastodes eutomelas
Anaplopoma fimbria
Ammodytes hexapterus
Tetragonurus cavieri
Atheresthes stomais
Liopsetta exelis

Cephalopoda

Loligo opalescens
Onychoteuthis sp.
Onychoteuthis banksti
Gonatus sp.

Gonatus fabricit
Gonatus magister
Gonatopsis borealis

Fishes
Lampetra tridentata
Clupea harengus pallasi
Oncorhynchus sp.
Oncorhynchus gorbuscha
Oncorhynchus keta
Oncorhynchus kisutch
Oncorhynchus nerka
Oncorhynchus tshawytscha
Mallotus villosus
Thaleichthys pacificus
Lampanyctus nannochir
Anotopterus pharap
Microgadus proximus
Theragra chalcogramma
Gasterosteus aculeatus
Sebastodes sp.
Sebastodes alutus
Anoplopoma fimbria
Pleurogrammus monopterygus
Reinhardtius hippoglossoides
Aptocyclus ventricosus
Trichodon trichodon
Ammodytes hexapterus
Bathymaster signatus
Anarhichas orientalis
Atheresthes stomais
Hippoglossus stenolepis

Cephalopoda

Loligo opalescens
Gonatus sp.
Gonatus fabricii
Gonatus magister
Gonatopsis sp.
Gonatopsis borealis
Other mollusks

91

113:

that the intake was practically identical in both cases. They consumed
more food in summer than in winter. The food intake by the various
age groups varied from 8 to 22% of the body weight, on average 10%
(Nakajima, Sawaura, and Oda, 1963).

Some tens of cases are known of the presence of various species of
marine bird remnants in the stomach of fur seals; these are obviously
cases of incidental intake.

_ Home range. Harems covering 10 т? or more represent definite sec-
tions of habitation on the coasts. The site for a harem is selected and
protected by the bull, who arrives first (see under “Daily Activity and
Behavior”). With the disbanding of the harems, these isolated sections
disappear. The young animals live on the fringes of harems and have no ©
specific section in the rookery.

Daily activity and behavior. Fur seals living on the beaches are more
active in the day and there is continuous movement among the animals.
They are quieter at night, with feeding apparently confined to the early
morning hours.

While at sea, the feeding regime shows certain regional features. In
the Sea of Japan, fur seals begin active feeding in the predawn hours,
which decreases toward midday. Between 1:00 and 4:00 p.m. the seals
rest, mostly on the water surface, and then set out for the second round
of active search; the time of feeding termination has not been established
(Panina, 1964). In the Pacific waters of Japan, in winter-spring, only a
single period of active search has been observed—from predawn to mid-
day. Ву 6:00-8:00 p.m. most of the stomachs are already empty (Panina,
1966a). These differences are the results of the varying accessibility of
food during the day in a given region of habitation.

The behavior of fur seals of different age and sex groups differs at sea
and in the coastal rookeries. In spring, in the latter half of May, bulls are
the first to arrive in any region of the range and occupy a small section
on the vacant beach (Fig. 64). No consideration is apparently given to
the convenience of the location of the future harem when selecting the
site. Possibly, many bulls attempt to occupy those sections in which they
spent the previous year (Kenyon, 1960). The bulls fight savagely with
their competitors to assert their right over territory, inflicting serious
wounds on each other during such battles. Usually the bulls arrange
themselves at a distance of 2-5 m from each other. During this period
there is frequent change of bulls in the sections because the stronger ones
arriving later drive away the weaker occupants. In such cases, however,
the new arrival has invariably to reckon with two or three neighboring
males, so that even a very strong latecomer is not always successful in
entrenching himself in the rookery.

114

92 Fig. 64. Formation of a fur seal rookery: bulls have occupied territories for their
harems. Bering Island, June, 1962 (photograph by S.V. Marakov).

After the bulls, usually from early June, immature males (bachelors)
begin to arrive. The first to appear are the four-to-five-year-olds, followed
by three-year-olds at the end of June, and two-year-olds from mid-July;
year-old males arrive only in August or even in September. A small num-
ber of bachelors of all ages can be seen throughout the summer. These

94 bachelors live along the fringes of the harems, forming a dense inactive
zone; but some bachelors move quite freely in a harem section before
the arrival of a female. At this time, bulls pay them little heed while the
appearance of a new bull invariably invites hostilities. In clear weather
bachelors visit the sea during the day but remain close to the rookery
and return at night for rest. In cloudy and foggy weather they spend most
of the day in the rookery, often playfully imitating the brawls of bulls.

The first of the females arrive in the first ten days of June (Fig. 65)
and their arrival en masse commences in the second half of June or
even in early July. Females in the last stages of gestation arrive singly,
more rarely in small groups. Bulls do not influence the females during
harem selection, leaving the choice entirely to them. Quite often, females
move freely from one group to another and only sometimes does a bull
prevent a female from leaving his harem, even using force if necessary.

92

UNS)

Fig. 65. Formation of the first harem. Tyulen’ Island, June, 1968 (photograph by
V.A. Arsen’ev).

He grasps the skin of the escapes with his teeth and throws her back
into the harem. Upon arrival on the beach the females join more enthu-
siastically the groups already formed rather than single bulls. With the
commencement of the large-scale (Fig. 66) arrival of females, the harems
gradually enlarge and merge, forming collective harems whose bound-
aries are difficult to demarcate. At the peak of harem life the rookery
represents a densely packed mass of females with bulls scattered among
them (Fig. 67). With the arrival of females, the bulls drive the bachelors
away from the harem area.

When moving on land, some fur seals put forward one of the fore
flippers, draw up both the hind flippers simultaneously and then, in the
same sequence, bring forward the other fore flipper and pull up the
two hind flippers. Alternatively, both fore flippers are directed forward
simultaneously and the body weight is shifted onto them before drag-
ging both hind flippers forward with a jerk. Such a mode of locomotion
resembles short hops and the animal moves more rapidly in this manner
(Mordvinov, 1968). Usually, fur seals move very slowly. But it is difficult
for a running man to catch an animal intent on distancing itself (for a
short distance).

Females undergo parturition’ soon after arrival in the rookery. The
time between arrival and parturition varies from 2 to 99 hr (average
22 hr). The female on arrival on the coast, sleeps much of the time
before parturition. The bull protects more attentively a female that has

93

116

Fig. 66. Increasing number of harems. Mednyi Island, June, 1969 (photograph
by S.V. Marakov).

delivered and does not allow her to leave his harem because mating
commences soon after parturition. The interval between parturition and
mating varies from two to seven days (average four days). During this
period the females are confined to the rookery and suckle the pups
(Fig. 68) but go to the sea for feeding soon after fertilization (Bychkov,
1964b). On leaving the rookery for the first time, the female stays at sea
for about a week; on returning, she quite easily finds her own pup among
the thousands present (Fig. 69). The females suckle only their pups and
do not entertain others. The periodicity of subsequent departures to the
sea and return to the beach cannot be tracked in such a massive gathering
of animals.

Marking the Tyulen’ Island bulls with a quick-drying paint showed
that not a single male remains in the rookery for the entire harem period;
rather, there is a continuous exchange of bulls. Of the 54 marked bulls,
57% stayed continuously for 1 to 15 days, 26% for 16 to 30 days, and
17% for over a month. In this period, however, all of them went out to
sea several times, returning to the rookery thereafter. None of the bulls
spent more than a week in the rookery without going out to sea. Most
often, the bull left for a brief period and, having been invigorated at

117

93 Fig. 67. Nearly full rookery. Some bulls still without harems. Mednyi Island, end
of June, 1969 (photograph by S.V. Marakov).

95 Fig. 68. Female fur seal suckling a pup. Mednyi Island (photograph by S.V. Marakov).

118

95

96

Fig. 69. Harem females returning to the beach after being out to sea оп a warm
day. In the background are haremless bulls. Tyulen’ Island, June, 1966 (photo-
graph by V.P. Popov).

sea, returned to his place. The main reasons for going out to sea are a
rise in the ambient temperature or fright, but in some cases sea outings
were prompted for no apparent reason. Along with brief absences, there
were also prolonged departures of marked bulls to the bachelor quarters
where the bulls rested peacefully shoulder to shoulder with the bachelors
and other bulls. Some of them returned to the harem after a few days
and exhibited their prowess. The bull rarely finds his old place since
three or even four bulls press him, compelling him to leave (Fig. 70). It
is for this reason that some marked bulls were noticed at different times
in six or seven different sections of the rookery (Bychkov and Dorofeev,
1962; Dorofeev and Bychkov, 1964). Evidently the bulls arriving first and
occupying a site farthest from the coast can remain the longest in their
territory (average 54.4 days) compared to those who come later and set
up a harem along the coast (average 34.2 days). However, the former can
hold a harem together only for a very short interval (average 19.1 days)
compared to the latter (27.2 days) (Kenyon, 1960).

Immediately after birth, the pups lie alongside their mothers for the
first few days (Fig. 71) and, after her departure, gather in groups forming
a nursery farthest from the sea coast. At this time they sleep soundly for

119

96 Fig. 70. Bulls barring the entry of a competitor into the rookery (in the fore-
ground). Tyulen’ Island, June, 1968 (photograph by V.A. Arsen’ev).

96 Fig. 71. Fur seal pup (“black pup”). Bering Island, July, 1969 (photograph by
S.V. Marakov).

most of the time and do not feed for the six or seven days their mothers
are out to sea. Later, the pups are fed at intervals. In the event of a
mother’s death, her pup inevitably dies of starvation since none of the
thousand or more other mothers in the rookery will take in a stranger.
The growing pups move throughout the rookery beyond their harems,

120

sleep much, and often play with each other for long periods. Later, they
learn to swim (Fig. 72), after which they spend considerable time at sea
in fair weather.

Pups are capable of independent movement on land almost

97 immediately after birth and those that have grown strong move long
distances throughout the rookery, mostly resorting to short hops. Their
speed is such that a man walking rapidly could keep pace.

With the disbanding of the harems, the division of the rookery into
age and sex groups ceases. Bachelors move freely and bulls and females
fill the bachelor quarters, pups are seen everywhere, brawls cease, and
bulls lie quietly beside bachelors and maturing bulls. By October the fur
seals of all the coastal rookeries begin leaving for the sea. The animal
population dwindles and by the end of November the coast is deserted.

Once in water, the animals become agile and quick. In the usual
method of swimming the seals use the fore flippers as the main locomotor
organs and can also use them as rudders or brakes. For a comparatively
small distance the hind flippers can also fulfill the role of a locomotor
organ but usually they serve as rudders and stabilizers. In rectilinear
motion the hind flippers are stretched out along the longitudinal [longer]
axis of the body and set vertically. The fore flippers are first deflected

97 Fig. 72. Pups learning to swim at the coast. Bering Island (photograph by S.V. Marakov).

98

121

forward and raised upward at an angle and then moved backward before
ultimately being pressed to the trunk. In the last stage of this process
the movement of the fore flippers can be very sharp so that it generates
the great thrust necessary for the animal to jump high above the water.
Quite often a herd is seen swimming rapidly by resorting to high long
jumps above the water [“porpoising”] but such maneuvers are used to
cover Only very short distances.

A fur seal, swimming steadily, can cover 5-6 miles an hour. When
resorting to jumps, the speed rises to eight miles or more (Mordvinov,
1968). |

Most fur seals return annually to the same ground where they were
born. The migration of some to other populations hardly disturbs this
pattern. The fur seals return to their own island even when the islands
are situated close to each other, for example: Bering and Mednyi Islands
(the Commanders), and St. Paul and St. George (the Pribilovs) (Nagasaki
and Matsumoto, 1957; Arsen’ev and Fedorov, 1964; Chugunkov, 1966).

Seasonal migrations and transgressions. The distances of seasonal
migrations of the northern fur seal, which reach 1,200-3,500 km, are
the longest among the pinnipeds. Autumn migrations commence at the
end of October to November. The animals set out singly or in small
groups. First to leave are the adult males, followed by the much younger
females and males. Old females and three-to-four-month-old gray pups
stay the longest in the rookery. Each of these three groups follows its
Own migration route, the study of which is largely based on an analysis
of the tags recovered.

The Kuril seals, inhabiting Tyulen’ Island in summer, almost
invariably take one of just two migratory routes. A part of the population
enters the Sea of Japan through La Perouse Strait, where they winter.
Another part enters the Pacific waters through the southern Kuril
Strait and, moving southward, disperses along the eastern coasts of
Hokkaido and Honshu islands. Many seals probably cross Sangarsk Strait
in December-January into the Sea of Japan and join those which have
arrived there from the north.

In winter months the disposition of fur seals in the Sea of Japan is
determined by the availability of food and the hydrologic conditions. The
bulk of them inhabit two regions of the sea: the western part from Peter
the Great Gulf to the Korean Bay and the central part in the region of
Yamato coast (38-40° N lat.). The older bulls apparently do not cross
south of 42° М lat. In the waters between these two regions only stray
fur seals are encountered.

The return journey from the Korean Bay commences in March and
is greatly intensified in April. The fur seals move northeast along the

99

122

west coast and reach La Perouse Bay in May. By this time, they have
already deserted the Korean Bay.

In the central part of the Sea of Japan the fur seals remain through-
out April. Some of them move north, joining the migratory course from
the Korean Bay in the region of Cape Povorotnyi, while others leave
through Sangarsk Strait from where, together with the animals that have
wintered in the Pacific Ocean, they move along the Japanese island coasts
toward Kuril Strait.

In addition to food, another vital factor determining the winter dis-
tribution of fur seals is the surface temperature of the sea. In the Sea
of Japan fur seals are encountered in the temperature range —0.6 to

_+12° but prefer not to go beyond 7 or 8°C. The southern boundary

of the winter habitat of fur seals is the line of convergence of the
surfaces of the much colder Sea of Japan and the very warm Pacific
Ocean water bodies, i.e., the line of the “polar front”. The northward
advancement of this line also compels the seal herds to move northward
(K.I. Panin).

In the Pacific waters of Japan fur seals are seen in December
and, moving along the coasts of Hokkaido and Honshu islands, reach
38° N lat. and, in some years, even 36° N lat. In February and March
the seals are widely distributed between 38° and 40° N lat., away from
the coasts on average up to 145° E long. Some can be encountered far
more eastward, at 160 to 175° E long. (Birman, 1966). The distribution
of animals is not even; they form very dense herds at some places over an
extensive area. However, even at these places the seals live in isolation
or in small herds.

The hydrologic regime of the eastern coast of Japan is determined by
the interaction of two massive currents: the very warm KuroShio and the
much colder OyaShio. The zone of convergence of the currents changes
with the seasons and thus the distribution of seals changes correspond-
ingly; the animals are confined mainly to the zone where the cold and
warm water bodies mix. Food is abundant in this zone. The temperature
of surface waters in the regions of habitat of the fur seal varies from 0
to 15°C but the animal is seen more often at temperatures of 6 to 11°C.

In the fur seal herds in coastal waters, males predominate over
females in all age groups; this predominance is accentuated with age.
In regions far removed from the coasts such a phenomenon is seen only
in groups of yearlings; females predominate in all other age groups.

The northward journey of the fur seals commences in April; by May
they abandon the southern wintering regions and concentrate in June
mainly along the coasts of Hokkaido Island. Later, practically all the
fur seals migrate to the summer rookeries on Tyulen’, the Kuril, and

123

the Commander islands. In 55% cases the fur seals migrate individually,
in 24% in twos, in 9% in threes, and in 4% in fours. In 8% cases,
groups of 5 to 25 seals were seen. In all the age groups up to-five years,
males migrated earlier than females; among males the much older ones
migrated earlier than the younger ones, but this difference in time was
very insignificant (Taylor, Fujinaga, and Wilke, 1955).

Females predominate in the winter herds of fur seals in the Sea
of Japan (since hunting eliminates only males and the females in the
population are usually more numerous). A large number of females are
older than 10 years and females of other age groups are roughly in equal
proportion but in small numbers. Most males are young, 3-6 years of
age. The much older males winter more northward and the one- and
two-year-olds almost never enter the Sea of Japan. In the Pacific waters
where, too, the females are generally more numerous than the males,
the females of all age groups are in equal proportion; unlike in the Sea
of Japan, old females do not predominate here. Among males, young
ones, including yearlings, are more numerous. The fur seals of all three
populations winter in the Pacific waters, though a good part of them are.
the fur seals of Tyulen’ Island.

According to earlier schemes, the fur seals of the Commander Islands
migrate (Suvorov, 1912; Boitsov, 1934; Ognev, 1935; and others) along
the Kuril range to the eastern coast of Japan, where they winter, and
return in spring by the same route to the Commander Islands. An analy-
sis of the large number of tags recovered did not confirm this route since,
in this region, the Commander seals constitute a very insignificant pro-
portion of the wintering herd. The wintering regions and the migration
routes of the Commander seals have not been clearly ascertained to date.

A very small number of seals is present in winter every year on
the southeastern tip of Mednyi Island. In November-December, some
are encountered in Olyutorsk Gulf, even in the region of ice floes
(Chugunkov and Prokhorov, 1966).

The migration of Alaskan seals also begins in October-November,
when they abandon the Pribilov Islands. The autumn migrations occur
rapidly in early December and the first seals are already sighted in
California at this time. The maximum number of animals winter there
in early February. In this period old females predominate; those older
than 10 years occupy the southern part of the winter station (up to
33° М lat.); seven —eight-year-old females remain slightly more north-
ward. The males and young (one- and two-year-old) females are rarely
seen. Immature females arrive later, their maximum number being seen
in the second half of March.

100

124

The bulk of yearlings of both sexes winter in the coastal waters of
British Columbia and Washington state, where they occupy the well-
protected straits and bays. Some seals of 3-4 years of age winter there
but do not venture more southward. Most of the yearlings and some of
the two- and three-year-olds remain in these regions throughout the year
and evidently do not return to the summer rookeries.

Migrations are shortest among bulls, which winter in the Gulf of
Alaska and partly in the southeastern part of the Bering Sea. This age
group is the first to arrive on the Pribilov Islands, from April end
to May.

During March most adult females of the Pribilov population leave
the southern wintering regions. Young females arriving there later
remain longer, usually until the latter half of April. At March end to
April old females are seen in the region of Vancouver Island. They
move gradually northward and are replaced by younger ones, and by
May end to early June almost all the animals found here are under 5
years of age. In May and June congregations of the fur seal are noticed
in the Gulf of Alaska and by mid-June mature females and young males
reach the Pribilov Islands. Young females join them later and the fur
seal population in the Bering Sea reaches its peak in July, August, and
September.

Some fur seals (mainly, perhaps, old males) are found in the Pribilov
Islands even in the winter months. Males with Pribilov tags have been
caught even in Olyutorsk Gulf. A small number of fur seals of both sexes,
but of different ages (mostly young), are seen in winter and summer in
all the areas of their long migratory route (Kenyon and Wilke, 1953;
Taylor et al., 1955; Arsen’ev and Fedorov, 1964; Spalding, 1964a).

Information on the transgressions of fur seals is very scant. At the
end of October, 1947, in Nogaev Bay (northern coast of the Sea of
Okhotsk), a fur seal was killed and some 20 more were sighted there
(Khmelin, 1950). In September, 1958, one was found in the northeast-
em part of the Sea of Okhotsk (Shelikhov Gulf), three strays in the

region of Tauisk Bay, and a group of three close to Okhotsk. The ani-

mals were not large, evidently young, and were spotted 1,000 km away
from the summer rookeries (Tikhomirov, 1964a). In 1959, three solitary
seals were noticed in Gulf of Anadyr (62 - 63° М lat.) and one was killed.
in the southern part of the Bering Strait (64° N lat.). All of them were
large males, perhaps of the Pribilov population (V.A. Arsen’ev).
Reproduction. In the breeding season, harems of varying strengths
are formed in the coastal rookeries. The most favorable ratio is 40-50
females to a bull. The harem population depends mostly on their dispo-
sition. Harems close to the sea are usually more populous. On St. Paul

101

125

Island the average population of 20 harems was 39 females. Instances are
known of a harem comprising as many as 250 females (Boitsov, 1934;
Bartholomew and Hoel, 1953). In such cases there was, perhaps, a fre-
quent change of bulls. As harem life reaches peak activity the harem
boundaries merge and it becomes difficult to count the females under
each bull. While enumerating the number of heads in a herd, the bull-
to-female ratio is taken as 1:40 -50.

The period of parturition and mating extends for about two months.
The first of the newborns are usually found in the rookeries on Tyulen’
and the Commander Islands seldom earlier than June 10. The number of
pups increases quite rapidly but maximum whelping occurs in the second
half of July, after which it begins to taper off. Some births and mating
are observed in the first ten days of August, though exceptions do occur
somewhat later. Because of the impregnation of the female soon after
parturition, gestation is believed to extend to about 360 days. Mating
mostly occurs on land, more rarely in shallow water, and can last from
5 to 40 min.

Young females arrive in the harem toward the end of the harem
period. On Tyulen’ Island four-year-old females (age determined from
recovered tags) begin to be seen in the latter half of July and some of
them are impregnated. Two-year-old females have not been recorded in
the harems before July 25 (Bychkov, 1964a).

Fig. 73. A group of “black” pups. Tyulen’ Island, July, 1968 (photograph by
V.A. Arsen’ev).

101

102

101

126

The male-to-female ratio among newborns is close to 1:1. Of the
4,276 embryos examined during investigations at sea in 1958-1961 in
the eastern part of the Pacific Ocean, 2,135 (49.9%) were males and
2,141 (50.1%) females. In the western part of the Pacific Ocean, of the
2,741 embryos examined, 1,393 (50.8%) were males and 1 348 (49.2%)
females (Arsen’ev and Fedorov, 1964).

The female usually gives birth to a single pup; twins are extremely rare.
In most cases, parturition is annual and sterility comparatively low, but
increases in old females. Under the conditions prohibiting the shooting
of females and normal herd growth, the annual increment of population
was 8- 8.5% (Dorofeev, 1964).

Table 8 gives an idea of the number of gestating females according to
age. Females in the eastern part of the Pacific Ocean begin reproducing
a year later than those in the western part (reason not established).

The period of latent growth of the embryo evidently extends over
two months. The embryo grows quite rapidly, and even at this stage the
growth of males surpasses that of females, as can be seen from Table 9.

Growth, development, and molt. The body length of newborns is
60-70 cm and their average weight is 5-6 kg. In individual cases the
weight of the newborns may exceed 10 kg. Within a few hours of birth,

Table 8. Age-related gestating females (Arsen’ev and Fedorov, 1964)
Gestating among them
Age, years No. of females Number Percent

in sample

Western part of Pacific Ocean

4 847 413 53
5 637 S31 83.
6 423 358 85
7 241 216 90
8 137 125 91
9 94 86 91
10 69 61 88
10+ 279 201 72
Eastern part of Pacific Ocean
4 375 16 4
5 403 180 45
6 445 340 76
a S45 434 80
8 609 519 85
9 555 501 90
10 513 455 89

10+ 2,641 2,171 82

102

103

127

Table 9. Change in length and weight of embryos in Japanese waters (Arsen’ev and
Fedorov, 1964)
Length, cm
Month Males Females
Number in Average Number in Average
sample length sample length
February 5 27.0 12 13.2
March 17 33.6 3 31.2
April 151 43.9 146 42.2
May 125 515 116 50.8
June 18 57.0 24 53.9
Weight, kg
Month Males Females
Number in Average Number in Average
sample weight sample weight
February 1 0.4 6 0.1
March 17 1:3 3 0.8
April 151 2.1 146 1.9
May 125 3.7 116 3.3

June 18 6.2 24 48

pups begin moving quite freely throughout the rookery. They gather
strength quickly and, after a few days of suckling, form separate groups,
mostly away from the sea. The stomach volume of a newborn pup weigh-
ing 4.5 kg was 1.1 liter, and of another 1 liter (fat content of seal milk
about 45%). After 3-4 months, by the end of the suckling period, the
body length reaches 70-80 cm and weight 15-17 kg (Scheffer, 1950).
The pups can withstand well all types of mechanical force and do not
suffer serious damage even when pressed by a bull weighing 200 kg.

Pups are capable of staying in water a few days after birth but avoid
the sea for 3-4 weeks. Only then do they begin to learn to swim in
shallow water in calm weather. In August the pups go out to sea on their
own and swim for long periods within sight of the coast. In September
they are as adept as their parents at sea, at times venturing far out; by
October-November they leave the islands for a pelagic mode of life.

On taking to independent feeding, the growth of young seals some-
what slows down initially but later proceeds steadily for up to 10-12
years. At this age the animals have reached full development and further
growth wholly, or almost wholly ceases. Throughout the growth period
males grow faster than females, as a result of which the adults of the two
sexes differ considerably in size (Tables 9, 10, 11).

A large bull can weigh 285 kg and a large female 63 kg (Dorofeev,
1964).

128

102 Table 10. Change in body length (cm) of the fur seal with age (Arsen’ev and Fedorov,

1964)
Males* Females
Age, years Number in Average Number in Average

sample length sample length

1 28 78.5 14 76.0
2 30 96.5 4 95.3
3 52 107.5 3 100.7
4 12 121.7 36 109.3
5 9 129.1 55 115.5
6 9 142.5 45 121.3
fl 7 148.2 66 121.4
8 5 162.9 105 123.6
9 8 168.7 143 125:3
10 5 175.9 129 124.3
11 137 127.9
12 106 126.6
13 120 126.5
14 108 128.4
15 67 129.9
16 51 130.4
17 46 127.6
18 23 129.1
19 19 130.7
20 6 130.0

* Adult bulls measure up to 200 cm or more.

103 Table 11. Change in weight (kg) of the fur seal with age (Arsen’ev and Fedorov, 1964)

Males Females
Age, years Number in Average Number in Average
sample weight sample weight

1 1 23.0 — =
2 47 24.0 1 25.0
3 404 315 11 25.6
4 190 40.9 40 291
5 21 56.4 26 30.7
6 9 71.0 10 355
7 3 116.0 5 33.0
8 5 162.6 1 41.0
9 = = 1 31.0

10+ == — 8 38.5

The age of the fur seal is determined from the annual rings at the
base of the upper canines (Scheffer, 1950). The number of layers in males
corresponds to the number of years and, in females the first layer from

103

129

the tooth crown zone is ignored when counting the number of rings.
The accuracy of determining the age based on these layers was verified
in tagged seals (Fig. 74).

Some females attain sexual maturity and are impregnated for the
first time in the third year but most mate in the fourth year, while the
vast majority begin reproducing by the fifth year (Arsen’ev and Fedorov,
1964; Craig, 1964). Some whelp only in the 7th, 9th, and even the 12th
year. The period of maximum fertility extends up to the age of 20 years
although by this time there is a relative increase in sterility. It has been
assumed that females retain their ability to reproduce throughout their
life (Boitsov, 1934).

Spermatozoa have been reported sometimes in the testes of three-
year-olds (Grebnitskii, 1902; Boitsov, 1934); usually, however, males
attain sexual maturity at 5-6 years of age and begin to mate even later.
Seven-year-old bulls, because of their physical immaturity, still cannot
occupy a place in the harem rookeries and are chased away by the more
powerful older bulls. Even in the 8th year the bull is not always successful
in forming his own harem (Fig. 75); if successful, he does not remain
for long in the harem rookery. Apparently, the bull attains maximum
strength by the age of 10 years (Fig. 75). The period of cessation of
sexual activity in bulls has not been established. There is no information
on longevity.

At 3-4 years of age the guard hairs on the nape of males begin
to grow and form a “mane” which represents a secondary sex character.
Even at 5 years of age the mane is quite visible and becomes very distinct
in the subsequent year. This feature is fully developed with the onset of
sexual maturity.

Fig. 74. Annual marks (layers) on the canines of fur seals. A—male, B—female
(figure by N.N. Kondakov).

104

104

130

Fig. 75. Large bull fur seal with а harem. Bering Island, July, 1972 (photograph
by S.A. Olekhnovskii).

_ Uterine molting has been noticed in embryos. During this period the
primary pelage is replaced by infantile hair, which is seen in the newborns
(Belkin, 1963). The hair coat of pups consists of soft black guard hair
(“black” pups) and very low, rather poorly visible underfur. The first molt
takes place roughly a month after birth. As molting progresses, the black
guard hairs are shed and brownish underfur grows vigorously; as a result
the pups change from lustrous black to dull brown. Later, the guard
hairs are shed and replaced by new hairs that are gray, changing again
the overall color of the pup. By four months of age, the pup completes
the first molt and acquires a beautiful silver-gray coat. The end of molt
coincides with the pups leaving the island and taking to a pelagic mode
of life.

Molting of young males (bachelors) occurs partially during their res-
idence in coastal rookeries; some even arrive on the island in the initial
stage of molt. The very young males (one or two years old) are the first to
molt, followed by the older ones. Molting peaks in September-October,
however, not all animals molt on the coast and the process is completed
at sea.

Molting of maturing and young bulls on Tyulen’ Island was observed
at the end of September. Three observations have been reported of traces
of nitrolacquer (used for marking in the preceding year) on the hair coat
of old bulls. It is possible that some old bulls do not molt every year. It is

105

131

thought that moderate molting continues until January and February of
the following year (Scheffer, 1962; Scheffer and Jonson, 1963; Bychkov,
1964). ;

Molting of females commences in August after whelping, and coin-
cides with the period of suckling. Evidently, molting is not yet complete
when the females leave the coast by November and is completed at sea.

Molting of the hair coat commences on the head, at the base of the
fore flippers, and at the tail end of the trunk. Molting then gradually
extends to the sides, back, and belly. During molt new hairs fully replace
the old. Even the fur hairs molt. Whiskers do not molt but, with the
onset of sexual maturity, they become gradually depigmented.

Enemies, diseases, parasites, mortality, and competitors. The fur seals
have no enemies in the coastal rookeries. At sea they are sometimes
attacked by the killer whale (Orcinus orca).

Newborn pups are known to suffer from uncinariasis caused by
intestinal parasites, most often causing the death of the pup. Instances
are known of inflammation of the respiratory tract and intestine. Adults
suffer from diseases of the eyes, sometimes resulting in blindness.
Skin diseases resembling scabies have been reported (Boitsov, 1934).
On the Commander Islands, bronchial, gastric, and intestinal catarrh,
infectious paratyphoid, and dermal herpes have been reported. On
Tyulen’ Island, there have been instances of bilateral pyelonephritis,
hemorrhagic meningitis, peritonitis, and endocarditis (Dorofeev, 1964).
In the blood of 18 fur seals (34 examined) on the Commander Islands,
microfilariae of an indeterminate species, were detected (Delyamure et
al., 1961).

Two species of lice parasitize the skin of the fur seal: Antarctophthirus
callorhini (Osborn, 1899, McAtee) and Proechinophthirus fluctus Ferris,
1916, Ewing. Parasitic mites have been detected in the naso
pharynx (Orthohalarachne attenuata Newell) and in the trachea and
bronchi (Orthohalarachne diminuata Newell).

Nineteen species of helminths have been recorded: one species
of trematode and six species each of cestodes, nematodes, and
acanthocephalans. The trematode, Phocitrema fusiforme Goto and Ozaki,
localizes in the intestine (known also in ringed seals and sea otters).
Of the cestodes, Adenocephalus septentrionalis Nybelin parasitizes the
large intestine (not known in other animals) and Clestobothrium
glaciale Cholodkovsky is found in the intestines. Diphyllobothrium krotovi
Delamure has been detected in the small intestine of only fur seals.
Diphyllobothrium (Bothriocephalus) lanceolatum Krabbe, quite a common
parasite of marine mammals, is found in the intestine of fur seals and
Diphyllobothrium tetrapterus Sibold, in addition to the fur seal, is found in

106

132

some other species of pinnipeds. Finally, Diphyllobothrium macrocephalus
Linstow, in addition to fur seals, has been reported only in bearded seals.
The nematode, Contracaecum osculatum osculatum Rudolphi, found in
fur seals, parasitizes the small intestine and stomach of many marine
mammals in both hemispheres; Phocascaris phocae Host, also known
in the Greenland seal, has been detected in the duodenum and in the
small intestine. Terranova decipiens Krabbe parasitizes the fur seal and
many species of pinnipeds of both hemispheres, and is also recorded
from some cetaceans. Terranova azarasi Yamaguti and Arima is known
only in pinnipeds. Anisakis pacificus A. Skrjabin, known also in the
three species of cetaceans, has been found in the intestine. In the small
intestine of fur seals, especially in yearlings, the nematode Uncinaria
lucasi Stiles has been detected. From the acanthocephalans, Corynosoma
strumosum Rudolphi has been reported in almost all the pinnipeds of
the Northern and Southern hemispheres, in two species of cetaceans, in
land mammals (cats and dogs), and in many species of birds. Corynosoma
ventronudum A. Skrjabin has been found in the intestines of only Steller’s
sea lions and the northern fur seals. Of the two species of intestinal
parasites, Corynosoma semerme Forsell is reported from many other
marine animals and birds, while Corynosoma villosum van Cleave only
from fur seals. Bolbosoma bobrovoi Krotov and Delamure, found in the
small intestine of northern fur seals, is also found in Steller’s sea lions
and Bolbosoma nipponicum Yamaguti, parasitizing the small intestine
and cecum, has been found in three species of whales and ringed seals.
The helminths found in various populations of the northern fur seal are
listed in Table 12.

The largest number of species (13) of helminths is known in the
Commander fur seals and only two species are common for all the three
subspecies. In the southern fur seal, seven species of helminths are found,
of which only one (Terranova decipiens) is known only in the northern fur
seal (Margolis, 1954; Delyamure, 1955; Olsen, 1958; A. Skryabin, 1958;
Delyamure and A. Skryabin, 1960).

Mortality is maximum in infancy, mainly in the first one-and-a-half
months after birth. With the disbanding of harems (from early August),
the mortality of pups declines considerably. Mortality varies widely in
different years but usually does not exceed 16-17% of those born. The
main cause of pup mortality on the Pribilov Islands is the intestinal para-
site Uncinaria lucasi, which causes anemia and emaciation. Other causes
are trauma inflicted by adult fur seals or injuries caused by falling from
cliffs where the pups gather quite eagerly. A relatively large number of
pups perish from starvation when they lose their mothers. Instances of
mortality caused by bronchial inflammation of the lungs are also known.

106

133

Table 12. Helminths in the northern fur seal (Delyamure and А. Skryabin, 1960)

Subspecies of fur seals
Helminth species eee eae ee eee ee eee ee
Commander Kuril Alaskan

Phocitrema fusiforme
Adenocephalus septentrionalis
Cestobothrium glaciale
Diphyllobothrium krotovi
Diphyllobothrium lanceolatum
Diphyllobothrium macrocephalus
Diphyllobothrium tetrapterus
Anisakis pacificus

Contracaecum osculatum
Phocascaris phocae

Terranova azarasi

Terranova decipiens

Uncinaria lucasi

Bolbosoma bobrovoi

Bolbosoma nipponicum
Corynosoma semerme +
Corynosoma villosum =
Corynosoma strumosum + +
Corynosoma ventronudum +

+++ ++++++

++ + +

* АЕ the end of the 1960$ 0. lucasi was also recorded from the seals of Bering Island

(V.A).

The extent of mortality increases in years of unfavorable climatic condi-
tions. In 1965, on Tyulen’ Island, a catastrophic mortality of pups resulted
from severe and prolonged storms; waves flooded the harems and washed
away pups which had yet to gather strength. Over 20,000 pups perished
(some 40% of those born that year). However, such cases are extremely
rare. Mortality also increases with increasing population, ie., crowding
in the rookery.

The mortality rate in the rest of the age groups in the coastal rook-
eries is very small, only a few tens [of seals]. Adults die mainly at sea.
Young ones, especially those born in that year and the yearlings inca-
pable of withstanding severe storm conditions, experience difficulties in
getting at food and die of starvation. The mortality of the Pribilov females
was mathematically computed as 18% for eight-year-olds, 20% for nine-
year-olds, 19% for ten-year-olds, and 25% for 12-year-olds and beyond
(average 20%) (Nagasaki, 1961; Chapman, 1961). Considering these val-
ues as high, American scientists use, in different calculations, three mean
mortality coefficients for females of all ages: 5%, 10%, and 15%. This
question has not been finally resolved.

107

134

At different times of the year, different types of competition arise
between the northern fur seals and Steller’s sea lions, these being most:
pronounced in the period of life in the coastal rookeries. These com-
petitions are mostly determined by the numerical ratios of these two
species. A relatively simple form of competition has been observed on
Mednyi Island (Commander Islands) where in summer some 100,000 fur
seals and 4,000-5,000 Steller’s sea lions live side by side (Fig. 76). The
fur seals form dense harems while the sea lions are represented mainly
by young immature males. At the end of the 1960s, the number of old
male sea lions began to increase, mature females appeared, and some-
times even small harems were formed. The interrelations of these two
species of animals vary slightly with the varying herd structure of the
sea lions. The reproducing sea lions (especially the bulls) are the most
aggressive toward the fur seals (S.V. Marakov). The fur seals spend only
the summer on the island while the sea lions live there throughout the
year, their numbers increasing significantly in winter.

Fig. 76. Steller’s sea lion, Eurmetopias jubatus, in a rookery of the northern fur
seal. Mednyi Island (photograph by S.V. Marakov).

108

135

In early spring Steller’s sea lions occupy much of the territory of the
northern fur seals and the fur seal bulls are compelled to put up with
the stronger and more powerful Steller’s sea lions. However, as the fur
seal females start arriving, the sea lions gradually vacate the rookeries
for the bachelor quarters and isolated boulders scattered abundantly on
the island coasts. It cannot be said that the sea lions are forced out
by the growing harems of the fur seals because they invariably leave
the rookery in the wake of the en masse invasion of female fur seals.
When hunting for fur seals commences, the Steller’s sea lions leave even
the bachelor quarters. However, some sea lions do remain and are even
caught along with the fur seals. As the harem activity abates, harems
disband, and hunting for the fur seals ceases, the sea lions again partially
occupy the harem rookeries in which the two species coexist peacefully
(Muzhchinkin, 1964). A similar picture, but involving a smaller number
of Steller’s sea lions, is observed on Bering Island.

Such interrelations are suggestive more of coexistence than of com-
petition, but there is a view that sea lions inflict damage on fur seals and
are, therefore, undesirable inhabitants of the rookeries (Marakov and
Nesterov, 1958).

Sometimes fur seals of unusual color—rust, light cinnamon, etc.,
have been found. In a rookery on the Commander Islands, a bright rusty
female was seen suckling a normally colored pup; at another place a large
well-fed pup of chocolate color was seen. On Tyulen’ Island a bull of
normal size but of yellow color was found in a harem. However, colored
fur seals are usually seen among small-sized animals, females and young
males (V.A. Arsen’ev). According to hunters (S.P. Naumov, 1933), male
sea lions ravish the female fur seals, but the fur of the offspring is poor
and is rejected in trade. There is no precise proof for this statement
and it is probably erroneous as successful crossing of such taxonomically
distant animals is impossible. Without ruling out attempts at crossing
Steller’s sea lion with fur seals, it may be said that fur seals exhibiting
aberrations from typical coloration are considered hybrids.

Another form of relationship has been noticed on some islands of the
Kuril range in which separate harems of both species are formed in the
same rookeries and their offspring born. But such rookeries are small and
the population of animals is much less than on the Commander Islands.
During June such rookeries are occupied by harems of the sea lion,
among which whelping is at its peak. The fur seal bulls arriving in early
June and the females in the latter half of this month, are forced to huddle
together along the fringes or occupy the free sections for raising their
pups. In this period, the large and powerful sea lions totally dominate
the fur seals.

136

108 Fig. 77. Female northern fur seal. Bering Island, November, 1965 (photograph
by S.V. Marakov).

In early July the harem life of Steller’s sea lions becomes calmer,
their numbers on the beach diminish rapidly, and pups begin to enter
the sea freely; most of the sea lions migrate to other sites or even to an
adjoining island. The small number of sea lions remaining in the rookery
no longer disturb the growing harems of the fur seals, whose harem life

109 is now beginning to reach its peak. But even at this time the sea lion
bulls remaining in the rookery move freely throughout, chasing the fur
seal females and bulls without heed for their belligerency. Such instances
are not frequent, however. The differences in the whelping periods of fur
seals and sea lions largely smoothen the competition between them and
help the weaker fur seals to occupy the harem territories vacated by the
sea lions and to spend a normal breeding season (Belkin, 1966a).

The fur seals and sea lions cohabiting in summer could quite often
face competition for food, as many items are common to the two species.
However, in most cases such competition is inconsequential. The fur
seals of the Commander Islands feed mainly on pelagic animals while
the sea lions feed on demersals (Barabash-Nikiforov, 1936). Fur seals
and sea lions are seen swimming along the coasts of British Columbia.
At this time the fur seals are dispersed farther from the coasts and feed

137

mostly on schools of small fish and squids at the water surface; meanwhile
the sea lions remain closer to the coasts and feed on demersal fishes and
octopuses, their primary food. In this region, however, there are about
ten species of animals which serve as food for both fur seals and sea
lions (Spalding, 1964а*).

Population dynamics. The dynamics of the natural populations of
all the fur seals has not been studied. It is perhaps insignificant since
instances of en masse mortality due to natural factors have not been
reported. Very soon after the islands on which rookeries are located
were opened to hunters, herds of fur seals and the populations of all the
three subspecies suffered large variations time and again, as a result of
human intervention.

The rapacious exploitation of the Tyulen’ Island stock began imme-
diately after the island was discovered in the mid-nineteenth century, and
some 100,000 fur seals were caught there during 1852-1855. The stock
was destroyed and only gradually replenished during the 14 years in which
hunting was banned. In 1870, the population was again destroyed and
unrestricted hunting prevented its restoration for a long time. The fur
seal population was so small (not more than 7,000 in 1911) that for
over 20 years (roughly 1895 - 1923) the annual catch did not exceed 500
fur seals. In 1911, the Convention for the Protection of the Northern
Fur Seal was signed and, as a result of rational exploitation, the popu-
lation was gradually reconstituted. This helped to raise the annual catch
to 2,000-3,000 animals by 1942. At the end of World War II, before
transferring the island to the USSR, the Japanese caught over 50,000 fur
seals in four years, which again brought the stock to the brink of disaster.
Besides hunting fur seals on the coast, their hunting at sea flourished
and was most rapacious as more than one-half of the sea catch comprised
females, most of them gestating.

In 1957, the USSR, the USA, Canada, and Japan signed a new Provi-
sional Convention for the Protection of the Fur Seal in the northern part
of the Pacific Ocean, the most important provision of which was a ban
on hunting at sea (a limited catch was permitted by each country exclu-
sively for research). This encouraged the development of a scientifically
organized, rational fur seal industry on the islands where, according to
the rules laid down, only the young males possessing the most valuable
coats were permitted to be killed.

A census was undertaken to determine the populations. Two groups
of northern fur seal bulls and newborn pups could be counted quite
accurately. The former were easy to enumerate in a rookery. The pups
were counted by chasing them with a chain past the counting device.
Simultaneous with live pups, dead ones too were counted and their total

110

138

yielded the tOvai number of mothers. By determining the number of
parents of both sexes, the number of young males that could be killed
was calculated in such a way that the required number of male parents
for normal reproduction was ensured (Dorofeev, 1958, 1960).

The ban on killing females, fixing a rational number of bachelors
that could be killed, and maintaining the natural regime of harem rook-
eries during the breeding season led to a comparatively rapid growth of
the population of fur seals on Tyulen’ Island. By 1960, the total pop-
ulation approached 100,000, and 6-7 years later to 160,000 - 170,000.
Adoption of the principles of rational killing and the ban on hunting
at sea, enabled a steady population growth. An obstacle to the growth

of stock on Tyulen’ Island is the limited free territory for enlarging the

harems, due to the island’s small size.

The fur seal rookeries on the Kuril Islands have always been small
and thus there was no coastal hunting there; but rapacious hunting at
sea led to the near total extermination of the rookeries. After the 1957
Convention, the fur seal stock was gradually restored to about 15,000 by
1966 (Uspenskii, 1955; Klumov, 1957; Belkin, 1965a* ).

Immediately following the discovery of the Commander Islands by
Vitus Bering in 1741, the fur seal stock there began to be exploited.
According to the eyewitness accounts of early investigators, the fur seal
population at that time was in the millions. By 1786, over 64,000 seals
had been killed. Individuals as well as organized companies hunted at
different periods. Each “owner,” before transferring his rights to the next,
tried to extract. the maximum benefit. Moreover, for many years gray
seals, i.e., pups were also killed. The killing of infants depleted the herds
50 much that hunting had to be almost prohibited. From 1843 through
1847, a complete ban was imposed, but this did little to restore the stock.
Later, restricted and rapacious hunting alternated; killing of the young
was banned in 1871 and judicious hunting of bachelors initiated. The
herds gradually began to replenish themselves. But by then hunting at sea
had begun anew and from 1891, in which year an agreement was signed
between the USA and the UK for immediate cessation of hunting in the
eastern part of the Pacific Ocean, there was extreme irrational killing of
the Commander and Tyulen’ populations by the Japanese. As a result
of the combined effect of all these factors, by 1911 the Commander
population had dwindled to a mere 9,000.

After several negotiations between Russia, the USA, the UK (also
representing Canada), and Japan, the Convention for the Preservation of
Fur Seals was concluded for the first time in 1911 and remained in effect
until 1940. During World War II, Japan unilaterally breached this Con-
vention. Although the Convention formally banned sealing at sea, there

112

111

139

was regular rapacious killing in the western part of the Pacific Ocean
during the six months in which the fur seals remained at sea under the
protection of no one. Further, the system of organized coastal hunting
was far from perfected. All these factors contributed to the extremely
slow restoration of the Commander fur seal stock.

After concluding the Provisional Convention of 1957, a gradual
restoration of the stock was achieved by organized hunting. By 1960, the
population of the Commander fur seal had increased to nearly 100,000
and later reached almost 200,000.

On the Commander Islands there are numerous deserted rookeries
and much free territory suitable for the formation of new rookeries.
Therefore, cessation of hunting, or properly organized hunting, could

encourage an increase in the number of rookeries and, perhaps, gradually

restore the fur seal population to its original glory.

The Pribilov Islands were discovered in 1786 but, unlike on the Com-
mander Islands, until 1799 seal hunting was done only by Aleutian emi-
grants. Later, as on the Commander Islands, an avaricious killing of seal
herds began, sharply reducing the animal population. In 1867, Alaska
(and the Pribilov Islands) came under the USA and from 1870 the 0$.
government restricted seal hunting, leasing only under the condition that
killing would be restricted to the islands and that females could not be
killed. Later, the hunting season and the annual quota were fixed. From
1910, the leasing system was abolished and the government established
its control over fur seal hunting. As a result of the measures adopted,

Fig. 78. Young female sleeping on the coast. Bering Island, July, 1967 (photo-
graph by Р.С. Nikulin).

140

the Pribilov stock was far better maintained than the Commander and
Tyulen’ stocks, and by 1911 comprised some 200,000 animals. From the
moment the 1911 Convention was signed, Americans constantly pro-
tected their stock over the migration route and also in the region of
wintering, using armed ships, thereby totally abolishing the carnage at sea
and establishing a quite rapid growth rate of the stock. The population
of the Pribilov fur seal had crossed the 1.5 million mark by the 1950s.

American scientists believed that the population had exceeded the
optimal level by that time, not only preventing further increase of stock,
but also leading to high mortality of the offspring and even to some loss
of adults. As a corrective measure, hunting of females was permitted on
the Pribiloy Islands, which in some years exceeded 40,000. Theoretically,
for a normal state of population, the annual addition of pups should not
exceed 500,000 - 550,000. This objective was achieved by 1966. In order
to maintain the stock at a stable level (about 2 million), hunting quotas
were proposed at approximately 60,000 males and 10,000 - 12,000 females
per year (Nagasaki, 1961; Chapman, 1961; Baker et al, 1963; Dorofeev,
1964; Roppel and Davey,* 1965).

111

Fig. 79. Year-old male fur seal sleeping on water. Bering Island, October, 1958
(photograph by S.V. Marakov).

113

141

The population dynamics of the northern fur seal could serve as a
striking example of the adverse influence of the activity of man on a
flourishing population, and of the possibility of restoring it by rational
utilization of the animal.’

Field characteristics. Adult males are large and females of moder-
ate size. The newborns are black in color, turning gray by the time
suckling ceases. The fur of the older animals is in various shades of
cinnamon-brown. The hair coat of bulls is a dark brown or gray, more
often monochromatic. Hairs on the nape of bulls are long and form a
“mane”. The pinnae are narrow and pointed. The fore and hind flippers
are long and their outer tips without fur; the hind flippers fold under
the trunk.

In summer the animals gather into large herds on the coasts and
form harems. The northern fur seal makes various sounds. Bulls groan
in a deep-drawn bass and during brawls make frequent guttural sounds;
the voice of females and the young sounds from afar like the bleating
of sheep, and that of newborns like the bleating of lambs. In water the
animals remain singly or in small groups; while resting on their back,
they raise the long hind flippers which, from a distance, resemble a sail.

The northern fur seal is readily distinguished from a sea lion, quite
similar to it, by much darker coloration, smaller size, and the absence of
an upturned nose. (V.A.)

Economic Importance

Concurrent with the increment in stock, which commenced after the
1957 Convention, there has been a gradual increase in number of fur
seals caught. By 1967, the total catch of the USSR had reached 20,000.

Because the fur coat is highly prized, seal hunting is of considerable
economic importance. Depending on world market conditions, the furs
are valued on average at US $100 apiece, with some fetching as much as
$160. Apart from the fur, fur seals provide a comparatively large quantity
of byproducts—meat, blubber, and liver. The carcass is used for feeding
ranch-maintained fur animals, while the oil melted from the blubber has
a commercial value. The liver is a source of vitamin A.

The technique for catching fur seals is extremely primitive but has
not changed over the years. The animal is stunned by a blow on the head

5 During 1963-1965 there was a marked decline in all the populations of fur seals,
which sharply reduced the number of bulls, the annual increment in pups born, and the
population of bachelors. This reduced the overall population. The reasons for the decline
have not been ascertained (V.A.).

142

with а club, preferably in the nasal region. More modern methods using
electricity, soporifics, and immobilizing agents are being developed.
Bachelors aged 3-5 years, settled on the fringes of harems or at
some distance from them, are the targets of hunters. In the predawn
hours the hunting party first surrounds the bachelor quarters, thus bar-
ring their access to the sea. The cordoned-off herd (sometimes up to
1,000 strong) is slowly chased aside to the slaughter zone, giving the ani-
mals frequent rests of 5-10 min. If rapidly chased, the fur seal warms
up and can die of heat shock (sunburn). In such animals the hair falls
out and the skin has to be rejected. On arrival at the slaughter zone, the
large herd is separated into small groups of 15-20 animals and graded.
Bulls, females, and small-sized animals are freed in the sea and the rest
clubbed to death as described above. After the blow, the seal is slit open
with a dagger-type knife in the heart region. As soon as bleeding com-
mences, the dead animal is skinned together with the blubber; the skin
at this stage is called a raw hide. Later, the blubber is removed using a
blunt knife. The skins, free of flesh are washed in cold water (sea water)
and salted, using common salt. The preserved skins are packed in drums
and dispatched to the fur factory for making semifinished products.
Seal hunting has acquired proper organization. The hunting period
(June 1 through August 1) has been fixed and the annual quota of kills
determined to ensure the future growth of the stock; killing of females
has been banned. Offspring and parents are regularly counted. If the
number of bulls exceeds the required number (in a restricted rookery),
the excess number is killed. The commercial catch of bachelors is lim-
ited to 16 to 20% of the population of each generation. All of these
measures promote the growth of the stock and help increase the catch
year after year without decimating the population. In this way it is wholly
possible to steadily increase the seal population. However, the decline in
population has greatly reduced the number of animals killed annually to
6,000 - 8,000 in the USSR, and from 50,000 - 60,000 to 32,000 - 35,000 in
the USA. Given such a situation, a partial or even total ban on seal hunt-
ing appears necessary for restoration of the population strength. (V.A.)

SUPERFAMILY OF EARLESS, OR TRUE, SEALS
Superfamily PHOCOIDEA Smirnov, 1908

Family of True Seals
Family PHOCIDAE Gray, 1825

These pinnipeds are extremely diverse in size, ranging from the small-
est to the largest in the order, and are least adapted to movement on

114

143

hard substrates because of their inability to raise the trunk on the hind
limbs (which are turned backward and incapable of bending forward at
the calcaneal joint) and also because of their shortened and weak fore
flippers. The fore flippers have well-developed long claws which are not
strongly uncinate or sharply compressed from the sides. The hind flippers
too have claws, though considerably reduced in some species, especially
among 8-incisored seals. The fore as well as the hind flippers are covered
with hair. The digits of the fore flippers, though enclosed in a common
skin, fold up to the last phalanx and possess relatively high mobility at
the joints, which enables the animal to clutch projections on the ground
and dig in, and to use them to puncture very dense snow or even to
scrape ice. However, the reach of the hand is extremely restricted, unlike
the hind flippers in which, on the contrary, the rays are covered in a bet-
ter developed and elastic skin membrane and are capable of considerable
movement by forming a broad fan-shaped oar.

Ear pinnae are altogether absent. The ear openings are densely cov-
ered and appear as small patches of exposed skin.

The head is usually rounded, shortened; the snout is not pointed but
only slightly compressed dorsoventrally and is covered with many labial
whiskers arranged generally in 5-7 arcuate rows oriented along the line
of the mouth. Their total number on each side of the mouth is about
50-60. With rare exceptions, the whiskers are flattened and have wavy
edges (Fig. 81). Whiskers are also present above the eyes (not more than
seven above each) and on the upper side of the rostrum close to the nos-
trils (One or two on each side). The cutaneous glands are not developed
to the same extent in the different species; the sebaceous glands gener-
ally function well, while the sweat glands are often poorly developed and
are not seen at all in some individuals. A scrotum is absent. The tail is
sufficiently well developed and flattened dorsoventrally.

In general build the species of the family are rather similar, varying
mainly in size and weight, shape of the head, snout projection, structure
of the fore flippers, depth of incision of the hind flippers, growth of the
claws, and number of whiskers.

The skull has a spacious cranium with a very narrow interorbital
space and widely separated zygomatic arches; the crests [sagittal, tempo-
ral] on the flattened upper side of the cranial portion are not developed
and the top profile is usually less curved. The bony tympanic bullae
(bullae osseae) are large and bulging, with a fairly smooth, hemispheri-
cal, sometimes slightly flattened surface of different profiles and a well-
developed bony [external] auditory meatus, which in most (subfamily
of true or 10-incisored seals, Phocinae) has a well-developed bony cav-
ity On its outer side. The mastoid process is relatively small, not fused

115

144

with the paroccipital and not directed downward. Ап alisphenoid canal
is absent. Nasal bones terminate posteriorly in a common wedge-shaped
apex protruding into the anterior portion of the frontal bones (Fig. 7).
The zygomatic bones lack a distinct anterior lower process (Fig. 8).

The molars and most of the premolars (except the first) have two
roots in the vast majority of the species. Teeth crowns are quite com-
pressed laterally and in most cases are complex with additional cusps at
the back and in front of the main cusp. The dental formula varies mainly
in the number of incisors from 3/2 to 2/1. The complete dental formula
for the main group of the family (10-incisored seals) is:

Saal (tard eeeeenl
Se в м

Canines are usually not well developed, but are longer and more mas-
sive predominantly in the large and more predaceous species (elephant
seal, hooded seal, leopard seal, monk seal, and larga.

The cecum is poorly developed, usually not longer than 3 cm. The
glans penis is smooth, without spines. The os penis is faintly curved,
quite thickened near the proximal end, and without transverse forked or
T-shaped tip distally.

The majority of the species have one pair of teats, very rarely
two.

The hair coat after shedding of the juvenile (or embryonic) fur is
short, tough, with weakly developed fur hairs that do not form a distinct
layer of underfur. The hair tufts have only one to five fur hairs 5 to 9 mm
long; the guard hairs are 12 to 18 mm long. The juvenile (embryonic)
coat [Lanugo] with which the pups of many pagophilic species are born
is, on the other hand, extremely luxuriant, dense, and thick, affording
good protection from wind and frost in the severe arctic and subarctic
climate. The juvenile hairs, mainly white, with cream, green, or gray
tones, are sported for not more than four weeks (often for a much shorter
duration); sometimes they are shed in the fetal membranes even before
birth (hooded, bearded, and common seals). After the first molt, the color
of the hair coat changes sharply but it takes several years for the definitive
features of the species to develop. Adults sport a spotted, sometimes
evern sharply contrasting coloration (Greenland [harp] and ribbon seals).
Nevertheless, seals with a wholly monochromatic coloration are not rare
(Baikal seal).

Sexual dimorphism is manifest among many species, mainly in col-
oration, body and skull dimensions, and other features. In some species

116

145

(elephant seal, hooded seal) this dimorphism is very sharply manifest in
body size, weight and structure of the snout.

The size difference between seals of various species is highly sig-
nificant. The largest member of the family (although not typical of our
fauna), 1.е., the elephant seal of the Southern hemisphere (Mirounga
leonina), measures (male) 6 m or more (up to 6.5 m) in length along
the dorsal surface (Lc) and weighs up to 4-5 tons. On the other hand,
most of the small-sized races of the ringed seal and Caspian seal attain
sexual maturity at 120-140 cm length (Lc); their weight at the lowest
feeding level (late spring) drops to 35 -40 kg. In the Okhotsk ringed seal
(akiba) the weight drops, on average, to 24.5 kg (females) and 26.0 kg
(males) (Fedoseev, 1971). In spite of such large differences in dimen-
sions and weight, the length ratio between the extremes is relatively
small (see p. 8) at roughly 1:5, but the weight ratio is quite significant
at 1:200.

In spite of the great similarities in general appearance, the species
of the family differ significantly in mode of life, site selection for breed-
ing and molt, and also in food specialization. Only one species, the
leopard seal (Hydrurga leptonyx), relies on warm-blooded animals for
food and feeds not only on penguins and seals, but also on large fish
(see p. 11). On the other hand, there are species feeding almost exclu-
sively or mainly on invertebrates—planktonic crustaceans (crabeater seal,
Lobodon carcinophaga) or various species of benthos, mainly mollusks
and crustaceans (bearded seal). Many other species of seals live mainly
on fish, some on larger (hooded seal, gray seal, and others), others on
smaller ones (ringed, Caspian, and Baikal seals). Some seals also con-
sume cephalopod mollusks (hooded, gray, Greenland, ribbon, and others)
or small crustaceans (ringed seal, sometimes larga, Greenland seal, and
others). The depth of submergence of these seals also varies. The deepest
divers are the hooded seal (perhaps the elephant seal also), Weddell’s
seal (Leptonychotes weddelli), which dives to a depth of 400-600 m, and
the ribbon seal, and probably even the gray seal.

No sharp differences in the daily behavior and activity of these ani-
mals have been recorded; they move around and feed at any time.

Most species perform significant migrations under the influence
of various factors—seasonal movements of the main food organisms,
changes of temperature conditions consequent to changes in ice
conditions, and also the need for specific conditions for reproduction.
Migration invariably conforms to an exact pattern—easily distinguished
from local movements—with some animals straying far from their usual
habitats (e.g., ringed seal in the circumpolar regions; hooded seal in the

146

White Sea or in the eastern Atlantic far south of the polar circle, and
larga on the shores of China).

True polygamy is rare among the members of this family (most dis-
tinct among the elephant seal, Mirounga leonina). On the other hand,
strict monogamy is likewise not very characteristic of true seals since the
males and females meet for a very brief mating season.

Males do not share the responsibility of raising the pups (as in the
other families). Pups are usually delivered one in a season (twins are
extremely rare). Apparently the pups are capable of displaying complex
forms of higher nervous activity.

The members of the family are distributed in all the seas north of
the subtropics: in the Atlantic and Pacific oceans and everywhere in the
Arctic Ocean, in some landlocked water bodies (Black, Caspian, and
Baltic seas and lakes Baikal, Ladoga, and others) and also in a fairly
broad ocean belt surrounding Antarctica, with some ranges (mainly those
of the elephant seal) projecting northward as “tongues” at places (along
the American, African, and Australian coasts).

The range of the family in the Pacific Ocean region encircles the
western, northern, and eastern fringes of the sea in the form of a narrow
arc. Its boundary passes from northern China and the Korean Peninsula
along the western part of the Sea of Japan, covers the coastal regions of
the northern half of Japan and extends along the eastern flanks of the
Kuril range including the Sea of Okhotsk, continues along the eastern
coast of Kamchatka to the Commander Islands, from where it bends
sharply to the east along the Aleutian Islands and reaches the coastal
belt of the American continent, from where it descends south to southern
California (Fig. 80).

The range of the family in the North Atlantic wholly coincides with
the range of the order (since the members of the family of eared seals
are absent there and the range of the walrus does not extend beyond
the range of the family). The range of the family of true seals in the
Southern hemisphere coincides generally with that of the order but does
not extend as far northward along the South American coasts, stops
short of reaching South Africa and only partly covers the coastal waters
of southern Australia (mainly Tasmania). In the Arctic Ocean the range
of the family coincides wholly with that of the order.

The range of the monk seal underwent the most significant variations
over the historic past while the Caribbean monk seal is totally extinct
(p. 500) and the range of the Pacific Ocean species is restricted to a
negligibly small relict section of the Hawaiian Islands. In fact, even the
European range of the monk seal, which is facing total extinction, is
presently fragmented.

*(пуз4ечо yy) э8иел ay Jo Алерипоа
эаеаол4 эцз sjoidap our usyorg ‘элэчазииэц и1э43поб э4з и! Алерипод Woy LOU oy) рие элэуазниэЧ илэЧлом э43 UI UMOYS $1
(eag yoerlg ou) 14э0х2) эЗиел ay) Jo Атерипо4 и1э41по$ sy], ‘эерюоца ‘s[eas эпл Jo Апшеу ay) Jo э8иел рэ}2п11$109Э3 ‘08 "Зы LIU

oC ° oC

В
oe

ANAT Ы
У.
Ва а Ss |
~ fl Lote
Ait ls |

x

aad : é
Sh om
№

Sas
“et,
. z
„<
т a

118

148

The taxonomic features of the family are so sharp and cover so
organically all the members of the family, at the same time distinguishing
them from all other Pinnipedia, that the isolation and status of the family
cannot be disputed. Although there is a tendency to question the entirety
and rank of the order of Pinnipedia based on serological data (see under
characteristics of the order), the unity of the family as such is beyond
doubt. If, however, the diphyletic origin of the Pinnipedia is supported,
then the family of mustelids (Mustelidae) is evidently closest to the family
of true seals from the canoid (arctoid) carnivores.

The earless seals, Phocidae, originated almost as long ago as the
eared seals, Otariidae. Although the finds of the latter pertain to a some-
what earlier period (Lower Miocene) compared to the true seals (Middle
Miocene), they are not so far apart to assume the origin of one group
from the other. The Lower Miocene Pinnipedia already possessed the
distinct features of the eared seals, Otariidae, and hence could hardly
have been the base stock for the Phocidae.

The prevailing division of the family into subfamilies is extremely
simple and has been generally accepted to date. The number of incisors
forms the basis of the division. Three subfamilies are usually recognized:
1) Phocinae, 10-incisored (1 3) seals covering the genera of ringed
seals and true seals Phoca (including Pusa, Pagophoca, Pagophilus, or
Histriophoca), gray seals (Halichoerus), and bearded seals (Erignathus);
2) Monachinae, 8-incisored (1 5) seals of the subtropical regions of
the Northern hemisphere covering monk seals (Monachus) as also the
antarctic seals Lobodon, Ommatophoca, Hydrurga, and Leptonychotes;
and 3) Cystophorinae, 6-incisored (I 5) seals covering the genera of
hooded seals (Cystophora) and elephant seals (Mirounga).

Some scientists (Gill, 1866, Gray, 1869; Kellogg, 1922; Simpson,
1945; and others) isolate Lobodoninae (Stenorhynchinae) into a dis-
tinct subfamily. These are the antarctic 8-incisored seals (Weddell’s seal,
Leptonychotes; Ross’ seal, Ommatophoca; crabeater seal, Lobodon; and
leopard seal, Hydrurga): There was, until recently, a tendency to place
the elephant seal, Mirounga, in this group (King, 1966), which is hardly
justifiable. There was also a suggestion to eliminate altogether the sub-
family Cystophorinae. In spite of the osteological features which bring
the genus Cystophora (hooded seals) close to the 10-incisored seals (spa-
cious for. lacerum posterius, presence of for. entepicondyloideum, and
so forth) and separating it from the genus Mirounga (elephant seals)
bearing some features characteristic of the antarctic 8-incisored seals
(highly reduced for. lacerum posterius in the skull, the same foramen |
in the humerus, etc.), there are many features (including some in the

149

basal and contiguous zones of the skull) which provide a no less weighty
justification in favor of maintaining the prevailing division of the family.

Various authors assign a different number of genera to the fam-
ily. There is a tendency to enlarge the number of Recent genera to 11
(Smirnov, 1929, 1935), 12 (Ognev, 1935; Chapskii, 1955), and up to 13
(Scheffer, 1958; Chapskii, 1963; King, 1966). The present publication has
adopted the principle of a more extensive interpretation of the concept
of the genus and thus 10 Recent genera are included in the family. Hence,
together with the fossils, the family consists of a total of 22 genera. Of
these, 10 belong to Phocinae, of which three are extant genera (Phoca,
Halichoerus, and Erignathus) and seven fossil (Prophoca, Callophoca,
Gryphoca, Platyphoca, Phocanella, Leptophoca, and Miophoca), nine to
Monachinae (including Lobodoninae), of which five are extant genera
(Monachus, Leptonychotes, Lobodon, Ommatophoca, and Hydrurga) and
four fossil (Pristiphoca, Paleophoca, Monotherium, and Pontophoca), and
three to Cystophorinae, of which two are extant genera (Cystophora and
Mirounga) and one fossil (Mesotaria).

The total number of extant species in the family is 18: eight belong to
the subfamily Phocinae (six in genus Phoca and two in the monotypical
genera Halichoerus and Erignathus), seven to Monachinae (three in genus
Monachus, the rest in the above-listed monotypical genera), and three to
Cystophorinae (Cystophora cristata and two species in genus Mirounga).

The Greenland harp seal (Ph. groenlandica), gray seal (Halichoerus

119 grypus), hooded seal (Cystophora cristata), and the aegialoid (pagopho-
bic) form of the common seal (Ph. v. vitulina) inhabit only the basin of
the North Atlantic and the adjoining portions of the Arctic Ocean; the
Mediterranean monk seal (Monachus monachus) inhabits the subtropi-
cal sections, especially the North African and Mediterranean coasts; and
the Caribbean monk seal (Monachus tropicalis) inhabited the Caribbean
Sea.

The following three species of this family are found in the North
Pacific Ocean: the Hawaiian monk seal (Monachus schauinslandt), ribbon
seal (Phoca fasciata), and two forms of the common seal (Ph. vitulina),
1.е., pagophilic (Ph. у. largha) and pagophobic (Ph. у. kurilensis). The
latter two species partly penetrate the Chukchi Sea also. Moreover, the
northern elephant seal (Mirounga angustirostris) is encountered along the
American coast. Phoca hispida and Erignathus barbatus are common to
the basin of the North Atlantic, seas of the North Arctic Ocean, and the
northernmost and northeastern parts, of the Pacific Ocean. The common
seal, absent in the Arctic Ocean (with the exception of the coastal waters
of Murman), thus enjoys a typical amphiboreal range. Two species of

150

the genus Phoca (Ph. caspica and Ph. sibirica) inhabit the landlocked
reservoirs of the Old World—the Caspian Sea and Lake Baikal. .

Four species of eight-incisored seals (Lobodon carcinophaga, Lep-
tonychotes weddelli, Ommatophoca rossi, and Hydrurga leptonyx) and the
southern elephant seal (Mirounga leonina) inhabit the Southern hemi-
sphere, mainly around Antarctica (in the notopelagic zone).

Many species of the family are of considerable economic importance
as valuable sources of fur, skin, blubber, and meat (for the state animal
farms).

One species (Hydrurga leptonyx) is a predator inflicting damage on
penguins and most other species of antarctic seals on which it survives.
Many seal populations of the genera Monachus and Mirounga have suf-
fered greatly as a result of rapacious hunting.

The fauna of the USSR contains (with the proviso stated on
р. 502) all the three subfamilies and the five genera. Three of the latter
belong to the ten-incisored seals (Phocinae—Phoca, Halichoerus, and
Erignathus), one to the subfamily of hooded seals and elephant seals
(Cystophorinae—Cystophora), and one to the subfamily of monk seals
(Monachinae—Monachus). These five constitute 50% of the genera in
the family. The total number of species of the family in our fauna is 10,16
1.е., 55% of the total number in the family. This constitutes about 3%
of the number of species in the USSR fauna. The range encompasses all
our territorial waters and the adjoining pelagic expanses.

Two species belonging to the subfamilies Monachinae and
Cystophorinae represent rather chance elements of our fauna. One of
them, the Mediterranean monk seal (Monachus monachus), became
totally extinct in the USSR waters of the Black Sea by the early twentieth
century and its casual finds on our coasts represent an exceptional
rarity from the Bulgarian or Anatolian populations whose numbers are
insignificant. Another species, the hooded seal (Cystophora cristata),
though not a regular find, is nevertheless fairly frequent, usually singly, in
the northernmost regions of the White Sea or in the adjoining sections
of the Barents Sea. Barring these two species (genera), there is only one
Soviet subfamily and the number of Recent genera of true seals in our
fauna is only 33% of the total number of genera of true seals in the world
fauna; the number of living species, however, constitutes about 45% of
their total number. Of the total number of species in our mammalian
fauna, the number of Pinnipedia of the family Phocidae known in the
USSR is 2.6%. The range of the family, not including the monk seal,
covers the entire territorial waters of the USSR except the Black Sea.

16 Combining Phoca vitulina and Ph. largha into a single species.

120

120

151

All the other species of the family, apart from the elephant seals and
the eight-incisored seals of the Southern hemisphere, are represented
in our fauna and, except for the two Atlantic species (Ph. vitulina and.
Halichoerus grypus), are of great importance in the field of game hunting.

(K. Ch.)

1 (16).

2 ( 3).

3( 2).

4( 5).

5( 4).

Key to Species of True Seals (Phocidae)

Identification Based on External Features

Upper incisors six (three on each side of jaw). Claws well devel-
oped on fore as well as hind flippers.

Whiskers slightly flattened, smooth (Fig. 81). Fore flippers short
and broad, with almost transversely incised anterior margin;
middle digit (with claw) perceptibly longer than rest. Color fairly
monochromatic, sometimes with dull spots. Dimensions large:
body length of adult not below 200 cm. Two pairs of teats. ...
BEERS Bearded seal, Erignathus barbatus (рр. 166-211)
Whiskers highly flattened, with wavy edges. Fore flippers rel-
atively long, with obliquely incised anterior margin. Length of
first two digits (with claw) longer than third. Coloration of dif-
ferent types. One pair of teats.

Head with long and high snout. Upper contour of its profile
straight or even convex on the nose bridge. Cheek teeth [post-
canines] quite large and generally with a single cusp. Color dull
and spotted, brighter on breast and flanks than on back. Body
length of adults in a straight line over 150 cm. ..............
Gray or long-snouted seal, Halichoerus grypus (pp. 454-495)
Head with moderately long, low snout. Upper contour of its
profile with fairly distinct small break in anterior portion of
forehead (on nose bridge). Crowns of cheek teeth, especially of

Fig. 81. Structure of the cheek whiskers of seals. Above—smooth whiskers of the
bearded seal; below—more flattened whiskers with wavy edges in the rest of the
true seals (figure by К.К. Chapskii).

152

6 (13).
7( 8).

121

8 7).

9 (10).

10 ( 9).

11 (12).

12 (11).

lower ones, with additional cusps set in front and back of main
tooth. When the anterior cusp is absent on lower teeth, main
cusp is not pointed and teeth in general are relatively small.
Color and dimensions vary. Body length not above 150 cm.
Color of hair coat usually dark and spotted, dorsally with
fairly sharp light-colored streaks or speckled, or altogether
monochromatic, dark.

Teeth relatively massive. Majority of lower cheek teeth with
longitudinally elongated crown; cheek teeth in relation to gen-
eral row of teeth often set obliquely; accessory cusps some-
what inclined toward the middle (main) cusp, with which bases
merge (Fig. 82). Color of hair coat spotted, with light-colored
streaks, or speckled with light-colored background. Body length
of adults in a straight line not less than 140 cm..............
......... Common seal, larga, Phoca vitulina (pp. 307-369)!’
Teeth not large, relatively thin. Lower cheek teeth usually with
highly disjointed crown, accessory cusps not inclined toward the
base (Fig. 83). Color of hair coat dorsally monochromatic or
spotted, with or without light-colored streaks. Body length in a
straight line less than 140 cm.

Cheek teeth with vertically disposed high accessory cusps.
Color monochromatic. Light-colored gaps or dark-colored spots
АЕ. Baikal seal, Phoca sibirica (рр. 290-306)
Cheek teeth with accessory cusps directed forward and backward
(fanlike). Color of skin not monochromatic, with pattern.
Color of skin with dark main background of back (ventral side
often paler), interrupted by light-colored streaks for the most
part in form of whole or broken lines. .....................
ОЕ, Ringed seal, Phoca hispida (рр. 218-260)
Color of skin variegated, dove-gray, dorsally mainly with light-
colored streaks but rarely assuming shape of closed rings, with
innumerable spots of diverse sizes and shapes, dark and some-
times almost black, and scattered haphazardly. ..............
о о Caspian seal, Phoca caspica (рр. 260-290)

See

121 Fig. 82. Structure of the crowns of upper and lower cheek teeth of the common

seal, Phoca vitulina (figure by K.K. Chapskii).

17 See note at the end of this Key.

153

не а

121 Fig. 83. Structure of the crowns of lower (cheek) teeth in the small species of
the genus Phoca. A—Baikal seal, Ph. sibirica; B—ringed seal, Ph. hispida;

13 (6).

14 (15).

122 15 (14).

C—Caspian seal, Ph. caspica (figure by K.K. Chapskii).

Large light-colored, almost white sections alternate with dark
brown, often nearly black sections in hair coat of adults. Color
of young ones not intensely spotted or altogether without spots,
dull gray; in animals of transitional age, young and adult, rudi-.
ments of above pattern seen vaguely (like a shadow). Light-
colored streaks totally absent in all stages.

Adults with well-defined (or diffused at edges) large dark-
colored, paired wing-shaped patches that stand out prominently
against light-colored background dorsally along flanks. Young
animals gray, with haphazardly scattered, large (angular) and
minute spots. Lower cheek teeth (apart from the first) with
well-developed accessory cusps and pointed main cusp (Fig. 84).
м. Greenland seal, Phoca groenlandica (pp. 369-436)
Adults with four light-colored closed bands against dark-colored
background: around neck, around base of fore flippers, and in
lumbar portion. In young animals dorsal side dull brown, with
distinct boundaries of dark field at site of future bands. Lower
cheek teeth small, with poorly developed accessory cusps (often
only one) and’ obtuse main Cusp: а...

Pe Ante Pa alte tea atl ae Ribbon seal, Phoca fasciata (pp. 436-454)

122 Fig. 84. Form of teeth crowns in the lower jaw. Bottom—Greenland seal, Phoca
(Pagophilus) groenlandica; above—ribbon seal, Phoca (Histriophoca) fasciata

(figure by K.K. Chapskii).

123

122

154
16 ( 1).

17 (18).

18 (17).

1 (16).

2 (3).

Upper incisors four (two on each side of jaw); color either dull
and dark or bright and spotted.
Lower incisors four (two on each side of jaw). Cheek teeth
extremely massive, often set obliquely toward general line of
tooth row. Color dull, brown, sometimes with large, nearly
rectangular patch in lower half of body. Whiskers smooth. Two
ран ор ее
Mediterranean monk seal, Monachus monachus (рр. 502-515)
Lower incisors two (one in each half of jaw). Crowns of cheek
teeth not large. Color of main background gray, covered with
rather rare, irregularly shaped, nearly black patches. Whiskers
flattened, with wavy edges. One pair of teats. ...............
.... Hooded seal, Cystophora cristata (pp. 524-547) (K. Ch.)

Identification Based on Skull Features

Three incisors on each side of upper jaw (total of upper and
lower incisors 10). Lateral contours of rostral part of skull,
viewed from above, reveal perceptible convexity anterior to
orbits (Fig. 85).

Alveoli of upper incisors with circular opening. Zygomatic
bones greatly reduced and broad, their smallest length (without
processes) exceeding smallest width by not more than 1.5 times
and smaller than interorbital space. Cheek teeth (when not
worn)-with poorly developed accessory cusps, firmly fused with
main cusp. Tympanic bullae flattened; their outline viewed from

Fig. 85. Rostral-facial part of the skull of seals. A—ribbon seal, Phoca (Histrio-
phoca) fasciata (lateral contours with convexity); B—hooded seal, Cystophora
cristata (concave lateral contours) (figure by K.K. Chapskii).

123

2)

4 (13).

5( 6).

6 ( 5).

155

below, resembles а шаре. .... ее:

Е Bearded seal, Erignathus barbatus (рр. 166-211)
Alveoli of upper incisors distinctly compressed lateratly. Small-
est length of zygomatic bones (without processes) more than
1.5 times their smallest width and more than interorbital width.
Cheek teeth with or without well-developed accessory cusps.
Tympanic bullae sharply convex and, viewed from below, reveal
more or less oval outline.

Lower posterior process of zygomatic bone considerably longer
than upper. Posterior edge of bony palate with deep arcuate or
angular notch. Bony septum in internal nares does not extend
beyond two-thirds of longitudinal suture between palatine
bones.

Skull, viewed from above, does not reveal infraorbital foramen,
which is concealed by carinate crest on orbital side of zygomatic
process of upper jaw (Fig. 86). Crowns of most cheek teeth of
upper jaw with single cusp (without accessory cusps posteriorly),
Upper contour of skull profile straight; anterior portion of skull -
in zone of nasal bones in adults almost as high as occipital por-
tion. Outline of nasal opening [nares] perceptibly broadened in
UPPER расе В
Gray ог long-snouted seal, Halichoerus grypus (pp. 454-495)
Sharp carinate crest absent on anterior wall of orbit and infraor-
bital foramen distinctly visible. Crowns of cheek teeth (except
the first) with additional cusps (at least one posterior to the
main). Upper contour of skull profile appears as a convex line;

Fig. 86. Position of the infraorbital foramen. A—seal of the genus Phoca; В and
С-—ргау seal, Halichoerus grypus: 1—anterior lower edge of orbits in all seals of
the genus Phoca $. lato; 2—crest. Infraorbital foramen (shown by arrow) in seals
of the genus Phoca viewed from above and in the gray seal covered from above
by crest (B) and seen only from lower side (C) (figure by K.K. Chapskii).

123

124

156

С»

8 ( 7).

9 (10).

10 ( 9).

Fig. 87. Contour of nostril. A—larga, Phoca vitulina [агейа]; B—gray seal, Най-
choerus grypus (figure by K.K. Chapskii).

facial part, even among adults, perceptibly below the occipi-
tal. Contour of upper part of nares not more broadened than

middle and lower parts.

general tooth row.

rostral part of skull less than length of orbit.

18 See note at the end of this Key.

Teeth relatively large; longitudinal diameter of alveolus of
canine and narrowest part of interorbital space (from above)
exceed maximum width of infraorbital foramen. Most of lower
cheek teeth with longitudinally elongated crown, quite massive
at base and not deeply divided; all cusps closely fused with
bases while accessory cusps are inclined toward the main. Cheek
teeth often set obliquely in relation to general line of tooth row
(Fig. 82). Common seal, larga, Phoca vitulina (pp. 307 -369)!®
Teeth relatively small; longitudinal diameter of alveolus of
canine and narrowest part of interorbital space less than width
of infraorbital foramen. Lower cheek teeth not massive but with
relatively high and deeply cloven crown; their accessory cusps
not inclined toward the main cusp; set straight in relation to

Tympanic bullae small and widely set, their length less than gap
between them. Bony lobe of external auditory meatus narrow:
its width less than gap separating it from crest of articular fossa
(Fig. 88). Length of rostral part of skull exceeds length of orbits
м er ois Caspian seal, Phoca caspica (рр. 260-290)
Tympanic bullae more or less large; their length not less than
gap between them. Bony lobe of external auditory meatus wider
than gap separating it from crest of articular fossa. Length of

157

124 Fig. 88. Bony lobe of external auditory meatus (1) and gap (2) between it and
crest of the articular fossa (3). A—ringed seal, Phoca hispida; B—Baikal seal,
Ph. sibirica and Caspian seal, Ph. caspica (figure by K.K. Chapskii).

124 Fig. 89. Ratio between length of the orbit (1) and length of the rostral (preorbital)
part of the skull (2) in the Caspian seal, Phoca caspica (figure by K.K. Chapskii).

11 (12). Lower cheek teeth closely set, with high, nearly parallel (like
crests) accessory cusps (Fig. 83, a). Anterior edge of nasal bones
without median projection. Length of tympanic bullae roughly
equal to gap between them or slightly more. ................

а Baikal seal, Phoca sibirica (рр. 290-306)

12 (11). Lower cheek teeth (except the first) with deflected (fanlike)

anterior and posterior cusps, considerably shorter than middle

125

126

158

13 ( 4).

14 (15).

15 (14).

16 ( 1).

17 (18).

18 (17).

(main) one. Anterior margin of nasal bones with median pro-
jections. Length of tympanic bullae perceptibly more than gap
between them. .......... Ringed seal, Phoca hispida (p. 218)
Lower posterior process of zygomatic bone not longer than
upper posterior. Posterior edge of bony palate without deep
notch. Bony septum in internal nares can extend far backwards,
even up to posterior edge of palate.
Lower cheek teeth (except the first) relatively large, with well-
developed acute cusps anterior and posterior to the main one,
which is also pointed (Fig. 84, a). Posterior edge of bony palate
without notch in midportion, sometimes with posterior projec-
tion. At least base of bony septum in choanae reaches poste-
rior edge of bony palate. Tympanic bullae not bent (Fig. 162).
Their external auditory meatus geniculately bent forward.
ее .. Greenland seal, Phoca groenlandica (рр. 369-436)
Lower cheek teeth small, with obtuse rounded main cusp and
poorly developed accessory cusps, of which anterior one gen-
erally absent. Posterior edge of bony palate faintly notched,
usually in form of shallow braces. Compact bony septum in
choanae rarely reaches posterior edge of palate. Tympanic bul-
lae strongly bent (Fig. 178). External auditory meatus not genic-
ulately bent. ...... Ribbon seal, Phoca fasciata (pp. 436 - 495)
Two incisors in each half of upper jaw (upper and lower incisors
not more than eight). Lateral contours of rostral part of skull,
viewed from above, concave anterior to orbits (Fig. 85).
Two incisors on each side of upper and lower jaws (eight incisors
in all). Nasal processes of premaxillary bones reach nasal bones.
Longitudinal bony septum in internal nares does not reach pos-
terior margin of bony palate, which has a sharp notch. Posterior
lacerate foramen far short of reaching basal suture. .... Medi-
terranean monk seal, Monachus monachus (pp. 502-519)
One incisor on each side of lower jaw (six incisors in all).
Nasal processes of premaxillary bones far short of reaching nasal
bones. Longitudinal bony septum in internal nares reaches end
of bony palate, which has no notch. Posterior lacerate foramen
extends far anteriorly, almost up to basal suture (Fig. 85): ...
и Hooded seal, Cystophora cristata (p. 524). (К. Ch.)

Note: There is a view (Chapskii, 1967, 1969) that the Pacific Ocean pop-
ulations of the common seal, Phoca vitulina, usually regarded as a sub-
species, represent, in fact, two independent species: one, the conspecific
Atlantic, i.e., the common seal (Ph. vitulina) and the other the specific

125

125

159

Pacific species, 1.е., the seal associated with ice floes or the “pagophilic”
larga (Ph. largha). This aspect calls for further investigation. In this book,
however, the prevailing broader interpretation of the species vitulina
(q.v.) has been adopted. The following are the main features distinguish-
ing the two groups of seals:

a) Transverse profile of tympanic bulla with high and fairly steep
curvature toward the base of the lobe of the external auditory meatus

(Fig. 90); fork of the posterior edge of the zygomatic bones arcuate

(Fig. 91); nasal processes of the premaxillary bones adjoin the nasal
bones over a considerable distance (Fig. 92). Posterior edge of the bony
palate usually arcuate (Fig. 93). Color of skin variegated: either gray
with whitish, often annular streaks and numerous dark spots or dabs, or
brightly speckled with light colored underside. ....Larga, Phoca largha

b) Transverse profile of tympanic bulla with low steplike interruption
toward the base of the lobe of the external auditory meatus (Fig. 90);
fork of the posterior edge of the zygomatic bones in the form of an angle
(Fig. 91); nasal processes of the premaxillary bones only just barely or
do not reach nasal bones (Fig. 92). Posterior edge of bony palate usually
in the form of braces or an angle (Fig. 93). Color of the skin darker,
with spots and clear spaces. .... Common seal, Phoca vitulina. (K.Ch.)

Fig. 90. Transverse contour of the tympanic bulla (1) and bony lobe of the exter-
nal auditory meatus (2) partly covered by the articular fossa (3) in different forms
of the common seal, Phoca vitulina, as seen in the lower part of the upturned
skull anteriorly (figure by К.К. Chapskii). A—pagophobic forms of the com-
mon seal: Atlantic, Phoca v. vitulina; Kuril (Island), Ph. v. Kurilensis; Richard’s,
Ph. v. richardi; B—pagophilic form of seal: Far-Eastern larga, Ph. v. largha.

Fig. 91. Articulation of zygomatic bone with zygomatic process of temporal bone

(1): angular notch on the posterior edge of the zygomatic bone in the pagophobic

form, Ph. v. vitulina, and arcuate in pagophilic form (larga), Ph. v. largha (figure
by КК. Chapskii).

125

127

160

nny

N

>
А В

Fig. 92. Nasal and ргетахШагу bones in different forms of the common seal,

Phoca vitulina. A—Atlantic common seal, Ph. у. vitulina; B—larga, Ph. у. largha

(figure by К.К. Chapskii): 1—nasal bone; 2—premaxillary bone; 3—facial part
of nasal bone; 4—maxillary part of nasal bone.

Subfamily of True, or 10-incisored, Seals

Subfamily PHOCINAE Gill, 186619

These are large-, moderate-, or small-sized seals. Body length [in adults]
including the tail along the dorsal curvature (Lc) varies from 100 to
almost 300 cm.

The hind flippers are longer than the fore flippers and both possess
well-developed claws. On the fore flippers the third digit is somewhat
shorter than the two preceding ones; it is noticeably longer than all the
rest Only in one species (bearded seal, Erignathus barbatus). A proboscis-
like growth is absent in the anterior upper portion of the snout. The
whiskers are usually highly flattened, with wavy edges; they are smooth
and slightly compressed only in one genus (bearded seal).

The teats are one pair, except in the bearded seal in which there are
two pairs.

The skull is usually not particularly large and is highly compressed
between the orbits which, in most of the species, with rare exceptions, are
very large. The maxillary bones directly anterior to them bulge markedly.

19 Strictly speaking, Gray (1825, 1850, 1866) enjoys the priority of classifying the true
seals (Phocidae) into subfamilies as also of establishing the subfamily Phocinae. He desig-
nated this subfamily as “Subfamily 2. Phocina”. His error in formulating this subfamily, i.e.,
inclusion of the walrus in it, was not an exception among the works of that time. Thus, Gill
(1866), to whom the formal priority was assigned for the classification of the subfamily,
Simpson (1945), and Scheffer (1958) also committed an error by including the monk seal
in the subfamily (K. Ch.).

126

161

А

Fig. 93. Structure of the palato-choanal region in different forms of the common

seal, Phoca vitulina. A—Atlantic common seal, Phoca у. vitulina; B—larga, Phoca

v. largha (figure by K.K. Chapskii): 1—contour of the posterior margin of the

bony palate; 2—uncinate process of the pterygoid bone; 3—horizontal part of
the palatine bone.

The preorbital processes are absent or highly reduced. The outer edge of
the external auditory meatus, with the exception of the genus Erignathus
(bearded seal), terminates with a bony lobe or is differently structured
(Greenland [harp] seal, Phoca groenlandica). The anterior edge of the
nasal bones is dentate and, when the median prominence is absent, with-
out an inverted angular notch. The nasal processes of the premaxillary
bones adjoin the nasal bones and are usually wedged between them and
the upper maxillary bones or, in a few cases (gray seal, Halichoerus gry-
pus, and to a lesser extent in the pagophobic members of the subgenus
Phoca s. str.), slightly short of reaching them. The posterior lacerate
foramen with one exception (genus of bearded seals, Erignathus), extends
considerably forward along the inner side of the tympanicum.

There are three incisors in the upper jaw and two in the lower jaw
on each side. The dental formula is:

3 1 4 1 20
15, Ст, Ра, т

The crowns of the cheek teeth when worn down, with one exception
(gray seal, genus Halichoerus), with well-developed accessory cusps while
these teeth (except for the first premolar) have two roots in most of the
species (except in the foregoing genus and also quite often in the ribbon
seal, subgenus Histriophoca of genus Phoca).

The postnatal hair coat is usually of the “embryonic” [lanugo] type,
with dense, long fur, and is shed toward the end of lactation. Among

20 Sometimes a second molar is seen.

129

162

the pups of the pagophobic members of the subgenus of ringed seals,
Phoca $. str. (genus Phoca) and the bearded seal (genus Ervignathus), the
embryonic coat is shed in the mother’s womb itself just before birth. In
the former case it is white (or creamy) and in the latter, brownish-gray.

Most of the forms are pagophilic but some (gray seal, genus Най-
choerus, and the pagophobic form of the subgenus Phoca s. str., genus
Phoca) whelp and molt on land.

Food specialization is not particularly distinct: it is most pronounced
in bearded seals, genus Erignathus (benthic feeders) and Phoca sibirica
(consuming mainly gobies and sculpins (Baikal oil-fish)) but specializa-
tion is not complete even among them.

These animals are distributed in the cold and moderate belts of the
North Atlantic and North Pacific oceans, in the Arctic Ocean, and also
in some landlocked water bodies of the Old and New World (Fig. 94).

Phocinae, with several features of a more primitive nature still pre-
served, apparently evolved earlier than the other subfamilies. Although
their oldest finds date back to the Upper Miocene, the actual appear-
ance of the group Phocidae should be placed in a much earlier period,
perhaps the Early Miocene or even Oligocene. The following arguments
indirectly support this view.

Firstly, even much earlier finds of Phocidae are known but presum-
ably placed among Cystophorinae (see p. 262).

Secondly, the fossil finds of Phocinae from the Sarmatsk formations
(Upper Miocene) represent the fully developed forms of the subfamily.

Thirdly, eight-incisored seals coexisted with them (see p. 498); the
former could not have evolved simultaneously with the Phocinae.

The landlocked water bodies, i.e., the seas of the Old World, some-
where in the Tethys and its sources, could perhaps be regarded as the
zone of evolution of the Phocinae.

By the beginning of the present century the system of the subfamily
was quite well established but became the subject of dispute in the mid-
1960s. The debate centered around the suggestion that the members of
the subfamily of hooded seals or the 6-incisored seals (Cystophorinae)
be included in it and doubts as to the affinity of the genus of bearded
seals, Erignathus, to this subfamily (King, 1966).”! Evidently, the classic
interpretation of the subfamily has been favored (see p. 148).

21 The karyotype of the bearded seal (2 п = 34, NF = 66) is well distinguished from
that of the Baikal (Ph. sibirica), Caspian (Ph. caspica), and Greenland (Ph. groenlandica)
seals (2 n = 32, NF = 62). It coincides with the karyotype of Weddell’s seal (Leptonychotes
weddelli) from the subfamily of monk seals, Monachinae (Anbinder, Mlekopitayushchie
[Mammals], Novosibirsk, 1970) (V.H.).

163

0

‘(ys
Osi [4
Bead г A
4
oc \

dey) 14
nie о $

БАС”

и у

‘ i :

a lV-6 Е LY К
LS Bd 2 р t
В. wa

130

164

The subfamily comprises 10 genera (45.5% of the total number of
genera in the family) of which seven are extinct and three extant (about
14% of all the extant genera of the family): Phoca Linnaeus, 1758; Erig-
nathus Gill, 1866; and Halichoerus Nilsson, 1820. The total number of
extant species is 8, which constitutes 44.5% of the total number (18) of
the Recent species of the family.

The economic importance of the subfamily is significant. Some
species (Greenland [harp] seal, Phoca groenlandica, ringed seal, Ph.
hispida, and Caspian seal, Ph. caspica) form the basis of the marine-game
industry. Their importance is primarily due to the fact that these animals
are concentrated in large numbers in some seasons, making it possible
to collect large quantities of hides of high commercial value (from which
furs of various types are produced) as also other raw materials (blubber
and meat for animals).

Three genera (100% of the genera of the subfamily) with 8 species
(100% of the species of the subfamily) are found in the USSR. These
are distributed in all the seas surrounding our country, except the Black
Sea, and in some landlocked water bodies such as Lakes Ladoga and
Baikal and the Caspian Sea. All the species, especially the Greenland
and Caspian seals, are of commercial importance. (K. Ch.)

Genus of Bearded Seals
Genus Erignathus Gill, 1866

1866. Erignathus. Gill. Proc. Essex. Inst., 5: 5, 9. Phoca barbata Fabricius
= Phoca barbata Erxleben.

These are seals of large dimensions with a massive trunk, relatively
small head, and a somewhat shortened neck.

The snout is moderately stretched, the eyes are relatively small, and a
narrow fringe of bare skin is seen around the nostrils. The extremely fluffy
upper lip has abundant (over a hundred) long, thick, smooth whiskers
(without wavy edges). The hand has a very long middle digit, protruding
noticeably farther than the rest.

The skull has a spacious cranium, relatively small orbits, and poorly
developed zygomatic arches. The interorbital area is not highly com-
pressed and the rostral part is broad. The zygomatic bones are sharply
reduced and broad; the length of the bone without the processes is only
slightly more (not more than 1.5 times), or even does not exceed, its
smallest width. The tympanic bullae, viewed from below, appear nearly
trapezoidal, with a distinct inner posterior angular projection in which

130

165

the carotid foramen is disposed. The nasal processes of the premaxillar-
ies reach the nasal bones and extend along them quite far behind their
anterior edge. A longitudinal bony septum in the internal nares (within
the palatine bones) is absent.

The dental formula, as in other seals of the subfamily, is:

3 1 + 1
5, ет, Pin M poo

The upper incisors have fine roots, circular in cross section, their
alveoli are also correspondingly circular in cross section. The crowns of
the premolars and molars, excluding the initial ones, have one (posteri-
orly) or two (anteriorly and posteriorly) obtuse accessory cusps (Fig. 95).
With the exception of the first premolar, all the cheek teeth have two
weak roots which are set apart. The teeth wear out and fall out early.

The body is rather monochromatic although the underside is usually
somewhat lighter than the upper; young animals have fine spots. The
primary, prenatal pelage is not of the embryonal type but cinnamon-
brown (sometimes incompletely) just before the pup is born. Age-related
changes are minor and mainly manifest in body size and partly in col-
oration, as well as in the structure and proportions of the skull. Sexual
dimorphism is negligible.

Two pairs of teats are present.

These seals are biologically associated with ice floes on which they

- reproduce and molt and also with shallow waters since they feed mainly

on benthic invertebrates. They do not form large groups on ice floes.
Migrations do occur but are not very distinct. Mating does not involve
the formation of harems and there are no special fights among the adult
males.

The animals are distributed in the circumpolar region in the seas
with an arctic regime and inhabit the periphery of the Arctic Ocean,

B
8.2 Oe ©.

A

Fig. 95. Structure of the crown of undamaged lower teeth in a young bearded
seal, Erignathus barbatus. A—from the inner side; B—from the outer side (figure
by К.К. Chapskii).

131

166

northern edge of the Atlantic Ocean, and the northwestern part of the
Pacific Ocean (Bering Sea and the Sea of Okhotsk).

Among the 10-incisored seals, the genus Erignathus occupies a spe-
cial position. Several morphological characters preserving features of
relative primitivity, to some extent the result of adaptation to benthic
feeding, prompted the inclusion of this genus under a distinct mono-
typic tribe, Erignathini (Chapskii, 1955). The taxonomic independence
is confirmed by data from serological investigations (V.I. Borisov) as
also by cytogenetic data (the diploid chromosome number in bearded
seals is 2 п = 34 and in Phocinae, 2 п = 32). Both these criteria com-
pel us to place the genus Erignathus somewhat away from the rest of
the 10-incisored seals in spite of considerable antigenic and cytogenetic
similarity. At the same time, its genetic relation with other members of
the subfamily are not yet quite clear. It is not clear to which genera of
Phocinae Evignathus is most closely related. It is only evident that its
predecessors separated from the main branch of true seals very long ago.
Whether or not the genus Ervignathus serves to some extent as a link
for all the typical northern seals (Phocinae) in the same manner as the
genus Monachus serves as the initial form for the formation of the typ-
ical (1.е., 8-incisored) seals of the Southern hemisphere is very difficult
to state; the origin of the genus beyond the Pliocene has not yet been
traced. Its phyletic lineage derives from the Pliocene form Platyphoca of
the Belgian formations.

The North Atlantic basin is evidently the center of origin of the
genus. The ratio between the extinct and extant species is 1:1.

The genus includes only one species, the bearded seal, Erignathus
barbatus (Erxleben, 1777).

The range of the genus in the USSR evidently encompasses the
whole of the Arctic and the Far Eastern seas (north of the Sea of
Japan).

This species is of much importance as a target of hunters, at the
same time posing no danger to the fishing industry. (K. Ch.)

BEARDED SEAL
Erignathus barbatus (Erxleben, 1777)

1776. Phoca barbata. Muller. Zoologiae Danicae prodromus. Nomen
nudum.

1777. Phoca barbata. Erxleben. Systema regni animalis. I, p. 590. Waters
of southern Greenland (Ognev, 1935).

1778. Phoca leporina. Lepechin. Acta Acad. Scient. Imper. Petropoli-
tanae. I, p. 264, Tab. 8, White Sea.

132

133

167

1811. Phoca nautica. Pallas. Zoographia Rosso-Asiatica, I, р. 108. Sea
of Okhotsk.

1811. Phoca albigena. Pallas. Ibid., р. 109. Kamchatka.

1817. Phoca lachtac. Desmarest. Now. Dict. Sc. Nat., 25, p. 581. Pacific
Ocean.

1828. Phoca parsonsii. Lesson. Dict. Class. d. Hist. Nat., 13, р. 414.
Northern seas. (V.H.)

Diagnosis
Only species of the genus.
Description

Body heavily built; head and flippers (especially the fore flippers), com-
pared to body length and weight, are not very large (Fig. 96). The hands
are short and broad as though truncated anteriorly. The fore flippers
are markedly shifted forward and set relatively closer to the anterior end
of the body than in other seals. Their claws are massive, with a quite
distinct, transverse, age-related structure, i.e., ribbed. The whiskers are
abundant, luxuriant, and set in 10 or 11 rows; five rows on the lower
side have 11 to 18 whiskers each. Their total number on each side of the
snout reaches 125.

The hair coat of the adult is perceptibly sparser than among other
seals. In the young, however, it is dense and uniform, and the skin
between the hairs is not visible. The structure of the hair coat is quite
similar to that of other seals. Three categories of hairs are distinguish-
able: guard, intermediary, and soft fur. Their quantitative ratios vary
perceptibly with age. Among adults the hair coat consists 50% of guard
hair, 30% of intermediary hair, and about 20% of fur. In pups, after
shedding the embryonic pelage, the proportion of guard hair is less
than 10%, intermediary hair slightly more than 40%, and the rest is
fur (V.A. Potelov). The hair on the trunk is sparser and long but on the
head, especially on the snout, it is short and dense; the nature of the
coat on the flippers, especially on the hind flippers in the region of the
ankle joint and on the extreme digits, is nearly similar.

The predominant color of the upper portion of the trunk is a
brownish-gray or blackish-olive, gradually turning lighter, into light gray
or dirty olive on the flanks and on the ventral side. Quite often, animals
with a light and more monochromatic pale ash coloration, almost devoid
of spots, are encountered. Along the median dorsal side, from head
to tail, runs a barely visible narrow band with rather blunt edges,
with a dark, sometimes almost black or slate-black coloration (“belt”).

132

168

Fig. 96. Bearded seal, Erignathus barbatus (photograph by С.М. Kosygin).

Somewhat large but usually very indistinct dull whitish spots with fine
uniform speckles against the background color are seen roughly from
the level of the shoulder blades farther to the end of the body dorsally
(Fig. 97). In some cases, mainly in adult animals, the spots are apparently
very distinct and bright, sometimes almost white, devoid of variegation
and distinctly visible from afar. Further, small dull spots, slightly darker
than the main background, are visible in places. Though few, these spots
catch the eye.

The head displays characteristic coloration in the form of whitish
patches that are prominent in the overall dark gray or even gray back-
ground (Fig. 98). One very large patch is located on the sinciput [fore-
head] and two pairs above the eyes and around the ear openings. The
whiskers are light in color, mainly muddy-white. The claws are dark gray,
almost black.

No color differences have been recorded in the hair coat between
males and females. Age-related color changes have not been fully stud-
ied. The skin of newborn pups, compared to that of adults, is usually
much darker, blackish-gray, sometimes light brown, with minor differ-
ences in color intensity between the upper and lower sides of the body.
In most cases the flanks and the upper portions, more rarely the ven-
tral side of the body, have numerous speckles, fused at places, usually

169

132

134

Fig. 97. Bearded seal. Devich’ya Luda Island, Kandalaksh Strait, White Sea,
August, 1962 (photograph by V.D. Kokhanov).

more abundant in the anterior part of the body. They extend somewhat
onto the head and fore flippers. The light-colored pattern on the head
(sincipital and supraorbital spots and also the lighter parts on the lips)
stands out contrastingly. The older juveniles and animals of transitional
age gradually lose the speckled pattern and become more monochro-
matic in coloration, with warm brown shades somewhat more promi-
nent. Sometimes the difference between the underside and the upper is
smoothened but, nevertheless, the upper side is quite frequently dark-
colored; a narrow dark band extending along the spine from the head to
the tail is nonetheless visible. The color of adults is paler, monochromatic
(but nevertheless usually more vivid on the spine), sometimes altogether
light-colored, silvery-gray or pale olive. The pattern on the head becomes
diffuse and almost disappears. Quite often, brownish, even rust-red tones
appear around the head, on the throat, and on the neck.

Coloration is subject to considerable individual variation. The large
light-colored spots on the trunk, tiny surface spots (dabs), and also the
light-colored pattern on the sinciput appear differently in different ani-
mals. The vividness of the color of the dorsal and ventral sides likewise
varies markedly.

In some specimens the dorsal whitish spots are barely visible while
the granular dark speckles are widely scattered and sometimes even

133

135

170

Fig. 98. Color of the head of a five-year-old bearded seal. Barents Sea, May,
1963 (photograph by V.A. Potelov).

totally absent; in other animals they are brighter, sometimes unusually
distinct and visible from a distance; they are also variable in size, dis-
position, number, and shape. The same is true of the spots/dabs on the
sinciput. Their number, shape, distribution, and color are highly vari-
able. Animals with no spots/dabs whatsoever are also encountered; in
most animals, however, these are disposed along the body flanks and are
absent ventrally, but specimens with a large number of ventral specks
(appearing mottled as a result) are also encountered.

The color of the hair coat of adults evidently undergoes seasonal
changes (fading). The color is a dark gray in autumn, gray in winter, and
yellowish-gray in spring (V.A. Potelov).

The geographic variation in color is not well understood.

The skull features supplementing the generic craniological charac-
teristics are as follows. Because of the small dimensions of the orbits
and evidently the shortened zygomatic bones, the width at the zygomatic

134

УТ

arches usually does not exceed that of the cranium at the mastoid pro-
cesses. The interorbital space is broad, without a sharp constriction, and
flattened at the top; its width at the narrowest point is about one-fifth the
mastoid width. The upper profile of the skull descends along a smooth
S-shaped line from the apex of the nasal bones to the base of the nares.

The infraorbital foramina are large, vertically elongated, and their
longer diameter slightly more than the longitudinal length of the alveolus
of the upper canines. A longitudinal crest extends on both sides of the
skull along the upper edge of the temporal bone; a notch is perceptible
between this crest and the base of the zygomatic process (Fig. 99). The
nasal bones are relatively broad; their posteior wedge-shaped end bears
a round apex which is not sharply pointed while the anterior edge has
long lateral projections between which a small, often slightly bifurcated,
median projection is seen in most cases. At the point of juncture of the
premaxillaries with the nasals, an angular notch is not formed on the
upper posterior edge of the nares, which appear oval in shape from the
front. The jugular processes are quite massive and bent backward sharply.

4 : k
go Nati oy

у сте ни т «Ру:
Se

SS

>

SNS
SS

=
- SOL,
ae

м

Fig. 99. Skull of the bearded seal, Erignathus barbatus (figure by М.М. Kondakov).

136

172

The posterior edge of the bony palate has a small, broad, slightly angular
notch.

The upper tooth row has a sharp S-shaped curvature in the hor-
izontal plane. The lateral (alveolar) edge of the upper jaw is sharply
displaced downward like a crest. The upper premolars (except for the
reduced first one) and the molar are usually without an anterior acces-
sory cusp and mostly with a single posterior one. The corresponding
lower teeth (except the first two) have two accessory cusps—one each
anterior and posterior to the main cusp.

Sex-related differences in the skull are so insignificant that practi-
cally almost or no differences are seen in averages even in Statistical
variance analysis. Thus, while on the basis of the limited material of the
1930s (Chapskii, 1938) the condylobasal length of adult males from the
Kara and Barents seas was 1.5 mm more than that of the corresponding
values for females, this difference narrowed down to 0.7 mm with quan-
titatively more complete data (Potelov, 1968). Almost no difference was
seen in the mastoid width (at the mastoid process) while the width of the
rostrum in males exceeded the average corresponding value in females by
only 1 mm. As a percentage of the mastoid width, the difference in width
of the rostrum was slightly more noticeable: in males 36% and in females
about 34%. Differences in other indices (height of forehead, length of
forehead, and length of cranium) were insignificant. However, contrar-
ily, indices of length of the nasal bones and skull height of females were
slightly higher than those of males. There were no sex-related differences
whatsoever in terms of percentages of condylobasal length (mastoid and
rostral width, length of nasals, their width at base of the apex, etc.)
(Potelov, 1968). According to other data (Ognev, 1935), the craniolog-
ical differences between males and females were slightly more. But this
statement is based on limited data since the sexwise analysis could not
have been made at that time with the desired degree of accuracy.

The age-related changes of the skull conform to the general princi-
ples. In pups the skull is relatively inflated, the forehead stunted, and the
rostral portion highly narrowed; the upper profile sharply dips forward
and the interorbital space is relatively wider than in adults. With growth,
the skull becomes more elongated due to the marked elongation of the
forehead and the relatively less enlargement of the cerebral section. The
index of the rostral width as a percentage of the mastoid increases from
30% among six-month-olds to 36% in adults, while the index of skull
height decreases from 72.10 64%. The age-related changes of the median
bifurcated projection of the coronal suture are significant: this projection
narrows and increases in length with age while its saddle-shaped notch
becomes deeper and acute. Rapid wear and shedding of teeth is highly

173

typical. With advancing age, the skull becomes more massive, a feature
associated with the thickening of the bones and also the elongation of
the interfacial (longitudinal) suture relative to the interparietal length
(Ognev, 1935; Chapskii, 1937).

The geographic variation of the skull, although manifest in some
indices (Ognev, 1935; Naumov and Smirnov, 1936; Chapskii, 1963), calls
for review based on a more comparable and wider series (see under
“Geographic Variation,” p. 181). Variation is perceptible in the total
length, form of the nasal bones, width of the nares, and the interorbital
space.

The os penis is very faintly curved and considerably more thick-
ened than in the other 10-incisored seals. Its cross section resembles an
equilateral triangle with smoothened apices.

The diploid chromosome number is 2 n = 34.

The bearded seal is the largest member of the 10-incisored seals
(Phocinae). The body length of adults (seen as a whole) measured from
tip of snout to end of tail along the dorsal curvature (Lc) is 195-255 cm
and in a straight line (Lcv) 175-240 cm. The average length of animals
from the Soviet western arctic along the body curvature (Lc) is 233.6 cm
in males and 239.2 cm in females”*; the corresponding measurements in
a straight line (Lcv) are 222.0 and 222.5 cm (V.A. Potelov). The average
values for the Bering Sea bearded seals do not, however, reveal differ-
ences between males and females along the body curvature (Lc) (males
240 cm, females 239 cm) while in a straight line (Lcv) males are larger
(225 cm) than females (217 ст) (Kosygin, 1966).7* The axillary girth of
adult bearded seals of the Barents and Kara seas averages 152.5 cm in
males and 156.0 cm in females; these values for the Bering Sea animals
are 161.0 and 148.8 cm respectively.

The condylobasal length of the skull of adults is 200-240 mm,
average 220.5 тт, width at mastoids 130-140 mm, and rostral width
40-50 mm.

The length of the os penis of adults varies from 10 to 15 cm.

The weight of the largest animals may reach 360 kg but the usual
range is 225-320 kg. Males are roughly 25-35 kg heavier than females.
The Bering Sea males average 277 kg, females 242 kg (Kosygin, 1966);

22 These sexwise differences (as also some others mentioned above) are difficult to
explain biologically and evidently are the result of an inadequate number of measurements,
especially of the males.

23 The difference in average length in a straight line is evidently explained by the inade-
quate number of males measured by this method (four males versus 20 females).

24 Data of Chapskii (1938), Potelov (1968*), and С.М. Kosygin.

138

174

the Barents Sea males weigh оп average 256 kg, females 265.5 kg
(V.A. Potelov). (K.Ch.)

Taxonomy
Only species of the genus.
Geographic Distribution

The basin of the North Atlantic Ocean, the arctic waters of the Atlantic
from the Canadian archipelago to the Norwegian and Barents seas, and
the northern and northeastern boreal-arctic zone of the Pacific Ocean
(Bering Sea and the Sea of Okhotsk).

It is quite difficult to establish the northern limit of the distribution
in the Arctic Ocean. Some wandering animals in summer Stray very far
from the coasts into the Central Polar Basin and are found among ice
floes in very deep waters. Particularly Е. Nansen (north of 82° М lat.)
and Ch. Sverdrup (around 85° N lat.) and Soviet explorers during their
sojourn in the icebreaker “Georgii Sedov” in 1938 reported the recovery
of a bearded seal. They also reported these records in the region of some
drifting stations of “Severnyi Polyus”. Some animals entered directly into
the North Pole region where they were reported by R. Amundsen in 1925
and also by the Soviet polar explorers of “Severnyi Polyus-I” in 1937 and
“Severnyi Polyus-3” in 1954, etc.

Geographic Range in the USSR

Constitutes roughly about one-half of the total range of the species, i.e.,
its Eurasian sector, and much of the Pacific Ocean region (Fig. 100).

The distribution is mainly the result of three natural factors: cold
waters, presence of ice floes, and shallow depths. The bearded seal there-
fore descends to 60° N lat. or more southward only at places where these
conditions are generated by cold currents, 1.е., near the Labrador and
southern Greenland coasts, in the Sea of Okhotsk, and the Tatar Strait.
Being predominantly a benthic feeder and hence biologically associated
with shallow depths, this seal usually does not stray beyond the conti-
nental shelf and prefers depths of up to 100 m.

Sightings of this seal in pelagic regions at great depths is more often
the result of drifting ice floes. In some cases it is evidently caused by
migrations also or is merely a random straying.

In the Barents Sea in the northwest along the Murman coasts, the
distribution extends in a narrow belt to east of the boundary with Nor-
way and enlarges along eastern Murman into the White Sea Inlet. Near

139

176

the meridian 40° Е long., depending on the position of the ice rims,
the boundary of distribution steeply turns northward; there, it encom-
passes not only the entire shallow water zone of the southeastern Bar-
ents Sea from Kanin Peninsula to the coasts of the Vaigach Peninsula
and the southern part of Novaya Zemlya including Cheshsk Bay, Капт:
Kolguevsk shallow water zone and the Pechora Sea to Yugorsk Shar and:
Kara Strait, but also the pelagic regions. The latter include in particular
the very broad Novaya Zemlya belt of the Barents Sea tending northwest,
north, and northeast to the northern extremity of Novaya Zemlya. From
there the distribution opens up like a fan toward the Kara Sea and also
in the direction of the Franz Josef Land archipelago, including it. All
other—central, northern, and western—generally pelagic regions of the
Barents Sea are practically outside the limits of the range.

The bearded seal colonizes the White Sea almost everywhere but
with different densities in different seasons and in different regions.
Apparently it very rarely enters the extreme southern parts of the
Onezhsk and Dvinsk bays (V.A. Potelov).

It is extensively distributed in the Kara Sea where it is quite common.
In the western part of the sea it is found mainly in the shallow waters
between the Vaigach and Yamal peninsulas, in Baidaratsk Bay, in the
region of White Island, as also on the eastern banks of Novaya Zemlya
and even in the pelagic zone of the western part of the sea south of Cape
Zhelaniya, drifting there on ice floes. Farther east of White Island, the
range covers the waters surrounding Shokal’sk Island, the northern part
of the Gulf of Ob, and Yenisey Gulf.

In many regions where the habitational conditions for this species are
less favorable (deficiency of food, continuous ice cover, extremely shallow
depths, freshwater conditions, etc.), the boundary recedes slightly north
of the continental coastline. Thus the bearded seal inhabiting the Gulf
of Ob usually does not enter its southern part although its presence
has been reported in the Ob’ estuary (Brandt, 1856); it has not been
sighted in the deep waters of Tazovsk and probably Gydayamsk bays.
Evidently it does not go far south even in the Yenisey Gulf though it is
encountered in its northern regions. It inhabits everywhere in the more
open Pyasinsk Bay and even enters the Pyasina River (Heptner, 1936).
It is encountered on the highly rugged western Taimyr coast, abounding
in islands, the Vil’kitsk Strait, and the coastal regions of southwestern
Severnaya Zemlya.

How far north the range extends into the Kara Sea sector is not
wholly clear but it has been reported time and again on the western coasts
of Severnaya Zemlya. It has been encountered at the same latitudes and,

177

more westward, in the pelagic portions of the sea between Severnaya
Zemlya and the Franz Josef Land archipelago.»

Information on the distribution of the bearded seal in the Laptev Sea
is still scant. It is encountered regularly in Vil’kitsk Strait, especially in
the spring-summer period, its breeding grounds are known in the Cape
Chelyuskin region (Tyulin, 1938; Rutilevskii, 1939). These animals reach
the breeding grounds in large number from the Laptev Sea (Rutilevskii,
1939). They were noticed, albeit not frequently, on the coasts of eastern
Taimyr (L.N. Popov, 1939), close to Begichev Island (Koshkin, 1937), and
on Preobrazheniya Island (M.P. Vinogradov, 1949). They were reported
on the Novosibirsk Islands (reports of the Russian Polar Expedition,
1900 - 1903).

The habitation of the bearded seal even farther eastward in the -
coastal waters of the eastern parts of the Laptev Sea and in the
East Siberian Sea, reported in the last century, has been confirmed
by other references also. One of them, though not very reliable,
pertains to the Yana estuary (Bunge,* 1887) and another to the Kolyma
estuary (Iokhel’son, 1898). Such limited information led, in turn, to the
assumption of a fragmented range for this seal in the East Siberian Sea.
Some latest observations reported from the eastern part of this sea, 1.е.,
from the region of Chaunsk Bay, coastal areas east of Cape Shelagsk,
and areas directly adjoining Wrangel Island (Mineev, 1936; Fedoseev,
1966; and others), suggest that the Pacific bearded seal is found on the
eastern fringe of the East Siberian Sea. On the other hand, we cannot
ignore other reports indicating that in the more western parts of this
sea, including the Kolyma region, the Pacific bearded seal is not only
extremely rare, but is totally absent (У. Arsen’ev, 1935; Mikhel’, 1938*;
Rutilevskii, 1962). Evidently the Asian boundary between the ranges of
the subspecies E. b. barbatus and E. b. nauticus traverses somewhere in
this sea and not in the Laptev Sea, as stated by some (Scheffer, 1958;
King, 1964).

The western part of the Chukchi Sea, including the Wrangel and Herald
islands wholly fall in the range. The bulk of the population here is in the
coastal belt between the Chukchi coast and the pack ice masses in the
zone of drifting ice floes, as also in the immediate proximity of the coasts.
These seals enter the much higher latitudes of the sea depending on ice

25 Distribution of the bearded seal in the Kara Sea is based on the following sources:
Nosilov (1911*), Zhitkov (1913, 1924), Heptner (1930), S. Naumov (1931), Kolyushev
(1933, 1936*), Probatov (1933), Lepin (1936), Rundan (1936), Esipov (1937), Kirpichnikov
(1937), Mikhel’ (1937), Antipin (1936*), Rutilevskii (1939), Laktionov (1946), Tarasevich
(1963), and others, as also V.L. Vagin, G. Galkin, L.I. Leonov, V.N. Nikitin, V.A. Potelov,
А.М. Tyulin, К.К. Chapskii, I.K. Yakimovich, and others.

140

178

conditions and degree of ruggedness of the permafrost in the summer-
autumn period (Leonov, 1953; Tikhomirov, 1966; Fedoseev, 1966; P.G.
Nikulin, and others). It is not entirely clear whether the Chukchi Sea is
one of the regions of their permanent habitation, i.e., whether these seals
are found here in all seasons or mainly insummer. Although Pacific bearded
seals have been reported northwest of Wrangel Island even on October 10,
it is quite likely that most of them move into the Bering Sea in winter. There
are, however, references leading to an opposite conclusion (RazumovskKii,
1931*; V. Arsen’ev, 1935; Jonson et al., 1966).

The Bering Strait region falls wholly in the range of the Pacific
bearded seal. Its boundary diverges from here southward: in the west
into the Gulf of Anadyr, in the southeast and south (roughly along the
meridian 175° W long.) toward the Pribilov Islands near which it turns
east and northeast. The western branch of the Bering Sea portion of
the range extends in a broad belt corresponding to the boundary of the
maximum extent of the range to the south of drifting ice floes and to the
west up to the coastal zone in the region of Cape Navarin (Tikhomirov,
1964, 1966; Kosygin, 1966). From here the range descends in a narrow
border to the southwest along the Koryaksk coast to Olyutorsk Gulf
and later slightly enlarging along the coastline, reaches Karagin Gulf.
Farther south, on the eastern coast of Kamchatka, the range is actu-
ally interrupted although some authors have included all the Kamchatka
waters in it (Freiman, 1936; Kurcheva, 1954*).

The Commander Islands also cannot be regarded as the zone of
normal distribution of this species since the finds there are only rare
(Barabash-Nikiforov, 1936) or extremely rare (Marakov, 1968). Similarly,
this seal is rare on the Kuril Islands. It enters the region of the Comman-
der Islands evidently from Karagin Island, most probably transported by
broken ice floes which ultimately break up and thaw.

The Pacific bearded seal does not inhabit the whole of the Sea of
Okhotsk. The range in this sea could be represented schematically in
the form of an arc open toward the central portion of the Kuril range.
Thus the southeastern part of the sea apparently falls outside the nor-
mal range. The boundary of the latter at this place describes a half loop
extending from the southern Kuril Islands roughly along the meridian
155° Е lat. to north-northeast and curves toward the midsouthern -por-

tion of the western coast of Kamchatka at some places on the latitude of

Bol’sheretsk or even more northward. The rest of the areas in the Sea of
Okhotsk, north, northwest, and northeast from this line (fluctuating to
some extent depending on the position of drifting ice floes) to the coast-
line of the mainland and the coasts of Sakhalin, constitute the Okhotsk

141

179

part of the range of the Pacific bearded seal. It thus covers the She-
likhov Gulf, Penzhinsk, and Gizhiginsk bays up to their northernmost
sections, the entire mainland zone to the north and northwest of the
sea including the portion known as the Shantarsk Sea. Farther south the
western region of the Okhotsk portion of the range extends in a broad
belt from the Gulf of Sakhalin along the entire flank of Sakhalin, ter-
minating in the southern portion of the sea between La Perouse Strait,
Japanese waters (on Hokkaido Island), southern Kuril Islands, and the
southeastern boundary of the range.”°

The Pacific bearded seal is practically absent in the Sea of Japan per
se but for extremely rare episodic finds whose reliability is doubtful. In
Tatar Strait, however, the boundary traverses evidently slightly south of
the De Kastri latitude (S. Naumov and N. Smirnov, 1936).

A characteristic feature of the range of this species in the west (in
the Barents Sea) and in the east (in the Bering Sea and the Sea of
Okhotsk) and also to some extent in the expanse of the Siberian polar
seas is its fairly well manifest seasonal variation due to the influence of
ice cover dynamics and ice drift. The maximum boundaries of the range
described above mostly pertain to the winter-spring season, while the
real picture of the distribution of the animals in the other seasons is
quite different. At places, for example in the Barents Sea, Bering Sea,
and the Sea of Okhotsk, the range narrows toward the end of summer
and in autumn and extends in a narrow belt into the coastal zone. On
the other hand, in the Siberian arctic seas it is more extensive due to
ice movements and thinning of ice floes in the summer (see “Seasonal
Migration,” pp. 196 - 198).

Geographic Range outside the USSR (Fig. 101)

In the European waters along the coasts of northern Norway, it extends
from Varanger Fjord to North Cape and even down to Troms. In more
severe winters the animals enter farther south up to Vesteralen, but
invariably in small numbers. The bearded seal is common in Spitsber-
gen but encountered only sporadically along the northwestern coast of
Iceland. Farther westward, the range covers the Greenland waters along
the eastern coast from Cape Farewell to Datsk harbor of slightly more
northward along the western coast up to Robson Strait; the eastern parts
of the Canadian archipelago in Kane Basin, Baffin Bay, and Davis Strait;

26 The review of the Okhotsk portion of the range has been compiled from the data
of Ognev (1935), Freiman (1936), Nikulin (1937), S. Naumov (1941), Pikharev (1941,
1948), Kurcheva (1955), Tikhomirov (1961, 1966), Fedoseev (1970), Fedoseev, Gol’tsev,
and Kosygin (1970), and some other sources.

180

‘Crysdey `М-Я) 57124404 зпуши8ия ‘eas рэрлеэа эцз о UOTINGINSIC “TOT "8 ТТ

06

081 ost oz ag 0
oc a Lo
Ne FR [>
ue | CEA
$ >
‘
| й 8)
А уе WA
af >
09 < Ye
DS
х
a

E . = iv A
wees:
Eee, “A “a at Я

Se
06 09 ot 0 oc. 09

142

181

and the coastal zone of Labrador in the south to the eastern coast of
Newfoundland (extremely rare here). The range in the west covers the
entire periphery of Hudson Bay, Foxe Basin, Gulf of Boothia, Lancaster
Strait, and the waters of Cornwallis Island. Still farther westward, along
the southern mainland straits, the range reaches the Beaufort Sea and
the Alaskan coasts where the range of the Atlantic bearded seal joins (?)
with the extreme eastern branch of the range of the Pacific bearded seal.
The range covers the entire coastal waters of Alaska from its northern
coast to Bristol Bay including the region of St. Lawrence, St. Matthew,
Nunivak, and Pribilov islands. South of the Sea of Okhotsk, it extends
into the waters of Hokkaido Island. (K.Ch.)

Geographic Variation

In spite of recognizing two subspecies, Atlantic (E. b. barbatus Erxleben)
and Pacific (E. b. nauticus Pallas) quite long ago (Miller, 1923; Ognev,
1935; N. Smirnov, 1935; S. Naumov and N. Smirnov, 1936; Vinogradov,
1949; Ellerman and Morrison-Scott, 1951; Scheffer, 1958; Chapskii, 1963;
and others), no one has conclusively defined their morphological charac-
teristics. “The distinctive features of Е. b. nauticus are generally not quite

distinct as they overlap each other, and many of them can be recognized

only when comparing in a large series” (S. Naumov and N. Smirnov,
1936). This situation is relevant even today.

1. Atlantic bearded seal, Е. b. barbatus Erxleben, 1777 (syns. leporinus,
parsonsii, lepechini).

The body dimensions and weight are slightly less; the skull in most
of the animals has very narrow and long nasal bones, imparting a very
elegant appearance.

The body length measured in a straight line (Lcv) averages 220 cm
and the axillary girth 152.5 cm. The condylobasal length of the skull aver-
ages 220-221 mm and the zygomatic width 130 mm. The length of the
nasal bones averages 21.5 mm and width 6.5% of the condylobasal length.

Ecologically, this subspecies exhibits a distinct pagophile tendency.
The western part of its range covers the White, Barents, Kara, and Laptev
seas.

Outside the USSR, the range covers northern Norway, Spitsbergen,
Greenland, and the Atlantic portion of the Canadian archipelago.

2. Pacific bearded seal, Е. b. nauticus Pallas, 1811 (syns. albigena, lach-
tak).

Slightly larger than the Atlantic’ subspecies. The color is quite
monochromatic, often with a rust tinge in the cervical zone. The nasal

143

182

bones in most animals of this subspecies are broad in the anterior part
and narrow very sharply toward the apex. ;

The body length in a straight line (Lcv) averages 227.5 cm and the
axillary girth 161 cm. The condylobasal length of the skull averages
225.5 mm; length of nasal bones 17.5% (average) and width 8.1% of
the condylobasal length.

Ecologically, this subspecies tends notably toward coastal land where
it forms rookeries.

The eastern part of the range in the USSR covers the western regions
of the Chukchi Sea and evidently the easternmost regions of the East
Siberian Sea as also the Bering Sea and the Sea of Okhotsk.

Outside the USSR, the range covers the Bering Sea portion of
Alaska, northern Bristol Bay, and the Alaskan side of the Chukchi Sea up
to the western extremity of the Beaufort Sea (Chapskii, 1963). The differ-
ences between the Atlantic and Pacific forms were later confirmed (Kosy-
gin, 1969) but doubts arose once again (Kosygin and Potelov, 1971*).
(K.Ch.)

According to the most recent data (Chapskii, in litt.), these: sub-
species are entirely valid.

Biology

Population. No accurate census of the bearded seal is available to date
either within our waters or in the other regions of its range. In the 1950s,
the total world population was approximately put at 75,000 - 150,000
(Scheffer, 1958; King, 1964).

In the 1960s, an attempt was made to approximate the total pop-
ulation in the arctic and subarctic seas surrounding Eurasia. Various
indirect data, including the probable eastern Canadian population put
at 200,000 (McLaren, 1958), were used for this purpose. The resultant
highly approximate figure of 400,000 (Chapskii, 1966) reflects more the
probable maximum and surpasses the actual numbers, which no doubt
do not exceed 300,000 at the most.

The distribution of this population is uneven. Its presence in large
numbers is confined to the following major sections of the range: (1)
southeastern part of the Barents Sea, (2) southwestern part of the Kara
Sea, (3) eastern extremity of the Kara Sea, (4) Sea of Okhotsk, and
(5) northern and northeastern regions of the Bering Sea. Each of these
regions holds a substantial concentration of the animal in different sea-
sons. It is probable that the maximum possible potential for the popu-
lation growth prevails in the Barents-Kara region although the Sea of
Okhotsk has been holding first postion in terms of catch in recent years.

143

183

The Bering Sea reserves are less than those in either of the first two
regions individually.

In the western arctic USSR and in the White Sea the maximum
concentration of the bearded seal (sighting or catching over 100 animals
per day of hunting or ship survey) was noticed in the following regions:
(a) northeastern part of the White Sea Inlet and the southern part of
the Barents Sea adjoining the northern extremity of Kanin Peninsula; (b)
sections of the Barents Sea between the Kanin and Kolguev peninsulas,
western part of Cheshsk Bay; (c) eastern belt of the Barents Sea near
Novaya Zemlya (close to the southwestern coast of southern Novaya
Zemlya Island; the section of the sea opposite Matochkin Shar Strait;
and zones of Admiralty Peninsula and Cape Zhelaniya); (d) Franz Josef
Land archipelago; and finally (e) the easternmost cornei of the Pechora
Sea, near Yugorsk Shar.

The regions with a somewhat smaller population (50 to 100 animals
sighted per day) are: Kandalakshsk and Mezensk bays, other sections
(apart from those mentioned) of the Kanin-Kolguev shallow waters (at
Barmin Headland and north of Kolguev Island) and the Pechora Sea (at

Fig. 102. A female bearded seal emerging from water. Barents Sea, April, 1967
(photograph by V.A. Potelov).

184

144

Russk Bend and Кага Strait). In the western part of the Kara Sea the
regions that are as abundant are primarily Yugorsk Shar, Baidaratsk Bay,
north of White Island, and then the Kara Strait zone and on Matochkin
Shar. In the eastern part of the sea bearded seals in largest numbers are
noticed north of Pyasinsk Bay, in the region of the Minin reefs and more
toward the open sea (slightly east of the arctic archipelago islands), and
also almost in the eastern extremity of the sea, slightly west of Severnaya
Zemlya (west of Shokal’sk Strait). The bearded seal is seen in smaller
but yet significant numbers north of Dixon Island, in the region of Nor-
denshel’d archipelago, and also right in Shokal’sk Strait and on Pioneer
Island (V.A. Potelov).

The disposition of the bearded seal has, however, undergone some
definite seasonal dynamics (see p. 187).

In the Sea of Okhotsk the bulk of these seals is concentrated in the
spring in the northeastern extremities of the sea, in the lower sections of
Shelikhov Gulf, including Yamsk Bay, around Zabiyak, Babushkin, and ,
other straits in the northern strip of the sea in the west to Ushka Bay; it is .
also seen not far from the northwestern coasts of Kamchatka (Freiman,
1936; Pikharev, 1948; Tikhomirov, 1961). In the northern part of the sea
the vast majority of the Pacific bearded seals is confined in the spring-
summer period to the “inner” edge of drifting ice floes turned northward
(toward the coast) (Fedoseev, 1966). Among the regions with abundant
Pacific bearded seals until comparatively recently were the western part
of the sea near Sakhalin, mainly Terpeniya Bay and, in a slightly later

season, the Gulf of Sakhalin and the region of Shantarsk Islands with the

adjoining Academy, Nikolai, Ul’bansk, Tugursk bays, etc. (S.P. Naumov,
1941; and others).

In the Gulf of Sakhalin and farther west of Cape Litke in the direc-
tion of Shantarsk Islands, hunters daily counted a few hundred Pacific
bearded seals at the beginning of the season (Pikharev, 1948). This sit-
uation is no longer true.

In the mid-1960s, some attempts were made to count the Pacific
bearded seal on the spring-summer ice floes of the Sea of Okhotsk
(Tikhomirov, 1968). The areas of some nurseries and molting colonies
did not generally exceed 50-100 km? and the total population was deter-
mined in such colonies (500-1,000 animals maximally). However, the
precise number of such “spots” was difficult to count and hence it was
impossible to arrive at a grand total for the population. The Sea of
Okhotsk stands fourth in the relative population of the Pacific bearded
seal, after the ringed seal, ribbon seal, and evidently the larga.

It is quite likely that two local populations of the Pacific bearded
seal exist in the Sea of Okhotsk. One of them is in the northein part

145

144

185

of the sea tending mainly toward the expanses from Lisyansk Peninsula
to Tauisk Bay and farther to Babushkin, Kekurnyi bays; and Shelikhov
Gulf. Another population inhabits the southwestern part of the sea,
including the entire region close to the eastern coast of Sakhalin Island,
the Gulf of Sakhalin and the region of Shantarsk Islands (Fig. 103). In
the pupping season this “Sakhalin” Pacific bearded seal is concentrated
in Terpeniya Bay and in the air holes and open pools on the eastern
coast of the island. With the breaking up of the ice floes in spring, the
Pacific bearded seal migrates to the Gulf of Sakhalin and the Shantarsk
Islands.

The population density of the Pacific bearded seal in the breeding
colonies estimated by aerovisual surveys in 1967-1969 did not exceed on
average 0.2-0.8 per sq. km.

There are no geographic isolations between the populations of the
Okhotsk Pacific bearded seal and they freely intermix. It is therefore quite
possible that in summer, when these seals are widely dispersed along the
shelf zone of the Sea of Okhotsk, their populations could partly intermix.
The 55-56° М lat. could be regarded as the tentative boundary between
the northern and Sakhalin populations of the Okhotsk Pacific bearded
seal.

ns
— SS
= SAASS SSS
м
RS SARSSS SAAS
7]

i)
re
ra
12)
Q

Fig. 103. Main regions of concentration of the Pacific bearded seal, Erignathus
barbatus nauticus, in the Sea of Okhotsk at the time of whelping and the migratory
course of its population for molting (G.A. Fedoseev).

186

The number of Pacific bearded seals in the Sakhalin population by
aerovisual surveys was put at 35,000 - 40,000 and the population in north-
ern Okhotsk at 145,000 - 160,000 (Fedoseev, 1970; Fedoseev, Gol’tsev,
and Kosygin, 1970*; G.A. Fedoseev).

In spring and early summer the bulk of these seals concentrate in
the central region of the northernmost part of the Bering Sea, in a belt
close to the edges of drifting ice extending between the St. Lawrence
and St. Matthew islands. The population here is only slightly less than
that of the other two proximate colonies in the strip of drifting ice floes
continuing in a southeastern direction: one directly to the southeast of
St. Matthew Island and the other even farther toward the Pribilov Islands
and Bristol Bay roughly at the meridian of Nunivak Island. The dispo-
sition of these groups depends on the actual ice conditions in a given
year.

One more fairly distinct population is grouped on the ice floes in
the western part of the Gulf of Anadyr?’. In the relative population of
seals during the icy period in the pelagic portions of the Bering Sea,
the Pacific bearded seal once again holds last place at 10-12% of the
total population (Tikhomirov, 1964; Kosygin, 1966). The factual relative
proportion of this species is even less since the calculations did not take
into consideration the ringed seal, as it is generally extremely rare in the
pelagic regions of the sea.

The total population in the Bering Sea section of the range is less
than the corresponding numbers in the Sea of Okhotsk. The population
is even less in our section of the Chukchi Sea. It is not clear whether or
not the populations of the Bering and Chukchi seas are totally isolated
since there is information that the Pacific bearded seal wanders from one
sea to the other (see p. 198). The populations in the Laptev Sea and in
the westernmost part of the East Siberian Sea are most dispersed. East of
the Siberian Sea, the population rises quite rapidly and in the Chaunsk
Bay region and more eastward, the Pacific bearded seal becomes quite
common though less numerous.

Habitat. The habitat varies in different parts of the range and in
different seasons. Regions with a high rugged coastline with bays and
islands which offer protection from storms and from the piling up of ice
floes are the most favorable. The bearded seal avoids very shallow open
coasts since they do not provide adequate food (“plowed” by ice) and are
exposed to coastal waves and are generally less favorable for habitation

27 Distribution of the most important concentrations of the Pacific bearded seal in the
Bering Sea is based on the observations of Tikhomirov (1964, 1966b), Kosygin (1966a, b),
and K.K. Chapskii.

146

187

of such a large animal. Its typical habitats are not in the continental shelf
but the shallow waters (a few tens of meters deep) where it is confined
because it essentially survives on benthic food. This is the primary reason
for the predominant distribution of the population in such shallow waters
as found in the Kanin-Kolguev, Baidaratsk-Yamal’sk bays, and the Gulf
of Anadyr, northeastern regions of the Bering Sea shelf, and the shallow-
water periphery of the Sea of Okhotsk.

The increasingly freshwater sections of the straits and bays in the
mouth zones toward the western (Atlantic) part of the range hold little
attraction for this seal; its transgression even into the major rivers is a
rare phenomenon. These animals are found rather infrequently even in
the mouth and estuarine zones. The affinity of the Far-Eastern Pacific
bearded seal for rivers is altogether different. It readily takes to some
rivers in the Sea of Okhotsk, Amur lowland, and also not infrequently
in the Gulf of Anadyr. The factors responsible for these transgressions
are not fully understood. It is possible that sometimes, under condi-
tions of exposed coasts (as on the western coast of Kamchatka), this is
the result of seeking sites well protected from storms and possibly the .
massive availability of some common tiny fish. In the western part of
the Sea of Okhotsk the Pacific bearded seal is regularly found in the
summer-autumn period in shallow bays, such as UI’bansk, Nikolai, and
Konstantin. Here, during low tide shoals and banks are exposed over
much of the river mouth zone which these animals use readily. The rea-
son for their affinity for such zones is apparent from the example of the
formation of rookeries in the upper courses of ОГЪапзК Bay in the fore-
estuary areas which dry up in low tide. The animals initially enter the
river (Syran) and later crawl onto the dried-up laidas [low coastal plains
dissected by tortuous rills]. In the immediate proximity of the sea, the
animals do not crawl onto the laidas probably because of the unusually
high viscosity of the bottom and the extremely significant shallowness in
the adjoining sections of the sea. The bulk of the animals take advantage
of the fairway (S. Naumov, 1941).

Something similar is noticed along the coasts of western Kamchatka
too. “In summer the Pacific bearded seals are encountered in the shallow-
water bays with abundant reserves of bivalves and tiny crustaceans and
sometimes ascend with the high tide into the river... . At the end of
October, more Pacific bearded seals than larga or ringed seals begin to
appear in the summer... . At this time, during low tides at night the
Pacific bearded seals love to rest on the dried-up river banks or the
laidas in the estuary” (Lun’, 1936).

The rookeries are formed on low pebbly banks or sandy shoals and
also on banks consisting of pebbles, quite often with an admixture of

147

188

significant quantities of silt and some small boulders. Only rarely, stray
animals or small groups are seen on smooth rocks or rocky platforms.
In Konstantin Bay the substrate under the rookeries on the drying-up
banks is extremely shallow on one side and very deep on the other; the
animals are seen on the deep side.

The coastal rooKeries of the Okhotsk Pacific bearded seal are formed
during low tide and are occupied from the second half of summer and
autumn until the appearance of ice floes on the coasts. From this time,
the concentration of animals decreases and they begin to stray from the
coasts and are seen singly on-ice floes. Depending on the ice conditions
on the coasts, they go out deeper into the sea and remain among drifting
ice floes. In the strip of fixed shore ice, locking up the coast with a con-
tinuous cover, this seal is encountered only on the edges where there are
open waters and mobile broken ice floes. For respiration and emerging
onto ice whenever possible, it uses the natural openings in the ice floes,
such as open pools, air holes, and crevices. Liberally using its claws, it
resorts to making air holes and the like only in extreme cases, though it
is capable of making them as well as the ringed seal (N. Smirnov, 1927).
Most recent investigators regard the bearded seal as an inhabitant of only
drifting ice floes interspersed with open water pools. In this context even
its ability to make and regularly maintain air holes in the ice has gener-
ally been doubted although some rare exceptions are possible (McLaren,
1958; Mansfield, 1963; and others). In the spring-summer period, it is
confined to drifting ice floes and when some food is available, it can be
found even at considerable depths.

The bearded seal exhibits not much choice of floes in the spring-
summer period but nevertheless prefers low ice floes that have not
become hummocky (Fig. 104). These may be very extensive or very small
and hardly capable of supporting the animal. It rests mainly on pure
white ice, avoiding “soiled” ice floes and uses the latter only in extreme
cases when there is no alternative. Quite often, it rests on very thin and
weak ice floes and appears from a distance to be lying directly on the
water (Pikharev, 1941). In most cases it selects the edges of drifting ice
floes but at some distance from the very edge, free from surges. It is also
found far from the edge and clearly avoids large, highly compacted edges
of ice floes. It rests singly, often in pairs, and in threes at the very edge
of an ice floe.

Food. The bearded seal gets at food predominantly from the bottom
or close to it, mainly from depths of up to 50-60 m. The limits of its
submergence have not been established but evidently 100 m is not Ше.
limit.

189

147

Fig. 104. Pacific bearded seal on an ice floe. Bering Sea (photograph Бу С.М. Kosygin)._

The animal food consumed by this seal is diverse but the preferential
or more accessible species are readily distinguished. In this respect, much
depends on the local features of the deep-sea fauna as also to some extent
on the time of year. In general, over 70 species of invertebrates and fish
have been identified in the diet of this seal. However, in a given region
and season, a relatively small number of species serve as the basic food.
Most often, crustaceans, mollusks, worms, and echiurids are found in the
food; it also consumes large amounts of fish, especially the polar cod.
Crabs and shrimps predominate among the crustaceans; amphipods and
isopods are consumed far more rarely. Mollusks are represented by some
massive species of gastropods, mainly of the genus Buccinum, more rarely
bivalves. Cephalopoda are seen in its food relatively rarely. From among
the worms, the various members of Polychaeta—Pectinaria, Arenicola,
Harmathoe, etc. are consumed. Echiurids (mainly Echiurus echiurus) play
a noticeable role in its food while Priapulus caudatus has a subordinate
role. Sponges, sea cucumbers, even sea anemones, and others, broods of
mollusks, and sometimes fish spawn have been found time and again in
the stomach of this seal.

_ The food composition of the White Sea bearded seal is as follows:
fish about 30% (of the total number of stomachs examined), crustaceans
77%, gastropods 10%, bivalves 5%, worms 2.5%, and others 2.5%. The

148

190

composition of the fish is quite diverse with the polar cod holding
first place, followed by plaice (Pleuronectes platessa), common sand eel.
(Ammodytes hexapterus), herring (Clupea harengus), and more rarely
navaga (Eleginus navaga), cod, haddock (Gadidae), and sometimes even
sea trout (Salmo trutta) and nelma (Stenodus leucichthys nelma). The last
two species are consumed very rarely when the seal transgresses into a
river, аз for example the Shoina. Crustaceans are represented almost
exclusively by decapods, mainly crab (Hyas araneus), rarely shrimps
(Sclerocrangon boreas, Eualus gaimardi, Hetairus polaris), and sometimes
amphipods (Anonyx migax). Gastropoda are represented exclusively by
the genus Buccinum and bivalves by Cardium ciliatum. From among the
rest of the groups, Priapulus caudatus, Cucumaria frondosa, and ascidians
are found rather rarely (V.A. Potelov).

The specific proportions of the most important components in the
food of the Barents Sea seals are as follows: fish 65%, crustaceans
(decapods) 67.5%, amphipods 12.5%, gastropods about 20%, bivalves
8.5%, cephalopods 4%, worms 14%, sea cucumbers 4%, and others 7%.
The fish food is even more diverse than in the White Sea. Apart from
the polar cod, flounder, and herring (consumed at places in large quanti-
ties), as also navaga, sand eel, and cod with haddock (consumed rarely),
the bearded seal quite often feeds on goby (Cottus tricuspis) and some-
times also blennies (Lumpenus sp.), and skate (Raja sp.) (V.A. Potelov);
according to earlier data (Wolleback, 1907), capelin and sea bass are
consumed from time to time.

From among the crustaceans, this seal consumes more often crabs
(Hyas) and shrimps (Sabinea), more rarely Sclerocrangon ferox, S.
boreas, and also sea slater (Mesidothea entomon) (V.A. Potelov). It also
consumes Eualis gaimardi, Eupagurus pubescens, Spirontocaris palaris,
Hippolytidae, and sometimes many species of amphipods: Anonyx nugax,
Stegocephalopsis ampulla, Acanthonoroma sp., Gammarus locusta, and G.
homari (Chapskii, 1938).

From among gastropods, species of Buccinum hold first place, some-
times supplemented by species of the genera Natica, Septunea, and Acry-
bia flava (Chapskii, 1938). The bivalve most often found in the stom-
ach is Cardium graenlandicum; also seen are mollusks, such as Macoma
calcaria, Astarte sp., and species of the genus Mya, and Saxicava arc-
tica, which are not characteristic of the food of the bearded seal (N.
Smirnov, 1903, 1908). Sometimes it also consumes cephalopods, espe-
cially cuttlefish and squids (genera Rossia, Ommatostrephes) (Kondakov,
1932*; V.A. Potelov). It often consumes echiurids—Echiurus echiurus
and polychaetes—Arenicola and Harmathoe, and from among the other

149

191

groups of benthos, the sea cucumbers Cucumaria frondosa, Psolus, and
sometimes sea anemones, jelly fish, ascidians, and sponges.

Some local food characteristics have been noticed although they
reveal no systematic pattern (because of paucity of data). Thus, in the
Kanin-Kolguev shallow waters and in the Pechora Sea where the ben-
thos abounds in the crab Hyas araneus, it forms an extremely significant
proportion of the food of the bearded seal. Moreover, other crustaceans,
mainly tiny shrimps (Sclerocrangon boreas, Hippolyte sp., Sabinea septem-
carinata), and sometimes the sea slater (Mesidothea entomon) account for
a large proportion of the food.

Crustaceans also play a predominant role in the food of the bearded
seal on the western coasts of Novaya Zemlya (shrimps, but sometimes
even amphipods). In its autumn-winter food, however,-the polar cod
almost takes first place.

In the Franz Josef Land archipelago, in the summer season, the
bearded seal feeds mainly on gastropods while amphipods play a subor-
dinate role.

In the Kara Sea the food composition reveals no perceptible changes.
The percentage content of fish and amphipods is the same as in the
case of Barents Sea seals; the indices for gastropods are quite close
although the total proportion of mollusks is higher due to the greater
intake of bivalves; the role of crustaceans here is low. The fish food of
bearded seals in the Kara Sea is far less diverse. It consists almost exclu-
sively of polar cod with an extremely small proportion of sand eel, and
evidently sporadic arctic char and omul (cisco). The most important,
frequently consumed crustaceans are crab (Hyas araneus) and shrimp
(Sclerocrangon and some others); sea slater’. and amphipods are sec-
ondary. Mollusks are almost invariably represented by gastropods (Buc-
стит, more rarely Natica and Neptunea), and in rare cases by scallops
(Pecten). Cephalopods are regarded as of lesser significance while worms
play a slightly greater role: polychaetes (Chione) and echiurids (Priapulus
bicaudatus, Echiurus echiurus); sea cucumbers and ascidians from among
the other groups are found rather rarely (V.A. Potelov).

Sometimes the animals are satiated with extremely diverse foods:
only mollusks or exclusively shrimps (Sclerocrangon boreas and S. ferox)
and sometimes isopods (Mesidothea sibirica, M. sabini) or fish alone. The
diet is mostly mixed.

28 Often found in the stomach of the bearded seal caught in the region of White Island
(Tarasevich, 1963) although, according to V.A. Potelov, the sea slater represents more an

obligatory than a favorite food. |

192

In the western part of the Sea of Okhotsk the main food items in the
spring are crabs (Hyas coarctata), echiurids (Echiurus echiurus), shrimps
(Sclerocrangon boreas, S. salebrosa, Pandalus hipsinotus), cephalopods,
gastropods (mainly Buccinum) and bivalves, sea cucumber, and often
sponges too, though the nutritive value of the latter is extremely
doubtful. The full list of the food items of the Okhotsk bearded
seal runs into several tens of species. In the deepest water region
(northern Sakhalin) where the minimum depth is about 140 m, the food
is less diverse: mainly shrimps (Pandalidae, Crangonidae), gastropods
(Buccinum, Chrysodomus) and cephalopods (Octopus sp.), fish (smelt),
‘and some others. Deep-sea animals, such as crabs, lamellibranchs,
polychaetes, echiurids, sea cucumber, and others are also typical. At
other places, these represent the most important food items of the Pacific
bearded seal but here they play a minor role or are totally outside the
diet of the seal.

In the northwestern extremity of the Gulf of Sakhalin where the
depth is considerably less, the food of the Pacific bearded seal is more
diverse but consists mainly of benthic components, the most important
among which are the crabs (Hyas coarctata), echiurids (Echiurus echiu-
rus), shrimps (Crangonidae, Pandalidae), as also gastropods and bivalves
(Pikharev, 1941). The Shantarsk archipelago bearded seal exhibits a fairly
“similar diet” with the exception of echiurids and gastropods, which
are totally absent; the importance of bivalves, especially Mya, is sharply
reduced (Pikharev, 1941; S. Naumov, 1941). In Tatar Strait, in May, the
Pacific bearded seal consumes mainly shrimps (Sclerocrangon) and more
rarely amphipods (Gammaridae) (Yu.A. Salmin).

In the Bering Sea the Pacific bearded seal feeds in the spring-summer
period mainly on crustaceans, mostly snow crabs (Chionecetes opilio),
shrimps, mainly Crangon dalii and Nectocrangon lar lar, and also species
of the genus Pandalus; it relies to a lesser extent on other genera, such as
Sclerocrangon and Eualus; the remaining groups of crustaceans, includ-
ing gammarids and mysids form a small proportion of the food. Gas-
tropods, mainly of the genus Polynices and others which were not iden-
tified fully, represent another group of preferred foods. Cephalopods
(not identified to the generic level) can be placed in the third group
of importance. Among the remaining invertebrates, worms constitute a
noticeable proportion: polychaetes, as also Priapulus caudatus, Echiurus
echiurus, and others (Kosygin, 1966, 1971*). Until recently, sand eel and
daubed shanny (Leptoclinus maculatus) almost exclusively represented
the fish constituents of the food; flounder (Pleuronectes) was very rarely
found (Kosygin, 1966). It is now known (Kosygin, 1971*) that fish occu-
pies an important place in the food of the Bering Sea bearded seal. The

193

150 stomach of 565 animals revealed 13 species of predominantly deep-sea
fish: sand eel, daubed shanny, rattail, eelpout, leatherfin lumpsucker, two
different genera of flounder, delta smelt, capelin, saffron cod (navaga),
some type of bass, prickleback, and pollock. Although these fish finds
were noticed only in 9% of the animals inspected, the stomach of some
seals actually contained some of them in quantities exceeding a hundred
(capelin, daubed shanny, and sand eel).

No direct observations have been reported about the manner in
which the seal obtains its food. However, from the intense wearing of the
claws of the fore flippers in some animals, the rather frequent presence
of sand and pebbles in the stomach, and also the presence of pits in the
dried-up coastal belt, which, according to some researchers, were dug by
the bearded seal (S. Naumov, 1941), it is assumed that the animal digs
food out of the bottom (Pikharev, 1941). The seal has to dig the bottom
to reach buried bivalves and worms. It is highly possible that this is done
with the fore flippers as they have powerful claws.

Nevertheless, the bearded seal seldom has to dig for its food as
most of its dietary items are not buried in the bottom (crabs, shrimps,
amphipods, and other crustaceans and gastropods). Insofar as food ani-
mals such as Priapulus caudatus, which are not deeply submerged in the
soft bottom are concerned, the seal senses them with its whiskers and
can evidently easily get at them without using its flippers. The possibility
of seizing some objects submerged in the soil directly by the mouth is
suggested by the repeated finds of broken siphons of some bibalves in
the stomach of these animals.

Assessments of the seasonal variations in the food of this seal are
somewhat contradictory. From April to June, the stomachs of most of
the Okhotsk bearded seals examined were empty. The conclusion was
therefore drawn that this seal ceases to feed in spring (S. Naumov, 1941).
According to other data, however (Pikharev, 1948), it continues to feed in
spring but less intensively. The data obtained for the Bering Sea bearded
seal (Kosygin, 1966, 1971*) point to some reduced intake in the summer
season. A nearly similar picture emerges for the bearded seals of our
western seas (V.A. Potelov).

Home range. This could not be ascertained with certitude although
it can be suggested that different populations which undergo seasonal
dynamics are confined to their own home ranges, as in the case of many
other species. The solitary life of the animals of this species for much
of the year is well known, but this characteristic varies depending not
only on the season, but also the geographic region. In general, however,
the western populations lead a more “cloistered” life while the Pacific

151

194

Ocean animals (in the Sea of Okhotsk) tend to form herds and beach
rookeries in the latter half of summer and in autumn.

Hideouts and shelters. The construction of hideouts and shelters is
not a general characteristic of this species. In the more severe arctic
regions, for example in the Kara Sea, the bearded seals probably make
snow holes or shelters, using for this purpose primarily the hollows in the
snow hummocks that form near their air holes in an ice floe. Such snow
holes were found far from the coast of Siberia (N. Smirnov, 1927) and
around Cape Chelyuskin (Rutilevskii, 1939; A.N. Tyulin). The bearded
seal was regarded as an inhabitant of these shelters based solely on the
size Of the hole and the dimensions of the lair, as these exceeded those
of the ringed seal. These criteria are not reliable, however. Such shelters
were not detected in other sections of the range.

The ability of the bearded seal to make pits in the ice has been
known from at least the 1920s (N. Smirnov, 1927). The animals usually
survive without them however, preferring zones of drifting broken ice
floes. When the open water pools are covered with thin fresh ice, these
seals, like others associated with ice floes, break the crust and keep the
opening from refreezing by frequent use. Several such holes have been
found close together, for example in the Sea of Okhotsk (Fedoseevo,
1971).

In the northern European part of the USSR the bearded seal usually
does not overwinter in the coastal belt which is permanently ice-bound
(“fast ice”). But some young animals stray. Instances of such strays have
been seen on the southern coasts of Novaya Zemlya. The young remained
scattered in the winter in bays and straits covered with fast ice and were

‚ compelled, like the ringed seal, to make air holes. Cases of young bearded

seals emerging from such holes onto thick spring ice have been noted in
Rusanov Bay as well as in Petukhovsk Shar Strait (V.A. Potelov).

Daily activity and behavior. Prolonged observations of the same ani-
mals in nature are almost impossible. Nevertheless, it can be said with
certainty that the activity of the animal does not weaken, but increases
with the onset of twilight. Such nocturnal activity is confirmed by the
numerous catches of the animals in fixed nets at night, particularly in
autumn when they are trapped more often. In summer the animals prefer
to warm themselves in the sun, resting on the ice.

At the end of March and early April 1967, in the northern part of
the White Sea and in the southern part of the Barents Sea, bearded
seals were found in the water until 8:00 a.m.; thereafter, until 12:00
noon, the number of animals in the water and resting on the ice was
nearly equal. In the late afternoon hours, about one-fifth of the animals
were in the water; from 4:00 to.8:00 p.m. the number of animals in the

152

195

water rose again to one-third (observations beyond this hour could not
be continued because of darkness). Similar observations at August end
in the Kara Sea also revealed maximum animal activity in the morning
hours. Until midday, not a single animal was found on the ice while six
were noticed in the water from 4:00 to 9:00 a.m. and 32 from 8:00 to
12:00 noon. In the course of two four-hour surveys in the afternoon, five
animals were found in the water on each occasion and 10 and 54 on
the ice. From 8:00 p.m. to midnight and in the first four hours of the
following day, 26 animals were noticed in the water on each occasion
and 24 and 12 animals on the ice.

In Baidaratsk Bay, from July 5 through 12, 1961, some 140 bearded
seals were sighted from a hunting ship. The number of animals resting
on the ice at different times of the day was (as % of the total number
of animals counted in the corresponding intervals, averages): 8:00 a.m.
to 12:00 noon—6%, 12:00 to 4:00 p.m.—31%, 4:00 to 8:00 p.m.—33%,
8:00 to 12:00 midnight—26%, and 12:00 to 4:00 a.m.—3%.

Thus in the morning hours almost all the animals remain in the
water looking for food while their activity decreases throughout the day.
Emergence onto the ice occurs exclusively in the second half of the day.
From May end through July inclusive, they rest on ice for a considerably
longer time than in other months (V.A. Potelov). A similar pattern was
observed in the Sea of Okhotsk also. The Pacific bearded seals there are
seen on the ice and in the water in spring and early summer at different
times of the day. They crawl onto ice in the largest numbers after 10:00
a.m., especially in good weather on clear sunny days. At first they are
quite restless but then fall asleep, sometimes very soundly. An animal
resting on the ice can often be approached within 3 to 5 m without
awaking it. The Pacific bearded seals sleep most soundly in good sunny
weather (S. Naumov, 1941).

This seal is most often found singly on the ice, rarely in pairs or trios
on a single floe, and extremely rarely in a group of five to seven animals;
when in such a group, the animals remain apart, invariably along the
very edge. The seal sleeps mainly on its side or on its back. In spite of
its heaviness, the animal clambers quite easily onto ice. First, it thrusts
its head high above the water for a survey, surfaces right up to the ice
and, holding onto it with the fore flippers, heaves itself upward with a
massive thrust of the hind flippers (Pikharev, 1941a*) or throws itself
up in one leap. While diving, it usually reveals a part of its back and
sometimes (evidently when at peace) the entire back and even the hind
flippers.

In the period of formation of the beach rookeries in the Far East,
activity is related not so much to changes in light and darkness as to ebb

196

and tide pattern, since the rookeries are formed on the receding water
front and begin to break up with high tide. Our western hunters regard
the bearded seal as an extremely timid animal while eastern hunters sub-
scribe to the opposite view. Both are evidently right since the prolonged
and intense killing of the animal in the Soviet western arctic seas has
led to greater fright and caution than in the Far East. It is possible that
even the name “sea hare,” given to it by Russian hunters in the west,
reflects to some extent the timid behavior of the animal.2? In the water,
however, it is bolder and permits the approach of man far closer than
when on ice.

Seasonal migrations and transgressions. Ordinarily, these seals are
regarded as settlers not given to en masse or prolonged migrations
(N. Smirnov, 1908, 1929; Ognev, 1935; Freiman, 1936; S. Naumov, 1941;
partly Tikhomirov, 1961; Shustov, 1964*; and others). In fact, the bearded
seal does not perform migrations of the type characteristic of the Green-
land [harp] seal. Nonetheless, the bearded seal cannot be regarded as a
completely settled animal since its population almost everywhere under-
goes fairly perceptible seasonal changes, largely due to changes in ice
conditions and also probably depending on the distribution of food and
selection of a site for reproduction, molt, and for setting up beach rook-
eries.

In the simplest case, the seasonal changes occur as follows: from
the second half of summer and in autumn, the bulk of the animals is
distributed in the coastal belt or in the shallow waters in the relatively
warm sections of the range where ice floes have thawed. With the onset
of winter and formation of the coastal ice cover, the animals usually leave
the shores. Throughout the winter-spring period, they remain beyond the
stationary coastal ice floes (fast ice) among drifting floes. In the eastern
part of the Barents Sea many animals are found far away from the coasts,
almost on the outer edge of the ice floes. As soon as the coast is free
from fast ice, part of the population is distributed along it and settles
preferentially in regions with a more rugged coastline, while the other
part moves away with the ice floes. From the second half of summer,
a reverse movement occurs: that part of the population which left the
coast with the ice floes now returns.

Such a simple pattern of seasonal migrations is characteristic in many
sections of the Sea of Okhotsk. At the beginning of winter, with the
appearance of stable ice floes and depending on their spread, the Pacific
bearded seal withdraws from the coasts to the outer edge of the coastal

29 Others suggest that it was so named because it “hops” while negotiating on ice (or on
land).

153

197

ice belt into the open water pools among mobile broken ice floes, moving
with them in the prevailing wind and current directions. In the regions
where the ice floes do not block the coast and where there are open
sections of water, as for example at places in the Shantarsk archipelago
straits and in the lower sections of bays (UI’bansk, Nikolai, and Kon-
stantin), the Pacific bearded seal is long confined to the coasts and small
numbers can always be seen almost throughout the winter (S. Naumov,
1941). The majority, however, go far away from the coasts and spend
the winter and also early spring far away from the Shantarsk Islands. In
the northern part of the sea the Pacific bearded seal moves toward the
massive drifting ice floes in winter and spring and onto their edges facing
the continent (Fedoseev, 1966).

In the second half of spring and early summer the animals are dis-
persed more extensively with the drifting ice floes but within their food
zones, confined to the shallow sections of the sea.

A reverse process occurs after the ice thaws, when the seals approach
the coasts, quite extensively scattered at first, and then concentrate in
the beach rookeries in the same place year after year from August end.
but mainly in September and October. They remain scattered deep into
autumn, at which time the coasts begin to freeze, after which a new cycle
of travel of the Pacific bearded seal commences on the ice floes in the
sea.

There is a view that one group of the Okhotsk population performs
distant migrations in summer and autumn from the southern regions
of the sea (especially from Terpeniya Bay) along the eastern flanks of
Sakhalin in the north into Shantarsk archipelago. It is these animals, per-
forming distant migrations, that form the coastal rookeries (Tikhomirov,
1961, 1966b). At the same time, there is a reference to the fact that “the
Pacific bearded seal moves northward into the Sea of Okhotsk follow-
ing the thawing ice floes in Tatar Strait” (5. Naumov, 1941). The real
situation can only be gauged by studying marked animals.

The seasonal migrations of the Pacific bearded seal in the Bering
Sea are evidently somewhat more complex. Its population winters in all
probability partly on the shores and partly on the ice edge formed in
the extreme northern part of the sea. As the ice recedes southward and
the shores are blocked, the fairly well-scattered population also moves
with the ice floes southward and ultimately turns up on the edges of the
pelagic expanses by early spring (see “Geographic Distribution,” p. 174).
It is very difficult to establish whether it is the Alaskan population alone
or the Chukchi-Anadyr population also that is concentrated in these
peripheral sections of ice floes drifting southwest of St. Lawrence Island
and southeast of St. Matthew Island. In all probability, both populations

154

198

are held there in some proportion but this can only be established with
certainty by studying marked animals.

As the summer approaches, the boundary of drifting ice floes with
seals begins to shift gradually in a reverse direction northward, to the
Bering Strait, with the bulk of the Pacific bearded seals carried along. It is
quite possible that a significant part of this population moves northward
even farther, into the Chukchi Sea, where it spends the summer in the
wide expanses and, possibly, early autumn. The seals, for the most part,
are evidently scattered in the coastal belt east and west of the Bering
Strait and their return commences in mid-autumn.

A new viewpoint has emerged in recent years on the seasonal migra-
tions of the bearded seal in the White, Barents, and Kara seas. Reports
of the wintering of this seal in the White Sea are reliable but the win-
ter range becomes extremely narrow and is restricted evidently only to
some small regions. In the southwestern part of the sea such regions are
the extreme north of Onezhsk Bay, southwestern sections of the Dvina,
and eastern sections of the Kandalaksha bays. Small numbers of these
animals, however, winter in all these places. Many more bearded seals
winter in Mezensk Bay and in Voronka (V.A. Potelov).

In the Barents Sea this seal winters in the extensive expanses of
drifting ice floes in the southeastern part of the sea. It has been assumed
that the migratory courses in the spring-summer months diverge in
different directions. One leads northeast, along the Novozemel’sk coast
to Cape Zhelaniya and probably terminates in the Franz Josef Land
archipelago. The other course runs mainly through the Kara Inlet into
the western part of the Kara Sea and perhaps extends not only to
Baidaratsk Bay and up to White Island, but even farther toward western
Taimyr (V.A. Potelov). A reverse movement to the wintering grounds in
the Barents Sea occurs in the autumn.

Reproduction. Even 20 to 30 years ago, it was thought that these seals
matured in three years and some females even in two years (Sleptsov,
1943). At the end of the 1950s, based on age established from the claws,
it became clear that the eastern Canadian female bearded seals attained
sexual maturity and conceived for the first time at the age of six years
while the bulls became productive at seven years (McLaren, 1958).

The female Pacific bearded seal is the earliest to attain maturity, at
the age of three years, but such quick-maturing animals are rare (8% of
generation). Mature animals among four-year-olds constitute about one-
fifth of the generation but not all the five-year-old females are mature
(only 83%); they are all sexually mature only from the 6th year onward.
The bulls are all immature at four years of age; 50% attain maturity at

154

199

Fig. 105. Embryo of a bearded seal. Kara Sea, August, 1965 (photograph by
V.A. Potelov).

five years and 66% at six years; all the males are mature only from the
seventh year (Tikhomirov, 1966а*).30

Time differences in maturity were detected even in the case of the
bearded seal of the western seas of the USSR; 15% of the females of
a given generation mature and become fertile at four years of age, over
one-half at five and six years, and some individual females at seven years.
Males, however, attain sexual maturity earliest, at the age of five years.
At six years, over one-half of the males are mature while at seven years,
immature animals are quite rare while all the eight-year-old bulls are
mature (V.A. Potelov).

Information on the mating season long remained contradictory.
From the 1930s, with the availability of information on the behavioral
characteristics and based on an analysis of the reproductive organs
(Chapskii, 1938; McLaren, 1958), spring, preceding the period of molt
and not summer or autumn, was judged as the mating season of the
Atlantic subspecies (Wolleback, 1907; Laktionov, 1946). The mating

30 All Pacific bearded seals of both sexes up to five years of age inclusive are regarded
as immature in field calculations of growth and weight increments (Kosygin, 1966c).

155

200

behavior involves а state of excitation in the animals, which pursue each
other, and perhaps “mating calls,” recognized as a manifestation of sexual
reflexes (Dubrovskii, 1937).

The bearded seal in the western part of the Soviet range shows
intense production of sperm from the second half of March to early
July, while ovulation occurs from March end through June but in most
animals in the last few days of April. Sperm were detected in the gen-
ital tracts of females toward the end of March, in April, and in the
first half of May. All this suggests quite an extended mating period
(V.A. Potelov).

Errors occurred in determining the mating season of the Pacific sub-
species: it was assumed that this seal mated in July (Sleptsov, 1943;
Kurcheva, 1955). Observations as well as a study of the reproductive
organs showed that the Bering bearded seal also mated in the spring,
from April 20 through May 15, mainly in the first 10 days of May
(Tikhomirov, 1964). The weight of the testes increases from early March
(115 g) to around mid-April (averaging almost up to 190 g) but decreases
(to 140-150 g) by the second half of May; this weight reduction continu-
es up to early July (Tikhomirov, 1966d*). The different dates of mating
and whelping gave rise to different conclusions regarding the duration
of the embryonic period. For the Far Eastern subspecies, it was formerly
assumed as nine months (Sleptsov, 1943) and for the Atlantic subspecies
as roughly 11 months including a lag of embryogeny in the first 1.5 to
2.5 months (Chapskii, 1938; McLaren, 1958).

According to the earlier data, sterility in the Atlantic and Pacific
subspecies covered up to 50% of the total eligible females. It was assumed
that sterility followed the year of whelping (Chapskii, 1938; Sleptsov,
1948; McLaren, 1958). According to the present data, not more than
25% (V.A. Potelov) or even 20% (Johnson et al., 1966) of the females
are sterile every year.*! In general, however, the mating period of the
Pacific bearded seal has not been adequately dealt with in the literature.

It is significant that mature males are encountered very rarely from
April through June in the whelping area of the Bering bearded seal on
the sparse ice floes south of the St. Matthew and Nunivak islands. Thus,
in 1963, of the 28 animals caught, only one was an adult; in 1964, among
the 43 bearded seals caught there, only one was a productive male. More
to the north, however, where hummocked ice floes commence, males
and females of all ages were caught (Kosygin, 1966b). It is possible,
however, that males remained on the St. Matthew and Nunivak islands
but lay in the water (animals swimming in the broad water pools between

31 An even smaller figure of 10% has been estimated (Tarasevich, 1963).

156

201

the ice floes were sighted quite often in 1964) and hence could not
attract the attention of the hunters. In any case, the mating period is
highly imperceptible. Probably, it proceeds without violent encounters
between the competitors, which is quite understandable considering the
remarkably peaceful disposition of this animal in general and the absence
of any seizure marks whatsoever on the skin of the males.

Some regions of pupping have been identified in the western seas of
the Soviet arctic.

Whelping occurs evidently only in the southeastern part of the Bar-
ents Sea and in the northern regions of the White Sea. The main region
of reproduction in the Barents Sea is the zone of drifting broken ice
floes with abundant open water pools between them in the expanse from
the White Sea to the Vaigach and Novaya Zemlya islands.*? Beyond
the limits of these ranges, whelping is extremely rare. Thus, some stray
newborn pups were detected in the central basin of the White Sea
(K.K. Chapskii), in the Kara Sea sections closest to the Barents Sea,
1.е., in Baidaratsk Bay in the zone of Yugorsk Shar and the Kara Strait
(V.A. Potelov). In recent years newborn pups have been found in the
White Sea only in Mezensk Bay and Voronka. In Dvina Bay and Gorle,
they were nowhere encountered though these regions were surveyed in
March of every year from airplanes and from hunting vessels. No data are
available on the finds of conceived or lactating females in Kandalakshsk
and Onezhsk bays (V.A. Potelov).

The whelping period in the western parts of the USSR range is
confirmed by numerous instances of finding almost completely developed
fetuses ready for birth and newborn pups still on the mother’s milk. In the
White Sea, gestating females caught in the first half of March included
fully formed pups which, on being delivered by dissection from killed
mothers, could crawl on ice (У. Smirnov,* 1927). One newborn pup was
found at Gorle at the end of March, 1947. In the 1960s (V.A. Potelov),
new and recently born pups were encountered in the White Sea in the
last 10 days of March, in the middle and last 10 days of April, and in the
first 10 days of May.

Premature fetuses were found in the Barents Sea on the Novaya
Zemlya coasts at March end and in early April, and in the Pechora Sea
from the second half of April. Newborns with mothers were noticed
in the easternmost part of the Barents Sea from the Pechora Sea to
Admiralty Peninsula from the end of the first 10 days of May to the

32 Along the Novaya Zemlya fringe of the Barents Sea, newborns with their mothers
were not noticed farther north of the Admiralty Peninsula. Pupping was not reported at all
in the Franz Josef Land archipelago.

156

202

Fig. 106. Fetus of the Pacific bearded seal, Erignathus barbatus nauticus, during
the process of birth. Bering Sea (photograph by G.M. Kosygin).

end of the first 10 days of June. Observations in the 1960s covered
the whole of April and the first 20 days of May, ie., the whelping
period. Considering that the apparently developed pup was still feeding
on milk (i.e., including the entire lactation period), the “pupping period”
could be regarded as extending there into the first 10 days of June
(V.A. Potelov).

Thus, pups in the western arctic seas of the USSR are born from the
last 10 days of March through mid-May, mainly through April, but some-
what earlier in the White Sea than in the Barents Sea where lactation
(and hence also whelping) is delayed by at least 10 days.

Information on whelping in the eastern arctic regions of the USSR
is extremely vague. The view was expressed that the Kara Sea bearded
seals migrated for winter to the Laptev Sea (Rutilevskii, 1939). From
this, the conclusion was drawn that the pups were perhaps born east
of Taimyr. There is no information on whether the Pacific bearded seal
reproduces in the East Siberian and Chukchi seas.

157

157

203

In the Sea of Okhotsk pupping probably occurs everywhere but
mainly at places where these seals concentrate (see p. 178) in the spring,
1.е., 1) in the western fore-Sakhalin area (including Terpeniya Bay) and
2) in the northern mainland area (mainly the eastern regions, depend-
ing of course on the position of drifting ice floes) (Fedoseev, 1971).
In general, however, information on the reproduction of the Okhotsk
bearded seal, including the conditions and period of whelping, which is
regarded as March end to April, needs to be more accurately established
(S. Naumov, 1941; Tikhomirov, 1961; Shustov, 1964). This period (from
February through April) for the region of Amakhtonsk Bay is even more
indefinite (N.A. Smirnov, 1911). A similar period has been indicated for
Tatar Bay also (S. Naumov, 1941). Most of the newborn pups in the
Bering Sea appear along the edges of drifting ice floes in the northern
part of the sea (region between St. Lawrence and St. Matthew islands
and the expanse between the Nunivak and Pribilov islands) and to a
lesser extent in the Gulf of Anadyr. The whelping period here is from
March end through early May. The peak of whelping here was recorded
in the first 10 days of May in 1963 (Kosygin, 1966) (Fig. 107).

In the whelping period the animals are not concentrated in a group
as in the rookeries. Instead the whelped females scatter far apart on the

Fig. 107. Newborn Pacific bearded seal. Bering Sea (photograph by G.M. Kosygin).

158

204

extensive expanses of the drifting ice floes. Thus in 1962, near Novaya
Zemlya, on the traverse of the Gulf of Sakhalin and later on the edge of.
ice floes 50 km away from Matochkin Shar, some 30 females with pups
were sighted, lying at 100-300 m away from each other (V.A. Potelov).

The female bearded seal is not particularly choosy about the icy
substratum for the newborn. She can make do with an even extremely
small portion of a floe or the very edge of a large ice field. The view
of some authors (Tyulin, 1938; Rutilevskii, 1939) that whelping occurs
possibly under the ledge of an ice floe or under cover of snow in special
lairs or holes made by the adult animals in the hummocky piles along the
shore ice or far away from it has not been confirmed. Pups everywhere
are born in the open. The statement that these seals reproduce on fast ice
in the bay and generally on shore ice is equally doubtful (N.A. Smirnov,
1927). The very act of birth has not been observed to date. Evidently it
is quite rapid. It was even suggested that the birth can take place not
only on an ice floe, but also in the water (Vibe, 1950).

Growth, development, and molt. Until recently, the size of the new-
born pup was usually established indirectly by comparing the large fetuses
ready for birth. Thus the length of a pup of the Atlantic form at the
time of its birth was put at 120-125 cm (in a straight line, Lcv) or
130-135 cm (along the dorsal curvature, Lc). According to the latest
data (V.A. Potelov), newborn pups from the White and Barents seas
measure 111-126 cm, average 121 cm (Lcv), and 121-141 cm, average
135 cm (Lc). The length of the Bering bearded seal at the time of birth
was put at 118-137 cm, average 127 cm (Lcv); the fetus before birth
weighed on average 30.2 kg (G.M. Kosygin).

Fetuses measuring about 100 cm long have a dense pelage consisting
of uniformly soft, slightly curled and flattened hairs roughly 15 to 25 mm
long. The color is mainly silvery-gray at the base with fairly vivid brown
tones in the upper portion. This primary embryonic coat undergoes some
changes as the time of birth approaches due to the growth of guard and
intermediary hairs and also thinning of the underfur (V.A. Potelov).

The embryonic hair coat is shed partly or even completely in the
mother’s womb immediately before birth. The newborn has the embryonic
hair coat, therefore, only on some parts of the body or this coat has been
totally replaced by a new coat. Clumps of matted embryonic fur are found
in the womb, in the fetal fluids (as small thickened disks), and also in the
intestine of the fetus itself (Bychkov, 1960; V.A. Potelov). The data for the
Bering pups do not conform to these observations; these pups are born
with a firm hair coat that changes only 2-4 weeks later (G.M. Kosygin).

The molted pup of the Barents Sea resembles somewhat a large,
full grown pup of the hooded seal in the color of its skin, but has a

159

205

somewhat more vivid silvery luster. The dorsal side is dark gray and the
ventral side light steel-gray (V.A. Potelov). Numerous tiny specks and
dabs of black or brown are sometimes seen scattered all over the body
or just on the front portion and flanks; these extend even onto the head.
The characteristic color of the head (light-colored spots) has already
been described.

The milk teeth perceptible during development in the womb
(Steenstrup, 1860) are preserved in the newborn although in an

intensely resorbed form. The permanent upper incisors and canines,

as also the second and third premolars, have already cut before birth
(G.M. Kosygin).

The lactation period extends for about a month in the western part
of our range (V.A. Potelov). In the Okhotsk and Bering seas too this
period is nearly similar, terminating in most females in the first 10 days
of May according to Tikhomirov (1966a), and in the Bering Sea in the
last 10 days of May according to G.M. Kosygin.

There are no detailed reports on the frequency of feeding or on the
amount of milk suckled by the pups in one feed. The maximum amount
of milk found in the stomach of a pup was 1 liter. The fat level of the
milk of the Pacific bearded seal varies from 30 to 60% (G.M. Kosy-
gin) and that of the western bearded seal 26 to 61% (V.A. Potelov and
A.G. Beloborodov).

It is difficult to trace the growth intensity during this period. From a
comparison of the average sizes of newborn and fully grown pups by the
end of May or early June (Lcv 140 cm), it is quite evident that during
lactation a pup of the Barents bearded seal roughly adds 10-15 cm (the
Lc length increases, on average, up to 150 cm).

In the following summer months and early autumn, the pup adds
roughly another 5 cm; a yearling measures roughly 155 - 160 cm (Lcv) or
170-175 cm (Lc).

In terms of percentages, the average adult growth dynamics can be
expressed as follows: length of newborns about 55% that of fully grown
adults; at the end of the lactation period, about 64%; five- to six-month-
olds in autumn, 72%; and yearlings, 76%. Six-year-old females in the
Canadian population (fully mature by this age) record 91% of their full
growth (McLaren, 1958).

The maternal instinct among the seals of this species is quite well
manifested. The female spends considerable time on the ice floe near
her pup, going into the water to feed only for a short duration. In this
period the female cannot make good the loss of her subcutaneous fat
reserves; she is highly emaciated and the thickness of the adipose layer
has decreased to 3 cm or less by the end of the lactation period. When

206

people come around, the female sometimes attempts to push the pup
into the water and dive with it (Tikhomirov, 1964).

The molting periods are not yet fully understood. In the Kara Sea
this period covers July and August (Chapskii, 1938; V.L. Vagin) but its
commencement there has not been precisely traced. Much information
was reported in the 1960s on the molting of bearded seals in the White,
Barents, and Kara seas in an unusually early period. Thus, in the White
Sea molted animals began being encountered from the second half of
April and early May; in southeastern Novaya Zemlya in the second half
of May; in the northern regions of the Barents Sea in June and early
July; and in Baidaratsk Bay from early July (and evidently much earlier
since fully molted animals were seen at this time) (Potelov, 1967*).

In the southwestern part of the Sea of Okhotsk molting continu-
es from early May through July end (Pikharev, 1940*; Nikulin, 1937;
$. Naumov, 1941; Sleptsov, 1943; Kurcheva, 1958*). This period is greatly
extended in different regions of this sea*>

In the northern parts of the Sea of Okhotsk molting proceeds gener-
ally at a very late period compared to that in the southern parts where it |
commences even mid-April (Freiman, 1936). According to recent data,
the chronology of molting in the Sea of Okhotsk is as follows: stray
molting animals, especially in Terpeniya Bay, were noticed already on
April 16; the number of such animals had increased in May and reached
maximum in June. Their number was even higher in July but the major-
ity had already molted; nevertheless, some stray unmolted animals were
also seen among them (Tikhomirov, 1961).

In the Bering Sea, too, the molting period is extended. It commences
with immature animals even in March and April (?; K.Ch.); adults molt
mainly from early or mid-May to mid- or end of June (Tikhomirov, 1964;
Kosygin, 1966a*).

Enemies, diseases, parasites, mortality, and competitors. Among the
large animals that can be reckoned as enemies of the bearded seal in
the arctic part of the range, the polar bear has long been regarded as
number one. Its stomach often reveals bits of the skin with blubber and
sometimes even the whiskers of this seal. The skin of many seals caught
in the Kara Sea showed distinct scratches made by the bear’s claws. With

33 Bychkov (1960) reported a much earlier period of molt: among the animals of different
ages caught in Terpeniya Bay from March 28 through 31, nine were in a high state of molt
(two adult males, two gestating females, and the rest immature animals of both sexes). In
six animals intense shedding of hair throughout the trunk had occurred; in two others there
were large sections of bare skin on the abdomen and intense shedding of hair on the back;
the ninth animal had small clumps of wool only on the neck and around the flippers, while
the rest of the body was bare. This pattern of molting is evidently pathological.

160

207

the polar bear population reduced at present, the damage caused by it
is no longer significant. At places on the coast of the Sea of Okhotsk,
during the period of formation of beach rookeries, the Pacific bearded
seal suffers also from the brown bear. Seals torn apart by this bear have
been seen time and again (on the Moroshechnaya River) and the bare
skin of a Pacific bearded seal left behind by a bear was found in the
region of Cape Amakhtonsk (Tikhomirov, 1966a).

Evidently, young animals are sometimes attacked here and there
by the Greenland shark although there are no concrete data to sup-
port these events. The damage inflicted by the killer whale is obviously
not significant although the stomach of these carnivores caught in the
Bering Strait contained the whiskers and claws of the Pacific bearded
seal (Zenkovich, 1938*). In exceptionally rare cases, the walrus, too,
attacks the bearded seal. At present, however, with the walrus popula-
tion decreased everywhere (particularly sharply in the western parts of
the range in the USSR), the damage caused by the walrus, if any, is practi-
cally negligible. Also, there is essentially no trophic competition between
these animals. The mollusks consumed by them are varied; while the
bearded seal prefers gastropods, the walrus thrives almost exclusively on
lamellibranchs. These differences have been strikingly demonstrated in
the case of the Greenland-Canadian animals too (Vibe, 1950). At places
on the Chukchi coasts, the gray whale, feeding mainly on benthos, is
perhaps a competitor of the Pacific bearded seal.

From among the ectoparasites of the bearded seal, especially in the
Far East, Echinophthirius horridus Olfers (Anoplura) is widely prevalent.

Heavy infection with helminths is reflected far more significantly in
the health of this species of animals. Among several dozens of animals
inspected even in the 1930s in the Barents and Kara seas (Chapskii,
1938), there was not a single case in which the stomach or intestines
did not contain a large number of cestodes and nematodes. Instances
are known of the highly diseased state of the animals, with their ali-
mentary canals choked with helminths. As an example, in one animal
the intestines, packed with cestodes, were ulcerated at several places to
the point of perforation. The animal was extremely sluggish and highly
emaciated and there was almost no adipose layer (L. Leonov; August,
1933, Franz Josef Land).

The helminth fauna of the bearded seal comprises 23 species and
three larval forms (Delyamure, 1955; V.V. Treshchev, M.V. Yurakhno).

From among the trematodes, Opisthorchis tenuicollis parasitizes the
bile ducts of the liver. Orthosplanchnus arcticus, infecting the gall blad-
der, bile passages of the liver, and the pancreas, is encountered more
often than other trematodes and in much larger numbers. Thus the

161

208

liver of the bearded seal can be a host for over a thousand О. arcti-
cus (V.V. Treshchev and M.V. Yurakhno). O. fraterculus is a parasite
of the gall bladder and Microphallus orientalis that of the intestines.
The cestodes infecting the intestines are: Diphyllobothrium cordatum,
D. hians, D. latum, D. lanceolatum, D. macrocephalus, D. schistochilus,
Diplogonoporus tetrapterus, Pyramicocephalus phocarum, and other gen-
era and species of Diphyllobothriidae; a single animal host could harbor
up to 44,000 specimens of these parasites (V.V. Treshchev). In some
bearded seals the cestodes parasitizing the intestines weigh one-sixth of
the weight of the organ itself. D. Janceolatum is encountered most often.
The nematodes infecting the stomach and intestines are Contracaecum
osculatum, Terranova decipiens, T. azarasi, and Phocascaris phocae. The
heart is infected with Skrjabinaria spirocauda and the lungs with Oto-
strongylus circumlitus, and Parafilaroides sp. The acanthocephalan worms
Corynosoma strumosum, C. semerme, C. validum, C. hadveni, and C. ven-
tronudum are intestinal parasites.

A comparison of the helminth fauna of E. b. barbatus and E. b. nau-
ticus revealed, in addition to worms common to both, helminths peculiar
to each.

The results of a study of 39 Atlantic bearded seals (V.V. Treshchev)
revealed only four which were unaffected while the remaining 35 (89.7%)
were infected with helminths and many of them quite heavily. All the ani-
mals in the age group of 1 to 15 years were infected. The most frequently
infected organs were the stomach (100%), small intestine (100%), duo-
denum (94.3%), and rarely the pancreas (34.3%) and the liver (22.8%).
A single animal could be parasitized by seven species of helminths, more
often five, rarely four, six, and even very rarely by three or seven species.
Of the 100 Pacific bearded seals studied (Delyamure and Serdyukov;
M.V. Yurakhno), seven were free from helminths while the remaining
93 were severely infected. Parasitic worms were recorded in the various
organs of all the animals in the age group 1 to 16 years. Infection of the
stomach and the intestines was 100%. The duodenum was often infected
(88.1%), less often the liver and gall bladder (48.3%), pancreas (25.8%),
and rarely the lungs (2.14%) and heart (1.07%). Parasitism of a single
bearded seal varied from 3 to 8 species of helminths. The animals were
more often infected by four and five species, rarely by six to eight and
more rarely by three species.*4

Population dynamics. Because of the lack of population data at any
initial level, its dynamics could not be determined accurately. At the end

34 These data were mainly compiled by helminthologists at the Crimean State University
under the guidance of Prof. S.L. Delyamure.

209

of the 1960s, the population evidently rose as a result of the cessation
of hunting from ships during the years of World War П and in the early
postwar period. The cessation of hunting by Soviet ships in the Barents
and Kara seas had a similar favorable impact from the mid-1960s.

The state of the Pacific bearded seal population in the Far East is
causing considerable anxiety. Its reduction in the Sea of Okhotsk has
been convincingly demonstrated by a sharp reduction in catch per ship.
From 1957 through 1963, the average number of Pacific bearded seals
caught by a single ship in the southwestern regions of the Sea of Okhotsk
decreased from 1,100 to 730, 1.е., by one-third. In the southern regions
of the sea (in Terpeniya Bay, on the eastern coast of Sakhalin), its popu-
lation has decreased so much that it has lost commercial importance. Its
total catch in the Sea of Okhotsk remained static until the mid-1960s only
by extending hunting into the northeastern regions of the sea, in She-
likhov Gulf, where the Pacific bearded seal was formerly almost left alone
by hunting ships (Fedoseev, 1966). Simultaneously, its population began
decreasing even in the Tauisk-Okhotsk region (Fedoseev and Shustov,
1964). The significant collections of this seal in the beach rookeries of
the Shantarsk Islands decreased. While in the 1930s thousands of this
seals could be counted in the rookeries, only a few hundred remained
by the 1960s. The hunting intensity of the Pacific bearded seal in the
Bering Sea should be established with allowance for exploitation of the
Okhotsk population.

Field characteristics. The Pacific bearded seal is a large, almost
monochromatic (sometimes with large light-colored spots visible from a
distance) animal with a relatively small head resting as though directly on
the shoulders, without a distinct neck. Light-colored dense and luxuriant
whiskers and relatively short, broad fore flippers, as though truncated,
are typical features. It is usually seen singly on ice floes and in water.
Encountered mainly on the coasts and in shallow waters in winter and in
spring, usually beyond the shore ice and among drifting ice. While diving,
its back is usually exposed and even the hind flippers on occasion. The
flippers and the air holes of the Pacific bearded seal are considerably
larger than those of the ringed seal. (K.Ch.)

Economic Importance

In the seminatural rearing of the local coastal population, the Pacific
bearded seal plays an extremely significant role in the European and
Western Siberian North (mainly for the Nenetz population) as well as
in the Far East (in the coastal Chukchi-Anadyr region, Koryak coast
and coasts of the Sea of Okhotsk). This valuable hunting target causes

162

210

no damage to the fishing economy. It is caught for its strong and thick
valuable hide, blubber, and meat. Its hide is a superior raw material
used locally for making soles of shoes, belts, etc. The skin (of fetuses
before bith) with strong well-preserved fur is used locally for making fur
goods. The skin of newborns with a strong hair coat is also sometimes
used. Compared with that of other seals, the meat of the Pacific bearded
seal contains the least blubber and is used at places even for human
consumption. Mainly, however, it is used by the collective and state farms
for feeding caged animals. The liver is toxic.

An adult animal, on average, yields 20 to 25 kg of hide (thickness
about 10 mm and area about 1.7 to 2.0 m2); for local needs the skin is
cut by cross “rings” into strips 60 to 75 m long with a width (in raw
form) of 2.5 to 3 cm.

The blubber weighs 75 to 125 kg in the summer when the animals
are least fed and the thickness of the fat layer averages about 4.5 to
5.0 cm. In the autumn-winter period a single adult yields up to 150 kg
or more of blubber. The meat of an adult without viscera often weighs
100 to 160 kg or more.

The popular method of catching is to shoot an animal resting on
an ice floe or sometimes showing above the open water. For this pur-
pose, a masked hunter in a whaleboat, motor boat, or canoe slowly edges
toward the animals among drifting ice floes or lies in wait with a rifle
along the edge of the shore ice. At places, mainly in autumn, the animals
are caught in a net. From the second half of summer, the animals are
killed in the coastal rookeries by beating them with sticks, as in the case
of the larga (p. 368).

Presently, the bearded seal is caught in the largest numbers in the Sea
of Okhotsk and the Bering Sea from ships which hunt for different types
of seals among the drifting ice floes. In the Sea of Okhotsk, until the
mid-1960s, the bearded seal held third place (next to ringed and ribbon
seals) among all the seals caught. Its catch there rose particularly from
1957, when 15,000 animals were caught with an average catch of 12,500
animals per annum for several years. These figures do not include a few
thousand caught by local organizations (Fedoseev, 1966a). In the Bering
Sea hunting of the Pacific bearded seal from ships occupies second place
(or shares it with the larga) after the ribbon seal.

The western hunting region, i.e., the southeastern part of the Barents
Sea, is now of lesser importance; from time immemorial our hunters went
there in the spring and later even the Norwegians. At present (1960s),
hunting operations are continued only by the Norwegian hunting ships in
the eastern regions of the Barents Sea where a few hundred bearded seals
are caught every year in the region of the “eastern ice floes”. In 1963, 324

163

211

animals were caught in this region while 1,239 animals were caught in the
“northern ice floes,” * 1.е., in the more western regions at Spitsbergen.

The locals hunt in the coastal zone in the Far East along the coasts of
the Chukchi Peninsula, Gulf of Anadyr, and along the Koryak coast, and
in the coastal regions of the Sea of Okhotsk, as well as in the west from
Kanin to Yamal and to a smaller extent at places in the eastern regions of
the Kara Sea and in the White Sea. As a result of such intensive hunting,
the Pacific bearded seal reserves in the Far East decreased sharply. To
prevent further depletion, hunting was regulated in the Sea of Okhotsk.
But hunting of the Pacific bearded seal by hunting ships should be totally
banned. The summer-autumn killing in the beach rookeries in particular
should be banned. In future, as and when the population of this seal is
restored, the nature and volume of its hunting should be fixed strictly in
accordance with the use of the raw material and the available reserves;
special attention should be devoted to census taking.

In the Bering Sea hunting ships should catch only that proportion
of the population as cannot lead to its depletion, and the native coastal
population should be involved, especially of the Chukchi Peninsula.

In the Barents Sea there is a need in the immediate future for more
coordinated hunting activity with the Norwegian expeditions based on
combined (or in any case coordinated) studies and rational distribution
of hunting quotas and also the institution of common hunting rules. To
increase the population in the White Sea, hunting should be banned
there (Potelov, 1969).

The entire system of economic utilization of the population of this
species should be reorganized. It is futile to catch pups of such a large seal
for the sake of fur (pups in the initial months alone are suitable for this
purpose). The fur quality of the skin requires further investigation. Also,
the quantity of such raw material cannot be significant because of the
peculiarities of hunting and the scattered distribution of the animal. It
would be more appropriate to use this animal for obtaining meat for ani-
mal food, the requirement for which has been steadily rising concurrent
with increased fur farming. The skin can be used as raw leather.

It is also extremely important that all countries ban the killing of
lactating females and pups in order to promote the normal restoration
of the Pacific bearded seal population. No less urgent is the problem
of systematizing the hunting and utilization of this seal by the coastal
population. (K.Ch.)

35 According to the annual report of the Norwegian Fishing Directorate on Seal Hunting
for 1963 (1964).

164

202

Genus of True Seals and Ringed Seals”°
Genus Phoca Linnaeus, 1758

1758. Phoca. Linnaeus. Syst. Nat. Ed. X, I, p. 37. Phoca vitulina Lin-
naeus.

1777. Pusa. Scopoli. Introd. Hist. Nat., p. 490. Phoca foetida Fabricius
= Phoca hispida.

1826. Callocephalus. F. Cuvier. Dict. Sci. Nat., 39, p. 544. Phoca vitulina
Linnaeus.

1844. Pagophilus. Gray. Zoology of Erebus and Terror, 3. Phoca groen-
landica Erxleben.

1864. Halicyon. Gray. Proc. Zoolog. Soc. Lond., p. 28. Halicyon richardi
Gray = Phoca vitulina richardi Gray.

1864. Pagomys. Gray. Proc. Zoolog. Soc. London, p. 31. Phoca foetida
Fabricius = Phoca hispida Schreber.

1864. Haliphilus. Gray. Ann. Mag. Nat. Hist., 17, p. 446. Phoca vitulina
richardi Gray.

1873. Histriophoca. Gill. Amer. Nat., 7, p. 179. Phoca fasciata Zimmer-
mann.

1904. Pagophoca ‘Trouessart. Cat. Mamm. Suppl.: 287. Substitute
Pagophilus Gray.>’(V.H.)

These seals are of moderate size or smaller, and are the smallest in
the family.

The facial portion of the head is moderately long (distance from the
eyes to the nostrils is a little more than the distance between the eye
and the ear opening). There is a narrow fringe of bare skin surrounding
the nostrils and between them. The whiskers have wavy edges. The first
digit or the first and second are the longest on the fore flippers.

The skull has: projecting rounded zygomatic arches and a narrow
interorbital space. The length of the zygoma, ignoring the processes, is
usually at least double its smallest width. The infraorbital foramen varies
markedly in size but in most cases is comparatively small and does not

36 Conforming to the relatively wide scope of the genus adopted in this publication,
almost all the true seals of the subfamily Phocinae have been grouped under the genus
Phoca (see below). (V.H.)

37 After Trouessart, 1904*, some authors (Ognev, 1935; and others) have used and
continue to use the generic name Pagophoca, considering that the name Pagophilus was
already assigned to Pagophila (birds—polar gull). However, according to Article 56a of
the International Code of Zoological Nomenclature, Pagophilus and Pagophila are not
homonyms and there is no need for Gray’s substitution of the name. (V.H.)

213

exceed or only slightly exceeds the maximum diameter of the alveoli of
the upper canines.

The nasal processes of the premaxillary bones are more or less
wedged between the nasal and the maxillary bones and do not reach
the nasal bones or just reach them only in one species (Ph. vitulina). The
nasal bones are fairly long and relatively narrow. The contour of the tym-
panic bullae proper (without lobes of the external auditory meatus) when
viewed from the lower surface, resembles a triangle with smoothened
apices or an irregular оуа1.38 The bony palate either terminates pos-
teriorly in a deep (more often angular, sometimes oval) notch, or is
without a deep notch, or forms a somewhat gentle, often double arc,
or even an almost straight transverse line. There is a compact bony
longitudinal septum in the choanae (within the palatine bones), or the
septum is almost lacking, or it terminates in the anterior or posterior
half of the palatine bones, or reaches the edge of the latter (Ph. groen-
landica).

The upper incisors have laterally flattened roots; the crowns of the
second and third premolars usually have additional cusps.

The color of the hair coat varies widely: usually spotted, in two
species in the adult state, consists of alternating large contrasting sec-
tions of light and dark colors (Greenland [harp] seal, Ph. groenlandica
and ribbon seal, Ph. fasciata) or is totally without spots (Baikal seal,
Ph. sibirica).

The primary hair coat in which the pups of all the species are born,
with one partial exception (in pagophobic members of the common seal,
Ph. vitulina), consists of long silky and dense wool mostly of a very light
color.*? Age-related color changes are characteristic of most species to
some extent or the other; they are most distinctly manifest in the Green-
land [harp] seal (Ph. groenlandica).

Sexual dimorphism is comparatively less prominent: adult males are
somewhat larger than the females but are not always distinguishable in
color. There are some differences in the skull proportions too.

There is one pair of teats.

The seals vary in ecological relations. Some are confined to the
coastal zone and are associated, especially during reproduction and molt,
with land while others lead an essentially pelagic life style and are associ-
ated with ice, on which they reproduce and molt. For the most part, these

38 Deviations of this characteristic are most common in the ribbon seal.

39 In the case of the Atlantic common seal and its ecological (pagophobic) Pacific coun-
terparts (see p. 355), the juvenile hair is shed in the mother’s womb itself, or at the time
of birth, or (very rarely) in the first few days after birth.

166

214

animals live in herds although they remain single during some periods
(especially when feeding and, in many species, during the pupping season
also). In most of the species, whelping occurs in comparatively narrow
limited periods of time, but this period continues from the end of Jan-
uary through July for the genus as a whole. The mating period proceeds
without harem formation. There is a latent period [delayed implanta-
tion] of 1.5-3 months in the development of the embryo. Mating is
followed by the onset of molt and later by a period of intense feeding.
Fish and various invertebrates, mainly crustaceans as also cephalopods,
serve as food objects. The fatness of the animal exhibits distinct seasonal
variation.

The geographic distribution is confined to the arctic and temperate
belts of the Northern hemisphere (Fig. 108). Within this zone, the range
encompasses the Atlantic, Pacific, and northern Arctic oceans in which
the seals are confined mainly to the continental zone; only the Green-
land and ribbon seals transgress regularly beyond the continental zone
while others do so mainly or occasionally with ice floes. Some species
inhabit the landlocked salt- and freshwater bodies (Caspian Sea, lakes
Baikal, Ladoga, and some others) and transgress into the lower courses
of rivers.

The southern boundary in the Atlantic Ocean along the North Amer-
ican coasts usually does not reach 45° N lat. while the boundary of nor-
mal distribution in the European seas does not cross south of Brittany
though some rare finds are known up to the Portuguese coasts. In the
Pacific Ocean, along the Asian coasts, the distribution zone extends from
the Bering Strait to the coasts of Japan (almost up to 35° N lat. on the
Pacific Ocean side of Honshu Island), the Korean Peninsula, and even
northern China (up to Shandung Peninsula and perhaps even up to the
Yantsiyang estuary).

In the Pacific Ocean, on the coasts of America, the range extends
down to Cedros Island, Baja California, Mexico (28°12’ М lat.). In the
North Atlantic Ocean at least one species of the genus (ringed seal,
Phoca hispida) inhabits all the peripheral seas, predominantly in the
continental terrace (or shelf) zone in the islands and archipelagos directly
adjoining the coasts as also away from the mainland, such as Severnaya
Zemlya, and the pelagic regions of the ocean. In the Central Polar Basin
it extends in rare cases almost up to the North Pole. One or the other
species of the genus inhabits everywhere in the seas covering the Soviet
coastline with the exception of the Black Sea.

Some species of the genus do not migrate far while others under-
take long and regular migrations. A characteristic feature of the latter
group is a special, fairly narrow localized region of winter concentration

215

"Алерипоа wisyinos ‘(irysdeyD yy) 2101255] (рэоцаошян) ‘ца Чеэз чодаы ay} рие 91рив]и2о48
(ппуао8ва) ‘ца ‘Теэз рииээло oy) Зирири! “| ‘$ 9эоца “еэз рэЗии эцз рие $1еэ$ эпл3 JO snuad ay) Jo э8ие1 [елэцэ0 ‘gol “314

Sol

a

т
ШУ 0605 COor 0606 0605 соот 0 Cool

167

216

where the seals reproduce and molt and a far more extensive region of
summer-autumn habitat (feeding ground) away from the former zone.
Not only the seals associated with the coast, but also those related to
some pagophilic forms when ice floes in their habitat thaw in summer,
form fairly concentrated rookeries on the coasts (larga, Caspian seal,
partly ringed seal, and Baikal seal).

The origin of the genus has been traced to Miocene ancestors which
morphologically are not exceptionally far removed from the present-day
species. Thus the sources of the genus extend into even older times. The
Pliocene finds (Belgium) help trace the branch quite reliably to the extant
species. Phoca vitulinoides Bened., Phoca halitschensis Blainv., and pos-
sibly others also are genetically related to the common seal. The species
Phocanella pumila Bened. and Phocanella minor Bened. lead undoubt-
edly to the present-day subgenus Pusa, a typical representative of which
is the ringed seal (Ph. hispida). Callophoca obscura Bened. from the for-
mations of the same age and place is regarded as a predecessor of the
present-day Greenland seal (Ph. groenlandica). The center of origin of
seals of the genus Phoca evidently falls in the Eurasian basin of the
Tethys.

The present level of study of the taxonomy of the genus covering
generic, subgeneric, specific, and intraspecific diagnosis is quite satis-
factory. Nevertheless, further research would lead to a more convincing
determination of the volume of the genus, to an accurate diagnosis of
the subgenera and species (particularly of the subgenus Phoca s. str.),
and to a better understanding of the actual intraspecific differentiation
of all the species. Much remains vague about the various aspects of ecol-
ogy, migrations, and population. The ribbon seal, Phoca (Histriophoca)
fasciata, remained the least studied species until recently, though there
is lack of adequate information even regarding other species, e.g., of the
subgenus Phoca s. str.

The genus consists of six extant species: common seal, Phoca vitulina
Linnaeus; ringed seal, Phoca hispida Schreber; Baikal seal, Phoca sibirica
Gmelin; Caspian seal, Phoca caspica Gmelin; Greenland [harp] seal, Pho-
ca groenlandica Erxleben; and ribbon seal, Phoca fasciata Zimmermann.
This genus is the largest in the family, covering 33% of its species. The
significant morphological difference between some of the species has led,
and continues to lead, some earlier as well as more recent authors (Schef-
fer, 1958; Chapskii, 1963; King, 1964) to divide the genus into some, usu-
ally up to four, genera. With the broader interpretation of the concept
of genus, also adopted by some recent authors (Ellerman and Morrison-
Scott, 1961*; and others), it would be more appropriate to regard the
natural groups of species as subgenera. Thus the genus comprises four

217

subgenera: Phoca Linnaeus, 1758, covering one species (PA. уйийпа);
Pusa Scopoli, 1777, with three species (Ph. hispida, Ph. caspica, and
Ph. sibirica); Pagophilus Gray, 1844, with one species (Ph. groenlandica);
and Histriophoca Gill, 1873, with one species (Ph. fasciata).

The species of the genus are almost equally represented in the basins
of the Atlantic, Pacific, and Arctic oceans. Three species each are encoun-
tered in the first two and four species in the peripheral part of the last.
Further, two species are found within the Palearctic in landlocked water
bodies: Caspian Sea (Ph. caspica) and Lake Baikal (Ph. sibirica).

The maximum population of these seals occurs in the arctic seas,
1.е., the northern and arctic Atlantic, especially in the polar regions, the
Sea of Okhotsk and the Bering Sea, the Canadian-Greenland waters,
the Barents and Kara seas, and also the northeastern part of the White
Sea.

All the species of the genus are of economic importance to some
extent while some represent the most important animals for hunting at
sea, especially the Greenland [harp] seal, Caspian seal, ringed seal, and
ribbon seal.

All the species of the genus are represented in the USSR fauna: in
the Baltic Sea—two species (Ph. vitulina and Ph. hispida); in the Bar-
ents Sea—three species (the above two and the Greenland [harp] seal,
Ph. groenlandica); in the Laptev and East Siberian seas and also in the
Central Polar Basin—one species (PA. hispida); in the Chukchi, Bering,
and Okhotsk seas—three species (Ph. vitulina, Ph. hispida, and Ph. fasci-
ata); and in the Sea of Japan (excluding Tatar Strait where three species
exist as in the Sea of Okhotsk)—one species (Ph. vitulina).

The species of this genus constitute 46% of the total number of
species of the order in the USSR fauna and about 1.8% of the total
number of species of mammals in the USSR.

In the Soviet territorial and internal waters as well as in the most
proximate international waters, the seals of this genus play an extremely
significant rele in marine-animal-based industries. The Greenland [harp],
Caspian, and ringed seals occupy first place in this respect. (K.Ch.)*!

40 As a result of recent studies (Chapskii, 1967, 1969), the monotypical state of this
subgenus has become extremely doubtful, if not erroneous.

41 For the key to species of the genus, see under the characteristics of the family (p.
151).

168

218

1776

1778.

1975.

1811.

1820.

1828.

1828.
1839.

1839.
1899.

1899.
1902.

1921.

1929.

1929.

1929.

1935.

Subgenus of Ringed Seals
Subgenus Pusa Scopoli, 1777

RINGED SEAL
Phoca (Pusa) hispida Schreber, 1775

. Phoca foetida. Fabricius in Muller, Zoologiae Danicae Prodromus,
р. VIII, Nomen nudum. Greenland.

Phoca vitulina botnica. Gmelin. Linn. Syst. Nat. Ed. XIII, 1: 63.
Gulf of Bothnia, Baltic Sea.

Phoca hispida. Schreber. Die Saugethiere, Table LXXXVI. 1776,
3: 312. Northern Atlantic.

Phoca ochotensis. Pallas. Zoographia Rosso Asiatica, p. 117.
Northern part of the Sea of Okhotsk, between Tauisk and
Gizhiginsk bays.

Phoca annelata. Nilsson. Scand. Fauna, 1: 362. Renamed
Ph. foetida Fabricius. Baltic.

Phoca schreberi. Lesson. Dic. Class. d’hist. Nat., р. 414. North
Atlantic.

Phoca communis. F. Cuvier. Dents mamm.

(Phoca communis F. Cuvier) B. var. Octonotata. Kutorga. Bull.
Soc. Imp. Nat. Moscow, p. 189. Neva.

(Phoca communis F. Cuvier) B. var. Undulata. Ibid., p. 191. Neva.
Phoca foetida var. saimensis. Nordquist. Acta Soc. Fauna Flora
Fenn., 15, 7: 28. Jake Saimaa, Finland.

Phoca foetida var. ladogensis. Nordquist. Ibid., р. 33, Ladoga.
Phoca (Pusa) hispida gichigensis. J. Allen. Bull. Amer. Mus. N. H.
16: 478. Sea of Okhotsk, Gizhiga.

Phoca hispida pygmaea. Zukowsky. Arch. f. Naturgesch. 87A, 10:
183. Barents Sea at 77°3’ М lat. and 49°40’ Е long. (pygmy ringed
seal); (V.H.)

Phoca hispida pomororum. Smirnov. Dokl. Ak. Nauk. (C. R. Acad.
Sc.) Leningrad, p. 95. Western coast of Novaya Zemlya.

Phoca hispida pomororum natio rochmistrovi. Smirnov. Dokl. Ak.
Nauk. (C. R. Acad. Sc.) Leningrad, p. 95. Sumsk environs, Onezhsk
Bay, White Sea.

Phoca hispida birulai. Smirnov. Dok. Ak. Nauk. (C. R. Acad. Sc.)
Leningrad, p. 96. Lyakhov Island, Novosibirsk Islands.

Phoca_ hispida krascheninnikovi. $. Naumov and Smirnov.
N. A. Smirnov, V. Adlerberg, Vinogradov, Smirnov, Flerov. Arctic
Animals. Leningrad, 1935. Bering Sea. (V.H.)

169

219
Diagnosis

The body length, including the tail, along the dorsal side (Lc) is not more ~
than 175 cm and the condylobasal length of the skull not more than
200 mm. The main background color of the hair coat on the upper side
of the body is dark, broken by a network of light-colored streaks, mostly
in the form of oval ringlets. The skull has a highly shortened rostral
portion: its length up to the commencement of the orbit is shorter than
the orbit (Fig. 89); the second to fourth lower premolars and molars bear
accessory cusps diverging from the base. The interorbital space is very
narrow (usually not wider than 7 mm in adults). The tympanic bullae are
relatively large, their length exceeding the gap between them, while the
width of the bony lobe of the external auditory meatus is-more than the
distance between its anterior edge and the crest of the articular fossa.
The anterior edge of the nasal bones bears three minute denticulate
processes. (K.Ch.)

Description

In body proportions the ringed seal resembles the other small seals, i.e.,
Caspian and Baikal seals. The fore flippers are shorter than the hind ones
and the first and second digits of the fore flippers are longer than the
third; the claws on them are of moderate size, with moderately elevated
upper margin, usually without such a narrow and high crest as in the
Claws of the Baikal seal. The head has a shortened snout (Fig. 109).
The labial whiskers are flattened and have wavy edges. The total number
of them (on each side) varies from 42 to 59; the supraorbital whiskers
number 3 to 6 and those near the nostrils one each. (Yu.I. Nazarenko)

nyt
aN
Wy Rte
AN)
|

ка DING NY era о
1 SOOT ered Г

Fig. 109. Ringed seal, Phoca hispida (figure Бу М.М. Kondakov).

169

220

Whitish gaps on a dark background, seen prominently in the form
of fairly distinct ringlets or curved streaks, are characteristic of the col-
oration of the hair coat. The distribution of these streaks in different
parts of the body surface is not very uniform. This pattern is most dis-
tinct and constant along the edges of the back and on the flanks but
absent throughout the entire median narrow zone of the back. It is also
absent on all the flippers and sometimes even on the ventral side which,
in most members of the population, is generally much lighter than the
back. In the anterior portion of the body, up to the shoulder blade and
the fore flippers, the ringed pattern is usually extremely fine and alter-
nates with light-colored variegations of different sizes and shapes. The
arrangement, size, shape, degree of clarity, and width of the rings vary
notably. They gather mainly along the edges of the back in wavy rows
like a typical network. Isolated ringlets scattered in different sections of
the upper side of the trunk are also seen. Sometimes the ringlets are
fused in twos, threes or more.

The main background color on the dorsal side varies from a оПуе-
gray to almost black. The ventral side in most of these seals is lighter
in color. When the color is uniformly vivid over the body, the underside
is generally slightly lighter because of the more intensely manifest and
diffuse ringed pattern (large number and greater width of the ringlike
streaks). When the dorsal side is darker than the ventral side, the color on
the flanks varies gradually. The rings are more often stretched along the
body length, slightly curved, and generally irregular in shape, especially
when the maximum number are crowded as though woven into a net.
Isolated clear zones are usually rounded and uniform. Their longitudinal
diameter does not exceed 10 cm, usually much less. In very rare cases
the ringed pattern is indistinct, dull, or lost altogether. These clear zones
become very small and disappear toward the head and the tail.

The upper side of the fore flippers is of the color of the adjoining
part of the body, sometimes with minute brownish specks or dabs. The
axillary portion is lighter in color. The color of the hind flippers is almost
monochromatic, dark, usually slate-black.

The color details of the hair coat, i.e., the ringlike pattern (number,
distribution, and degree of sharpness of contour of circular clearances,
their shape and size) and the main background color vary widely in dif-
ferent animals. Animals with a very bright or dull annular pattern are
also encountered; some have many rings forming a somewhat circular
pattern while these are few and isolated in others; the rings are some-
times large, sometimes small and light-colored streaks forming them are
not always closed; sometimes they are thick and sometimes. narrow and
their shape varies from nearly circular to oval; sometimes the rings are

221

170 highly elongated as though fused. There are по significant age-related

170

color changes (apart from the replacement of the embryonic pelage in the
pups). There are fewer ringlike streaks of light color on the skin of pups,
especially in their first year (after the first postnatal molt). Sex-related
color differences are absent.

The skull has a highly enlarged cranium, short and narrow snout, and
narrow interorbital space (Fig. 110). The width of the cranium imme-
diately above the mastoid processes exceeds its length measured from
the orbits to the posterior edge of the occipital condyles. The width at
the zygoma in most cases is nearly the same as the skull width at the
mastoid processes, sometimes perceptibly exceeding the latter. In adults
the length of the auditory bullae not more than their width, constitutes
20-23% of the condylobasal length. The anterior margin of the nasal
bones has three denticulate processes while the width of these bones at
the point of wedging into the frontals constitutes 15-20% of their total
length; they are wedged into the frontals to one-third to half of their
total length. The posterior margin of the bony palate is bracelike or has
a simple angular notch, or, in rare cases, is smooth. The longitudinal

\ Nt we ща `
ААУ Хх NS ь ei

SQ

Fig. 110. Skull of a ringed seal, Phoca hispida (figure by N.N. Kondakov).

171

222

bony septum in the choanae гип$ posteriorly not beyond the anterior
half of the longitudinal palatine suture.

The teeth, including the canines, are relatively thin and small. The
upper premolars, commencing from the second, usually have a single
accessory cusp anteriorly and another posteriorly; rarely, the anterior
accessory cusp is lacking or there are two accessory cusps posteriorly.
The second to the fourth lower premolars have a split fanlike crown
which usually has one accessory cusp anteriorly and two posteriorly. The
upper molar usually has three cusps while the lower one often has four;
gaps between the molars from the upper first to the fourth are usually
present.

The difference in the cranial dimensions of males and females is
slightly in favor of the former; differences are seen in the length and
various dimensions of the skull width. Age-related variations, however,
fall in the general pattern (see p. 221).

The diploid chromosome number is 32.

The body length of adults measured from tip of nose to tail end
along the dorsal curvature (Lc) varies widely in different populations,
individually and geographically, depending on age and sex, from 101
to 125 cm, possibly even more. The feeding conditions of the juvenile,
which evidently leave a mark on its entire life, play quite an important
role on the body length. The newborns which, for some reason or the
other, are underfed, remain far from attaining normal size by the time
they reach sexual and complete physiological maturity and quite often
remain puny. The body length of such pygmy seals differs little from that
of young ones.

The overall weight of the adults varies in different populations from
30 to 80 kg and, in the largest and well-fed animals, especially among
the Baltic ringed seals, the males can weigh up to 133 kg and females
125 kg (V.A. Zheglov). In the western arctic USSR the average weight
of adult males is 56.3 kg, of females 53.2 kg.

The relative weight of the internal organs (as percent of total body
weight) in Bering ringed seals weighing 24-32 kg was as follows (g): heart
23 - 87; liver 22 - 43; and lungs with trachea and larynx 9-22. The weight
indices of these same organs in Ladoga ringed seals weighing 30-42 kg
was: 47-105; 15-41; and 215-238. The small intestine was 14.7 m long
(Sokolov, Kosygin, and Tikhomirov, 1966).

The skull measurements of males from the eastern regions of the
Barents Sea and western part of the Kara 5еа“? were: condylobasal
length 161.5-182 mm; mastoid width 96.5-108.5 mm; width at the

42 According to the data of the Zoological Museum of Moscow University.

172

223

zygoma 94-109 mm; width at the upper canines 22-28.5 mm; and
interorbital width 3.8 - 6.2 mm. The skull measurements of females (from
the same regions) were: condylobasal length 159-173 mm; mastoid
width 95-109 mm; width at zygoma 99-105 mm; width at the upper
canines 22-29 mm; and interorbital width 4-7.3 mm. The dimensions
of ringed seals within the USSR reveal significant geographic variation
(see pp. 231-234). (K.Ch.)

Taxonomy

Taxonomically, the ringed seal is most closely related to the Baikal
(Phoca sibirica Gmelin) and Caspian (Phoca caspica Gmelin) seals along
with which it forms the subgenus Pusa or the true seals. Some authors
have assigned this subgenus the rank of genus (Scheffer, 1958; Chap-
skii, 1963; King, 1964). Concomitantly, attempts were also made to clas-
sify Ph. sibirica and Ph. caspica only as subspecies of Ph. hispida, a
view that did not find favor. Among the other extant members of the
genus Phoca 5. 1., the common seal or larga (Phoca vitulina Г.) is clos-
est to the ringed seal in systematic position and evolution. Sometimes
the two are combined under a single subgenus. Both the subgeneric
(or generic) branches have been identified at least from the Upper
Miocene.

The interrelations between members of the group Pusa are not yet
entirely clear. The prevailing view is that the ringed seal is closer than all
the others to the original ancestor in the line from which the two other
Closely related species evolved by some type of invasion from the north
and isolation in landlocked reservoirs at a very early stage of its geo-
logical history. Another view holds that the primary subgeneric branch
evolved in a more southern landlocked sea derived from the Tethys, if
not in the Tethys per se (Chapskii, 1968).

A morphological comparison points to a closer affinity of the ringed
seal to the Baikal seal rather than to the Caspian seal though a crosscheck
of the craniological features reveals a fairly variegated picture (Chapskii,
1955b). (K.Ch.)

Geographic Distribution

Covers the peripheral seas of the polar basin, rarely its central regions,
the arctic and subarctic Atlantic seas, the northernmost and northeastern
parts of the Pacific Ocean (Bering Sea and the Sea of Okhotsk), Baltic
Sea, Lake Ladoga, and the Saimaa lake system.

224
Geographic Range т the USSR

Accounts for no less than half the total range of the species. In the seas
of the arctic belt the distribution of the ringed seal extends continuously
from the western to the eastern boundaries of the USSR (Fig. 111).

It inhabits the southern, eastern, and northern marginal portions of
the Barents Sea, encompassing the belt’ of ice floes from Novaya Zemlya
to the meridian of Spitsbergen, but is not reported from the pelagic
central and western parts of the sea. The boundary of the range in this sea
runs close to the Murman shores in the east, turns then to the south of
the White Sea Inlet, turns arcuately roughly from the meridian of Cape
Kanin Nos to the northeast and, having reached roughly the latitude
of the northern extremity of Novaya Zemlya, turns westward. In the
coastal waters of Murman it is distributed almost everywhere with a
preference for sections with a more rugged coastline quite protected
from the surf, especially river expanses in estuarine zones. It may be
assumed that the ringed seals present on the coasts of western Murman
represent recent arrivals since suitable biotopes for breeding are not
available there.*?

The range in the White Sea covers all the bays except the inlets and
straits. The ringed seal often swims into the lower reaches of the rivers.
In Dvinsk Bay it is distributed almost everywhere in the peripheral zone
in the south to the very mouth of the Northern Dvina but is evidently
scarce in its northwestern open part as also in the Central Basin. It is
quite common almost everywhere in Kandalaksha Bay, right up to its
uppermost (cul-de-sac) part; in the central regions of the bay, however,
it is sporadic. It is obviously confined to the northern half in Onezhsk Bay
as also in its western regions abounding in islands and with a particularly
rugged coastline.

It is quite common in Mezensk Bay but prefers the coastal belt.
On the whole, the distribution in the White Sea is quite extensive but
it usually avoids the extreme central portion of the sea and the straits
away from the coasts. This is true of the summer-autumn as well as
winter-spring period. In summer it is common even in the northern

43 Scant, fragmentary, but nevertheless concrete information on the distribution of the
ringed seal on Murman coasts is scattered in the works of Pleske (1887), Knipovich and
Yagodovskii (1902), Knipovich, Yagodovskii, and Zhikharev (1902), Soldatov (1902), Bre-
itfus (1903, 1905, 1912, 1915), М. Smirnov (1903), Wolleback and Knipovich (1907*),
Formozov (1929), Sdobnikov (1933*), and Surkov (1965*).

(ysdeyD УМ) USSN ou UI ири1и vooYg ‘fees рэ8им ay) Jo эвиеч “TTT “314 ELI

174

226

regions of the White Sea and forms rookeries Шеге“ on the bars and
spits. In the eastern inlets of the White Sea the ringed seal inhabits
the Kanin-Kolguev shallow waters, Cheshsk Bay, the Timansk coastal
waters and the Pechora Sea up to Yugorsk Shar, Vaigach, Kara Strait
and southern parts of Novaya Zemlya inclusive, with a preference for
peripheral coastal expanses.

In the more northern regions of the eastern extremity of the Barents
Sea, the distribution extends in a strip that is not very broad along Novaya
Zemlya up to the latitude of its northeastern extremity. Here the ringed
seal is confined preferentially to the bays, especially those which project
deeply into the land with highly rugged coasts, and not to the open spaces
or capes jutting into the sea. It also inhabits Matochkin Shar.*°

From the northeastern extremity of .Novaya Zemlya the boundary
of distribution turns morthwest and extends through the zone of drifting
ice floes into the region of Franz Josef Land archipelago. Here and in
its immediate icy environs the ringed seal is quite common and is found
almost everywhere. It is particularly numerous in the straits in summer
but not sighted in winter, evidently because concealed by the ice cover.*”

The completely isolated Baltic portion of the range of the: ringed
seal covers the regions of all our territorial waters in the Gulf of Finland
including Nevsk Bay (where, however, it is rare), the region of Sarema
and Hiiumaa islands from the contiguous strip of the Baltic Sea itself,
extending northwest, and also the entire Gulf of Riga.*

Lake Ladoga constitutes a distinct section of distribution.

44Some data of a very special nature on the distribution of the ringed seal in var-
ious regions of the White Sea were drawn from Danilevskii (1862), Knipovich (1897,
1907*), Zhitkov (1901, 1904), N. Smirnov (1903). V. Nikol’skii (1927), Bianki (1965),
Golenchenko (1961, 1963*), Nazarenko (1967, 1968), and others. The unpublished data of
А.Р. Golenchenko, Yu.I. Nazarenko, V.A. Potelov, and others were also used here.

45 For the characteristics of distribution of the ringed seal in the southeastern regions
of the Barents Sea, data from the following published and unpublished sources were used:
Golenchenko (1961), Zhitkov (1903*, 1904, 1913), Klumov (1935), Moskalenko (1945),
А.Р. Golenchenko, Yu.I. Nazarenko, V.A. Potelova, Tsapko (1958*), and others.

46 Based on the published and unpublished observations and information of several
researchers: Krivosheya (1884), Gorbunov (1929), Lepin (1935*), Klyuge (1936),
M.I. Vladimirskaya, А.Р. Golenchenko, А.М. Dubrovskii, A.I. Zubkov, V.A. Potelov, and
К.К. Chapskii.

47 More detailed information on the distribution of the ringed seal in the Franz Josef
Land archipelago can be found in the works of Nansen (1901*), Al’banov (1917), Pinegin
(1934*), Esinov (1935), and Tsalkin (1936). The oral communications of L.I. Leonov,
hunters, and others were also used here.

48 This general review of the distribution of the ringed seal in the Baltic was drawn
mainly from the works of Greve (1906*), Schubert (1929), Freund (1933), Ropelewski
(1952), Аш, Ling, and Paaver (1957), Bergman (1960*), Leis (1960), and V.Z. Zheglov.

175

227

The distribution of the ringed seal in Lake Ladoga changes depend-
ing on the time of year, ice drift, and finally on the migration of some
of its food objects. In summer and autumn the seals confined predom-
inantly to the coastal zone go farther away from the coasts in winter,
especially in the shoals; in spring they may be transported even into the
central portions of the lake together with drifting ice floes. In summer
and autumn these seals are concentrated at places in the immediate prox-
imity of the coast or toward the islands and rest on individual boulders,
rocky “аз” [littoral shoal or islet], and sand bars.‘?

In the Asian sector of the arctic the southern boundary of the range
runs all along the coast of Western and Eastern Siberia to the Bering
Strait. Northward, however, the range extends up to the Central Polar
Basin (Chapskii, 1949; Rutilevskii and Uspenskii, 1957). A more com-
plete picture of the distribution of the ringed seal is available for the
Kara Sea. Here it is found everywhere to some extent but preferentially
in the coastal belt, from Novaya Zemlya to Vil’kitsk Strait and Sever-
naya Zemlya. On the eastern coasts of Novaya Zemlya, it is distributed,
although not very uniformly, all along the stretch from Cape Zhelaniya
to the Kara Strait. Southeast of the latter, it inhabits the coastal belt of
Vaigach and Yugorsk Shar. It inhabits Baidaratsk Bay and the western
coast of Yamal. The central regions of this part of the sea also fall into
the range since the ringed seal is encountered there at different places
along with ice floes, at least in the summer season. Farther east, it inhab-
its Malygin Strait and generally the waters of White Island, Gulf of Ob
with much of Tazovsk Bay, the expanse from Shokal’sk Island to Yenisey
Gulf (including Gydayamsk Bay), the region of Dixon Island, Pyasinsk
Bay, and the coastal belt of western Taimyr to Vil’kitsk Strait. Along the
western rim of Severnaya Zemlya, the range rises to its northernmost
extremity and runs far northward. The ringed seal is encountered on all
the islands of the Kara Sea, even such islands as Uedineniya and Vize~°
which are far away from the continental coasts.

4 The distribution of the Ladoga seal has been described from the data of Chapskii and
co-authors (1932*), Golenchenko (1935), М. Smirnov and colleagues (1954*), Sorokin and
co-authors (1957, 1958*), A.S. Sokolov (1958*), А.А. Antonyuk, A.I. Zubov, P.V. Fil’kin,
and others.

°° This summation of the distribution of the ringed seal in the Kara Sea is based on the
observations of several individuals and on data from the literature, including Zhitkov (1913),
Heptner (1930, 1936), S. Naumov (1931), Kolyushchev (1933), Probatov (1933), Klu-
mov (1935), Urvantsev (1935), Kirpichnikov (1937), Mikhel’ (1937), Chernigovskii (1935),
Antipin (1939*), Rutilevskii (1939), Laktionov (1947*), Ushakov (1953), Mikhailov (1958),
G.G. Galkin, L.I. Leonov, У.Е. Nikitin, V.A. Potelov, А.М. Tyulin, К.К. Chapskii, I.K. Yaki-
movich, and others.

228

Information on the ringed seal in the extreme east is highly
incomplete. It is quite widely distributed in the Laptev Sea though very
unevenly, as in thé western part, close to the coast of eastern Taimyr (on
Komsomol’sk Pravda Island, in Faddei Bay, and in the Pronchishcheva
Bay region), in Khatanga Gulf, in the region of Begichev Island, and
in Nordvik Bay, and very far in the east along the coastal strip. It is
encountered even in the bays of such rivers as the Anabar, Olenek,
Lena, and, in the more northern regions, especially in the eastern part
of the sea, on the threshold of the Dmitrii Laptev and Sannikov straits,
and in the zone of the Novosibirsk Islands. More than at other places,
ringed seal sightings are reported from the region of Begichev Island and
southwest of it (Koshkin, 1937). In general, however, this sea is regarded
as far from abounding in ringed seals (Mikhel’, 1937).

In the East Siberian Sea the ringed seal inhabits the coastal belt
and evidently all the bays, estuaries, and foredelta sections as well as the
pelagic regions of drifting ice floes; it reaches the Novaya Siberia and De
Long islands. In the western regions of the sea, west of the Medvezhii
Islands, the ringed seal is considerably less numerous than east of the
Kolyma estuary; it is most numerous in Chaunsk Bay and the adjoining
coastal belts. It was encountered, though not often, in the more northern
pelagic regions, including in the proximity of Wrangel Island.>!

It is common in the Chukchi Sea along the entire coastal belt but
because of the shallow waters, it is quite often confined more toward
the open sea. Its most abundant regions are near Cape Serdtse-Kamen’,
Kolyuchin Bat Inlet, and northwest of Kolyuchii Island (Fedoseev,
1965c). The northern boundary of distribution in the Central Polar
Basin is not amenable to precise determination. It is drawn (Scheffer,
1958) tentatively along 85° lat. but innumerable cases of the appearance
of ringed seal are known even more northward, even at latitudes 88°
and 89° or a few minutes farther away (Chapskii, 1949; Uspenskii and
Rutilevskii, 1957*). The entire Central Polar Basin should perhaps be
included in the range of this species though the ringed seal is extremely
rare in the fore-polar regions and is seen evidently only in the spring-
summer months.

In the Far Eastern waters, south of the Bering Strait, the range of
the ringed seal encompasses the entire mainland rim of the western part
of the Bering Sea. On the eastern and southern coasts of the Chukchi
Peninsula, the ringed seal is extremely common and numerous and is
most concentrated in winter and spring in a wide belt of coastal ice floes

51 According to the data of Iokhel’son (1898), Buturlin (1913), Arsen’ev (1935), Mikhel’
(1937), Fedoseev (1966b), and others.

176

229

all along the stretch from Cape Dezhnev to Provideniya Bay and farther
west in the regions of Rudder Spit and especially Krest Bay (Fedoseev,
1965c). The ringed seal is also common in the Anadyr drowned river
valley [Gulf of Anadyr] and the surrounding coastal waters; it often
transgresses in summer into the lower courses of the Anadyr and other
rivers, Sometimes even extremely small rivers and rivulets; it is seen mov-
ing along the Gulf of Anadyr coasts, remaining long and even winter-
ing at places in the bays, straits, and on open beaches (Portenko, 1941;
V.N. Gol’tsev). It is encountered, though less numerously, on the Koryak
coast (N.B. Shnakenburg).

Southwest of Cape Olyutorsk, the ringed seal inhabits the entire
coastal belt, descending to Apuka, Гр! Bay (Anastasia), Parapol’sk
Valley and Karaginsk Island, and the eastern coast of Kamchatka
(Samorodov, 1939; Averin, 1948; L.A. Portenko and F.B. Chernyavskii).
It has been reported from time to time in the sea, far away from the land,
mostly in the season of drifting ice floes. In the Bering Sea the ringed
seal usually inhabits only the coastal zone (Rozanov, 1931; Razumovskii,
1933; Shustov, 1967, 1968*; E.A. Tikhomirov; and others). This is also
confirmed by the 1964 observations of K.K. Chapskii.

It is extremely difficult to draw the boundary of distribution in the
open sea beyond which the ringed seal does not enter the south and the
east. The mean position of the ice edge in the winter-spring period can
partly and only tentatively serve as such a boundary line. Only in the
proximity of the continental coast does the boundary descend to the lati-
tude of the Commander Islands >* (Barabash-Nikiforov, 1935*; Marakov,
1964*, 1968); still more southward, in the coastal zone of Kamchatka,
it barely reaches Cape Lopatka along the eastern coast. A find on the
northernmost edge of the eastern side of the Kuril Range is quite difficult
to explain.

The distribution of the ringed seal-in the Sea of Okhotsk covers the
entire northern part of the sea in a broad belt from west to northeast,
from the Gulf of Sakhalin and the Shantarsk Sea up to Gizhiginsk -and
Penzhinsk bays more deeply projecting southwest, and also the Tigil’sk
region and the adjoining sections slightly south of western Kamchatka.
From the outer edge of this strip along the eastern edge of the sea,
along the eastern coast of Kamchatka, the range arcs into a somewhat
less broad strip southward and comes to naught on the northern islands
of the Kuril range. In the southwestern part of the sea, however, the space
covered by the ringed seal extends in a broader strip along Sakhalin up to

52? On the Commander Islands themselves, “random finds of stray animals” (including
gestating females) have been reported at various points on the coast (Marakov, 1968).

176

177

230

Fig. 112. Distribution of the Okhotsk ringed seal, Phoca hispida ochotensis, in
the breeding period and its migratory course for molting in the Sea of Okhotsk
(cross-hatching) and in 1969 (horizontal hatching) (G.A. Fedoseev).

its southern extremity and up to the southern Kuril Islands. It is difficult
to draw the exact boundary of distribution in the central pelagic regions
of the sea since this depends to some extent on the movement of ice floes
and is therefore extremely variable not only in the course of the annual
cycle, but also from year to year. It could be represented schematically as
slightly receding northwest of the long-time mean position of the winter
ice edge. As a result, the southern boundary of distribution in the Sea of
Okhotsk runs along a fairly steep arc whose curve faces northwest, with
one end approaching almost the northernmost and the other almost the
southern Kuril Islands.

In the Sea of Japan the ringed seal inhabits only Tatar Strait, entering
it in the south roughly up to the latitude of De Kastri or slightly more
southward (Dorofeev, 1935*).

Geographic Range outside the USSR

In the arctic and North Atlantic Ocean the ringed seal occupies the
northern regions of the Norwegian waters (usually this is the Finnmark
region but in particularly cold years the boundary shifts to Lofoten or
more south), Spitsbergen archipelago and the strip of arctic pack ice
in the west up to the coastal waters on the eastern side of Greenland,
and in the north almost up to 75° N lat. Farther west, the range covers
the strip along the western coast of Greenland in the north to Kane

178

23

Basin inclusively, the Labrador coast, northern edge of Newfoundland,
the northernmost part of St. Lawrence Bay (in the south up to 50° М
lat.), almost the entire Canadian archipelago, Hudson Bay, and Hudson
Strait (Fig. 113). :

In the Baltic Sea Basin the distribution covers the Gulf of Bothnia,
the zone of Aland Islands, the Baltic Sea proper adjoining this zone,
and the entire northern part of the Gulf of Finland west of the USSR
boundary. Lake Simaa in Finland represents an isolated section.

East of the Bering Strait, the distribution covers the eastern part of
the Chukchi Sea and southern regions of the Beaufort Sea; in the Bering
Sea it covers the northeastern extremity of the Bering Sea adjoining
Alaska, and to the south up to the northern edge of Bristol Bay. In the
Sea of Okhotsk the distribution includes the northern coastal waters of
Hokkaido. (K.Ch.)

Geographic Variation

The extensive but fragmented distribution of the ringed seal suggests a
fairly significant intraspecific geographic variation. Soviet waters are host
to six subspecies (N. Smirnov, 1929, 1935; Ognev, 1935). At present, due
to lack of adequate conclusive proof, not all are recognized (Chapskii,
1952). The diagnosis of the entire subspecies should be reviewed afresh
with the exception, probably, of the Far East ringed seal, which has been
fairly well detailed (Fedoseev, 1965c; Fedoseev and’ Yablokov, 1965).

The following subspecies have been recognized in the ranges falling
within the USSR.

1. Baltic ringed seal, Ph. (P.) h. botnica Gmelin, 1788 (syn. annelata).
This subspecies is almost the largest in size (V.A. Zheglov). The
color is deep dark, often almost black, especially the dorsal background
which bears a “lacy” network of light-colored ringlets. The condylobasal
length of the skull is 158-187 mm (average 163.2) (Ognev, 1935).
Eastern part of the Baltic Sea and Gulfs of Finland and Riga.
Outside the USSR, it has been reported in the northern, western,
and southern parts of the Baltic Sea. 53

2. Ladoga ringed seal, Ph. (P.) h. ladogensis Nordquist, 1899.

The size is only slightly smaller than that of the preceding subspecies.
Proximate in color to the Baltic ringed seal but usually slightly lighter
(V.A. Zheglov). The average body length of the female including the tail

53 The greater morphological proximity of the Baltic subspecies to the Ladoga and
Saimen subspecies than to the Pomorsk subspecies (White Sea populations) has also been
confirmed by the latest craniometric data (Muller-Wille, 1969).

232

Fig. 113. Range of ringed seal, Phoca hispida, southern boundary (К.К. Chapskii).

177

233

along the dorsal curvature is 125.3 cm. The condylobasal length of the
Skull is 167-184 mm (average 173.3) (A. Sokolov, 1956).

Lake Ladoga.

Not reported outside the USSR.

3. Pomorsk ringed seal, Ph. (P.) h. pomororum Smirnov, 1929 (syn.
Pygmaea rochmistrov1).

The size is relatively large. The color shows no black tones and the
ventral side is usually lighter.

The body length including the tail is 111-153 cm along the dorsal
curvature; males average 127.6 cm and females 126.6 cm (Chapskii, 1940).
The condylobasal length of the skull is 167-189 mm (х = 177.2) (and
even 201) (Ognev, 1935).

White and Barents seas and at least the western part of the Kara
Sea.

Outside the USSR, it is probably found in the adjoining regions of
Norway.

4. Siberian ringed seal, Ph. (P.) h. birulai Smirnov, 1929.

Its size is large. The color of the hair coat is lighter than that of the
Pomorsk ringed seal. The condylobasal length of the skull is 179 - 195 mm
(x = 185.7) (Ognev, 1935).

Seas of the eastern arctic sector: from the eastern part of the Kara
Sea to the threshold of the Chukchi Sea.

Not reported outside the USSR.

5. Bering Sea ringed seal, Ph. (P.) h. krascheninnikovi Naumov and
Smirnov, 1935. |

It is of moderate size, smaller than the Siberian subspecies. Its color
characteristics have not been determined. The condylobasal length of the
skull is 152.8- 189.5 mm; males average 174.2 mm and females 168.2 mm
(Fedoseev, 1965b).

Bering Sea.

Outside the USSR, it has been sighted in American waters of the
Bering Sea.

6. Okhotsk ringed seal, Ph. (P.) h. ochotensis Pallas, 1811 (syns. gischi-
gensis, nummularis).

It is of small dimensions, the smallest form in the USSR waters.
Color characteristics have not been established. The body length includ-
ing the tail is 101-135 cm along the dorsal curvature; males average
117 cm and females 116 cm (Tauisk Bay; Fedoseev and Yablokov, 1965).

179

234

The condylobasal length of the skull is 152.2 - 176 mm (x = 163) ($. Nau-
mov and N. Smirnov, 1936).

Sea of Okhotsk.

Outside the USSR, found in the waters of Hokkaido.

* * *

The following subspecies are usually recognized in the waters outside
the USSR: (1) Ph. (P.) h. hispida Schreb., 1775—western Atlantic Ocean
including Greenland, Canadian waters up to the Beaufort Sea; (2)
Ph. (P.) h. saimensis Nordquist, 1899—Lake Saimaa in Finland; (3)
Ph. (P.) h. beaufortiana Anderson, 1943—-western polar Canada, Beaufort
Sea; and (4) Ph. (P.) h. soperi Anderson, 1943—Baffin Island lakes.

The last two subspecies require thorough verification; they are some-
times placed (Scheffer, 1958) among the synonyms of P. hispida hispida.
Judging from the latest data (Fedoseev and Nazarenko, 1970), no differ-
ences whatsoever could be identified between the populations inhabiting,
on the one hand, the southeastern part of the Barents Sea and, on the
Other, the northern part of the Bering Sea. Evidently the entire arctic
belt of Eurasia is inhabited by only one “Pomorsk-Siberian” subspecies
or even the monotypical subspecies Phoca hispida hispida Schreber, 1779.
In the latter case the rest of the forms of ringed seals indicated for our
northern seas from the Barents to the Bering, 1.е., the Pomorsk, Siberian,
as well as the Bering subspecies, should be regarded as synonyms of the
nominal form. (K.Ch.)

Biology

Population. Among the total population of seals in our waters, the ringed
seal holds first place. A preliminary assessment of the total population
in the USSR waters gives a figure of three million (Chapskii, 1966).>4
The majority, about 2.0-2.2 million, inhabit the polar sector from the
eastern part of the Barents Sea to the Bering Strait.

The quantum of ringed seals inhabiting the Sea of Okhotsk, roughly
calculated by applying the age-related structure analysis of animals
caught at random, recognizing the relative proportion of the mothers
(21%) and the proportion of annual births, etc., was placed at a maximum
of 800,000 (Fedoseev, 19665, с). Similar figures, 1.е., 600,000 to 865,000
were arrived at in subsequent aerial surveys (Fedoseev, 1968). The

54 The calculation is based on a quantitative assessment of the reserves of the ringed
seal in the waters of the eastern part of the Canadian archipelago, where they are put at
almost one million (McLaren, 1958) and extrapolating this value to the USSR range (with
allowance for the varying habitation of the ringed seal in different sections of the range).

180

235

differences in the calculations are explained as due to error in the method
of calculation (С.А. Fedoseev).

In the Bering Sea ringed seals are evidently fewer than in the Chukchi
and East Siberian seas (Tikhomirov, 1966b). In 1964, aerovisual surveys
estimated 12,000 ringed seals on ice floes in the western part of the
Bering Sea (Shustov, 1969b).>°

From our present knowledge, it is impossible to establish the actual
population of ringed seals in the Chukchi, East Siberian, and other arctic
seas. Evidently in each of the arctic seas the reserves of ringed seals are
extremely divergent. It can only be assumed that in the USSR part of
the Chukchi Sea the ringed seal is more abundant than in the Bering
Sea and less so in the Laptev than in the East Siberian Sea. Evidently
the two latter seas are next to the Kara Sea in this respect. Possibly the
Barents Sea has a higher population of ringed seals than the Kara Sea
and evidently the White Sea also.

In the Sea of Okhotsk, as in the other seas, the ringed seal is very
unevenly distributed. In April, during the breeding and lactation period,
most of the animals localize mainly in three regions: the northwestern
part of the sea (near the mainland coast from Tauisk Bay to Ayan),
Shelikhov Gulf, and on the eastern coast of Sakhalin, including Terpeniya
Bay. The bulk of the population is seen every year in the northwestern
part of the sea. In the other regions (in Shelikhov Gulf and on Sakhalin),
the population of the Okhotsk ringed seal varies significantly from year
to year depending on the position of ice floes suitable for breeding.
Thus in 1968, during the first 20 days of April, only 3,500 pups and
25,000 adults were counted (aero-visual survey) on ice floes and in the
waters of Sakhalin; in the very next year, however, in the first ten days of
the same month, there were 32,000 pups and 106,000 adults (Fedoseev,
1970). An animal density on ice floes during the pupping season of, on
average, 1.5 animals per km? is not high.

Rookeries of molted animals are found in May when intense ther-
mal and dynamic break-up of the ice floes occurs. The animals actively
migrate to those regions where the ice floes remain intact longer. Large
collections of the molted Okhotsk ringed seal are usually noticed in the
region of Eirineisk Bay and in the Sea of Okhotsk, Khanyangda-Ayan,
on the Shantarsk Islands, in Sakhalin and Shelikhov gulfs, and on the

55 This figure cannot, in fact, reflect the actual size of the population (only the animals
seen on ice floes were taken into account). The actual population in our part of the Bering
Sea is higher. The figure indicated constitutes only a third, until recently, of the total annual
catch of the ringed seal on the coasts of the entire Chukchi Peninsula (see p. 257) and only
double the number of ringed seals caught in 1960 on the southern and western coasts of
the Chukchi.

’_ 236

northeastern coast of Sakhalin. This seal is encountered in small numbers
on the northeastern coasts of Kamchatka. Stray ringed seals are encoun-
tered in the northwesternmost part of the sea adjoining La Perouse Strait
(G.A. Fedoseev).

In the Bering Sea almost all the animals are confined year round to
the coastal belt extending northeast of Karaginsk Island to the Gulf of
Anadyr. The ringed seal is evidently concentrated in large numbers in
the Gulf of Anadyr. On the southern and eastern coasts of the Chukchi
Peninsula, the maximum concentrations during reproduction and molt
are noticed in Krest Bay, in the region of Rudder Spit, and all along the
coast of the Bering Strait to Cape Dezhnev.

In the Chukchi Sea, the sections close to Cape Serdtse-Kamen’, the
region of Kolyuchin Bay, and slightly northwest of Kolyuchin Island, are
prominent in abundance of the ringed seal (Fedoseev, 1965c). In the
East Siberian Sea, the ringed seal is more numerous, even abundant in
summer on the Medvezhii Islands and generally in the eastern part of
the sea; on the contrary, on the Novosibirsk Islands it is rare (Mikhel’,
1937). The southwestern part (region of Begichev Island-Nordvik Bay of
the Laptev Sea), as far as can be judged from the scant data (Koshkin,
1937; L. Popov, 1941*), is quite rich in the ringed seal.

In the Kara Sea the ringed seal is more numerous on the north-
western coast of Yamal, in the region of White Island, and the northern
portion of Gulf of Ob, on Dixon Island, and in the adjoining sections of
Yenisey Bay, in the region of the Pyasine estuary, Minin Sea cliffs, and
on Cape Sterlegov. Significant congregations are seen in the midpart of
the Novaya Zemlya strip of the sea, especially from Matochkin Shar to
the Pakhtusov Islands. At almost all these places the ringed seal is not
a permanent resident; it is seen in spring and sometimes in summer or,
on the contrary, abandons these regions in summer and autumn.

The chief sites of concentration of the Barents Sea ringed seal are
the southeastern coastal regions of the sea in which the largest number of
arrivals is seen in the autumn-winter months: the regions of Kambal’nitsa
and the adjoining islands (northeastern coast of Kanin), some sections of
Cheshsk Bay, regions of Sengeisk Island (with the strait), Kolokol’kovsk
Bay, Pechora Bay, Varandei Island, Khaipudyrsk Bay, western “mouth”
of Yugorsk Shar, southeastern coast of Kolguev Island, rock cliff in the
southern part of Novaya Zemlya including the region of Mezhdusharsk
Island, Belush’ei Bay, and others.

The population and regions of maximum concentration of ringed
seals in the White Sea are as yet not adequately known.

In the USSR territorial waters of the Baltic, based on aero-visual
observations, about 12,000 ringed seals were estimated (V.A. Zheglov;

181

182

237

Zheglov and Chapskii, 1972*). Of these, 8,000 were in the USSR
territorial waters of the Gulf of Finland and about 4,000 in the Gulf
of Riga. The total population of the Baltic ringed seal at present is
estimated at roughly 50,000 (V.A. Zheglov). In winter the seals stray far
from the coasts and are confined mainly to deep-water zones; depending
on the formation and thawing of ice floes, these animals move into
shallow-water sections in the environs of islands (V.A. Zheglov).

The Lake Ladoga population is less than in the past (Chapskii,
1932*; and others); it probably does not exceed 5,000-6,000 (A.A.
Antonyuk).

Habitat. As the ringed seal belongs to the group of pagophilic seals
(associated with ice floes), it usually inhabits those water bodies which
are icebound, at least in winter. It selects for breeding predominantly the
coastal, stationary ice floes. Only the Okhotsk ringed seal deviates from
this rule and probably the Chukchi ringed seal at places. As a result of
intense tidal currents in the Sea of Okhotsk, stable shore ice is not formed
and the Okhotsk ringed seal is forced to use the broken-up ice floes drifting
in the relative proximity of the coast for breeding and molt. Further, this
seal undergoes parturition at some distance from the edge turned toward
the coast, selecting fairly firm, somewhat piled-up ice floes. Any fairly
firm ice floe with air holes in the vicinity serves as a whelping site. In most
cases, pups lie in the open and not under a snow cover (N. Smirnov, 1911;
Tikhomirov, 1961; Fedoseev, 1964b*, 1965; and others).

In all the other regions the ringed seal is confined during the breéed-
ing season to shore ice or coastal ice floes concealed under a snow cover
from the gaze of passersby. The pups are delivered in snow caves on
the ice floe, near an air hole, or in hollows formed among heaps of
broken-up ice floes.

The young animals (Fig. 114) not participating in reproduction (and
also evidently even a part of the adult males) remain beyond the range
of stationary shore ice in nearby sections of broken-up and drifting ice
floes. The arctic ringed seal remains even much later, in the period of
molt, in air holes mainly on the same coastal stationary ice floes that are
breaking up with time and warmth. In addition to pups of the current
year, young animals also gather here and after lactation and shedding of
the embryonic pelage depart from the shore ice. During this period the
ringed seal greatly enjoys resting on the ice floes preserved for a long time
along the highly rugged coasts and in the straits between islands. Such,
for example, are the southern coasts of Novaya Zemlya, the coastline
of the Bering Strait, and many other parts of the range. Nevertheless,
ringed seals do not avoid even the shallow sections with a fairly even
coastline, such as the Yamalsk shallow-water zone in particular or the

238

northern coastal strip of the Chukchi Peninsula. Naturally, under such
conditions the ringed seal colonizes far away from the coast, beyond the
limits of compact ice masses.

In the Sea of Okhotsk, however, during molt the ringed seals rest
once again on small, preferably isolated ice floes. At this time they evince
no interest whatsoever in site selection and can be seen resting on clean
or soiled, hummocky or smooth ice floes; sometimes they are seen even
on top of a hummock (Pikharev, 1941).

Beach rookeries are not very typical of this species, especially in the
arctic zones and in the Far East.

The ringed seal does not rest often in summer or autumn on the
coast in the western parts of the Soviet arctic and subarctic sections of
the range. Its rookeries are known on dried-up spits in the White Sea
Inlet (e.g., on Lidtke Spits, etc.), in the estuarine zone of the Eastern
Kambal’nitsa (along the northeastern side of the Kanin), and in the
estuaries of some Murman rivers, including Voron’ei. Further, ringed
seals rest on rock ridges, “ludas,” and “kirevyadi” at several places on
the Ladoga coast, and at places on the coasts of the Gulf of Finland and
Gulf of Riga in the Baltic Sea. At places they enter the rivers but do not
usually ascend far upstream.

181

Fig. 114. Immature ringed seal, Phoca hispida ochotensis, Bering Sea, June, 1964
(photograph by G.M. Kosygin).

183

239

Food. The ringed seal feeds on fish and crustaceans; other animal
groups (mollusks, worms, cephalopods, etc.) are consumed only very
rarely and are not at all characteristic of its diet. It feeds mainly in
the upper water layers on animals that are available en masse. Benthic
food is resorted to only in shallow places.°° The quantity-wise ratio of
fish and invertebrates consumed varies in different seasons according to
the periodic concentration of a given food type. In the autumn-winter
months the importance of fish increases noticeably; in some regions, e.g.,
Novaya Zemlya, with the en masse arrival of the polar cod, fish food can
be highly predominant and be even the mainstay in its diet.

In the spring-summer months, contrarily, the ringed seal feeds mainly
(sometimes at places even exclusively) on the various forms of crus-
taceans abundantly available. Thus in the Kara and Barents seas ringed
seals were caught with their stomach packed with either amphipods
(Themisto sp.) or mysids (Mysis oculata).

Like other seals, the ringed seal feeds sometimes more intensively
and sometimes less so in different seasons. The main season of its feeding |
covers the middle of summer, the entire autumn, and early winter. In
spring, 1.е., at the time of mating, feeding is evidently poor. It is less
intensive in the protracted period of molt because of the long residence
of the animals on ice floes. Nevertheless, there is no prolonged total
abstinence from food by this species.

In the Sea of Okhotsk, during spring (from February through June)
the ringed seal feeds mainly on the black-eye euphausid (Thysanoessa
raschii) while the amphipods (mainly genera Gammarus, Themisto, and
Anonyx), Shrimps (genera Pasiphaea, Pandalus, and Spirontocaris), mysids
(Mysis), and partly sea slaters (Mesidothea and Idothea) play a lesser
though significant role. Fishes such as pollock, pond smelt, navaga, and
more rarely herring (S. Naumov, 1941) are consumed in these months
in a comparatively small quantity (roughly at 10% by weight) (Fedoseev,
1965e) and represent over 35% of the number of species identified
(Pikharev, 1946).

In summer, at the end of molt, the ringed seal feeds intensively far
away from the coasts, evidently mainly on the planktonic organisms and
primarily on the black-eye, which clearly predominates; it also thrives
apparently on the schools of capelin which are attracted to the black-eye
and other planktonic species (Fedoseev, 1965e).

56 The depth of submergence can sometimes reach 60 m or more based on the fact that
the food of the ringed seal often includes the common sand eel (Ammodytes hexapterus)
which prefers to settle and spawn at such depths.

240

In autumn and early winter the ringed seal feeds more on fishes
(navaga, smelt, small herring, sometimes goby, sand eel, etc.). Since it
quite often transgresses even into rivers, especially in Kamchatka, the
ringed seal is assumed to consume salmon also (Lun’, 1936; Freiman,
1936). Although precise data are not available on this subject, the alleged
consumption of salmon by the Okhotsk ringed seal is erroneous. From
among the crustaceans, it consumes at this time predominantly shrimps
and amphipods, fewer euphausids, and least of all, mysids (Fedoseev,
1965e).

The food of the ringed seal in the Bering Sea has not been investi-
gated fully. In the transition period of spring and summer it feeds almost
equally on fishes (mostly the polar cod) and crustaceans (shrimps includ-
ing amphipods, and more rarely on mysids and black-eye). At this time
on the USSR coasts of the Chukchi Sea the ringed seal feeds more on
amphipods and shrimps and less on fishes (mainly on navaga, and some-
times flounder) (Fedoseev, 1965c). In winter, however, the ringed seal in
the Chukchi Sea survives almost exclusively on fishes, mainly the polar
cod (Johnson, Fiscus et al., 1966).

In the western seas of the Soviet arctic (the Barents, including
Pechora, and the Kara) the ringed seal consumes the very same two
groups of abundantly available foods: fishes and crustaceans. Polar cod
plays an exceptional role among the former and serves as the main food
in the autumn-winter period when huge schools of this small fish arrive at
the coastal regions to spawn. Seasonal concentrations of polar cod attract
the arrival and at places significant concentrations of the ringed seal in
the autumn-winter period on the coasts not only of Novaya Zemlya and
Vaigach, but also Timansk tundra, Cheshsk Bay, White and Kara seas.
In this manner, but on a smaller scale, concentrations of the ringed seal
are also stimulated by other schools of fish, e.g., the arrival of navaga
in the Pechora Sea and other regions. At the beginning of this century,
herring was abundantly found in the stomach of ringed seals caught on
the Murman coasts (Soldatov, 1902) when this fish was available there
en masse; the ringed seal also consumed capelin on these coasts. It was
assumed that the ringed seal noticed in the river estuaries of the Murman
coasts and transgressing fairly long distances upstream could even thrive
on salmon (N. Smirnov, 1903) but not accurate data are available (see
p. 257). In the Gulf of Ob ringed seals falling in fish traps ate omul
but this fish was not found in the stomach of seals caught in the open.
Young char were sometimes found in the stomach of the Novaya Zemlya
ringed seal. In fact, in the spring of 1904, the remains of a salmon were
found in a salmon net (Breitfus, 1908). There is now hardly any doubt
that the real culprit is the common seal. Yet the view that the ringed

184

241

seal preys on salmon persists here and there. This is evidently promoted
by the fact that the ringed seal can actually live at some places on the
very large fishes caught in nets; otherwise it could not have survived.
Whatever the reason, it is still firmly believed that “the whitefish plays a
significant role in the food of the ringed seal” (Kirpichnikov, 1937) and
the small population of the ringed seal in the Yenisey-Pyasinsk region
of the Kara Sea is castigated for the poor arrival of the omul (ibid.), or
that the ringed seal in the western seas of the Soviet arctic consumes
salmon and omul along with other fishes (Golenchenko, 1961).

In addition to fish, even in the west these seals consume amphipods
(mainly Euthemisto sp., Gammarus sp., and Gammarocanthus loricatus
and Anonyx пирах), a mysid (Mysis oculata), а euphausid—black-eye
(Thysanoessa inermis), and shrimps (Eualus gaimardi).

The food of the Baltic ringed seal has scarcely been studied in our
territorial waters. Presumably this seal feeds on fishes and crustaceans on
the Estonian coasts too but crustaceans play a minor role and are con-
sumed mainly in the winter and spring; Baltic herring and sprat take first
place among fishes; other fishes consumed include cod, eel, omul, and
even pike-perch (Aul, Ling, and Paaver, 1957). According to other data,
the Baltic ringed seal consumes mainly sprat and also goby, snailfish,
and amphipods; in the spring, however, it feeds mainly on the sea slater
(Schubart, 1929). Recent investigations showed that, in the Baltic Sea,
the ringed seal feeds mainly on the slater, goby, eelpout, small Baltic
herring, smelt, sprat, and stickleback (V.A. Zheglov). Large fishes are
evidently inaccessible to the ringed seal and it can only steal them from
fishing nets.

In Lake Ladoga the ringed seal feeds for the most part on smelt, ruff,
and small vendace. The stomach of this seal also contained stickleback
and small crustaceans (A.S. Sokolov, 1958; S.M. Sorokin). The view that
the Ladoga ringed seal “feeds on whitefish, char, vendace, and other
fishes” (A. Smirnov, 1961*), here too is evidently based on the instances
of this seal stealing these fishes from nets.

Home range. Although ringed seals are usually regarded as settlers,
their population is not confined to any one section of the sea year
round. They migrate (see “Seasonal Migrations and Transgressions”)
with changing ice, food, and other conditions. It is therefore practically
impossible to establish the definite boundaries and sizes of expanses
occupied by one single animal or even the population as a whole.

The mothers exhibit the most distinct stationary distribution
(excluding the Okhotsk ringed seals) and that, too, in the winter-spring
period when the gestating and whelped female is confined to a single
site selected by her for its suitability for pups to be born or already

184

185

242

Fig. 115. Ringed seal on а sandy shoal. White Sea, Severnye Koshki, September
20, 1970 (photograph by A.G. Beloborodov).

birthed. Such animals are scattered several hundred meters apart. The
density of their disposition depends on the size of the population, extent
of ruggedness of the coastline and the nature of the coast in general,
the depth of the coastal zone, ice conditions, and so on. Juveniles of
both sexes and even adult males are confined beyond the limits of the
stationary shore ice, along its edge, and in the zone of broken drifting ice
floes with open water pools. The degree of their concentration is much
greater but yet variable under the influence of the above factors, mainly
food conditions.

Hideouts and shelters. Ringed seals, like many other true seals, make
air holes in the ice floe through which they crawl out of the water for
respiration. The mechanism of making such holes is not known with
certainty but it may be assumed that initially, when the crust of the ice
floe is not yet very thick, the animal pierces it with its head by diving in
the same manner as does the Greenland seal. The Baltic ringed seal can
pierce an ice crust up to 2-2.5 cm thick in this manner (V.A. Zheglov).
Subsequently, these holes are kept open by repeated use for respira-
tion and later for crawling out of the water when the ice is thick. It is
possible that the walls of the hole are worked up with the claws also

186

243

(М.А. Smirnov, 1927; Kirpichnikov, 1937); in any case, the claws of the
ringed seal are powerful enough for this purpose.

The ice around the air hole builds up gradually and, when it is
used exclusively for respiration, a semispherical arch is formed above
the opening over the course of time, in which only the head or even only
the tip of the snout protrudes (and hence the hole is not large). The arch
is formed by snow built up from inside as a result of alternate thawing
by the exhaled warm air and possibly the water spray freezing during the
intervals in respiration. At the center of the arch, i.e., the dome, there is
usually a small hole but it can altogether be concealed under the snow
and wholly imperceptible from outside.°’ The coastal people call such
“air holes”.

There are many variations of the above-described typical air hole: it
can be hidden in the hummocks, under a hanging ice floe, pressed out
as a result of compression on the surface, in a niche formed under a pile
of ice floes, in a fresh fissure, and so on. It is not quite clear whether
the animal works on such a conical opening and, if so, in what manner.
Probably, from time to time, the ringed seal has to enlarge it, working
on it not only with the claws but also with its teeth.

Such openings quite often serve two functions: for respiration and
egress from the water. In such cases they are considerably wider and less
sharply narrowed upward, with a diameter of 30-40 cm or more on the
surface of the ice floe. They are made initially like air holes and then,
when the ice is quite firm to support the animal, are used for crawling
onto the ice repeatedly. Due to constant use, such holes are not sealed
by the ice nor are they highly frozen from inside.

Snowdrifts and the location of air holes promote this process.
Although the holes are made under extremely diverse conditions, they
are mainly concealed. When the hole is made by a gestating female, she
often selects a section of the ice cover damaged by a fissure followed by
compression, causing further opening up of the ice floe and the piling
up of broken lumps. In such places there are open water pools which
are rapidly closed by young ice along with randomly disposed ice blocks
with niches and voids between them. Sometimes the ice floes invert and
pile up one over the other in such a way that they form a typical roof
with a pointed peak rising above the edge of the young ice floe. Ringed
seals are particularly drawn to such natural hideouts for whelping. ,

Quite frequently, the ringed seal gives birth to pups in special lairs or
holes made in snowdrifts among hummocks, around an ice floe elevated
during compression, and at the base of drifting ice floes. Such holes are

57 Air holes can also be concealed under a snow cover as a result of drifting snow.

244

also encountered directly in deep snow at almost a level surface. Such а
lair can be detected only with the help of a dog. No one has ever seen
how a ringed seal constructs its “nest”. It is perhaps not very difficult
in loose snow. It probably makes a hole in the snow, partly using all its
physical might and partly by shaping it with its fore-flippers. Moreover,
the heat radiated by the animal’s body and its respiration somewhat
thaw the snow and it evaporates. Further, the arch from inside the lair
is sometimes lined with snow, which increases its strength. The roof of
many such snow lairs found in southwestern Baffin Land could only be
broken by applying force and after repeated trampling (McLaren, 1958).

The snow lairs made in the Kara Sea are usually longitudinal, about
170-185 cm long and 60-70 cm in height; they extend sideways from
an air hole whose diameter varies from 34-45 cm. Sometimes there is
a passage in the snow from the air hole into the snow chamber. The
same animal can usually make more than one air hole which is used by
other animals also. Only a single lair is usually built although according
to some workers (A.N. Tyulin), there can be spare ones. Judging from
all the information, mainly, if not exclusively, adult females make lairs,
and essentially for protecting their pups.

Among the Chukchi ringed seals, the snow lairs are much larger
in size with an area of 3 to 7.5 m? and 2-3 air holes. In winter such
“caves” can afford protection to several animals. Individual stray air holes
(without lairs), opening outward, are disposed quite close to the lair. The
upper diameter of the ice holes opening into the lair varies from 80 to
120 cm while the thickness of the ice in which it is made can be 95 to
110 cm and the height of the snow cover 85 to 125 cm (Fedoseev, 1965c).

Daily activity and behavior. Precise data on this aspect are not avail-
able. Visual observations in different regions, different seasons, and in
different environments provide no categorical answer. In the summer-
autumn season the ringed seals regularly remain in water, feed there,
move from place to place, rest, and sleep at any time of the day. In
spring and early summer they can be seen resting by the side of an air
hole or pool and sleeping much of the time for days on end, especially
in good weather.

Judging from the fact that ringed seals are trapped more often at
night in the nets set for them (and even in fishing gear), it can be con-
cluded that these animals are also active at night. Evidently even in the
season of intensive feeding, they sleep several times during the day, 1.е.,
after every full meal.

The more active spring behavior among the Okhotsk ringed seals
disposed on broken drifting ice floes is somewhat different. “Rookeries

187

245

are always buzzing with activity, with the seals diving into the water and
crawling onto the ice time and again” (Fedoseev, 1965f).

Seasonal migrations and transgressions. These aspects are relatively
less manifest among ringed seals and have not yet been fully studied.
These are largely passive activities, resulting from seasonal movements of
the ice floes with which this seal is associated almost from the beginning
of the arctic summer.

In the Sea of Okhotsk, corresponding to the predominantly anti-
clockwise flows, the bulk of the Okhotsk ringed seals move in the spring
from the northeastern regions of the sea (from the Penzhinsk, Gizhi-
ginsk bays and Shelikhov Gulf) to the southwest (see Fig. 112). This
was hinted at even in the early 1930s (Razumovskii, 1933). There were
contrary opinions too (S. Naumov, 1941). From the end of March when
the whelping season commences and the animals spend much of their
time on the ice, their maximum concentrations are seen in the region of
Tauisk Bay, on the threshold of Shelikhov Gulf, and in its southwestern
part. By May, the animals extend to the expanse in the Khanyangda-
Okhotsk zone; by June, they move into the Ayan region in the western
part of the Shantar Sea; still later, from June end through July end, they
are concentrated in the southwest from Shantarsk Bay to Sakhalin Gulf
and in the regions east of Sakhalin. Such is the usual passive drift of
the lactating and molting ringed seals and those simply “enjoying a sun
bath” under the influence of snowdrifts (Nikulin, 1937; Pikharev, 1940*;
Tikhomirov, 1961; Fedoseev, 1965f; and others).

Some basis for the possible active migration of the Okhotsk ringed
seal was reported very recently. While observing the concentrations of
these seals in the western regions of the Sea of Okhotsk from a plane,
it was noted that, in spite of powerful, stormy western and northwestern
winds driving the ice floes toward Kamchatka, the Okhotsk ringed seals
did not move with them; they were sighted a few days later en masse in
the northern and western parts of the sea (G.A. Fedoseev).

At the end of the season, the ice floes become very thin and the
ringed seals abandon them and begin to feed voraciously. Their move-
ments now are associated with the migrations of schools of fish and
with the distribution of planktonic and nektonic crustaceans. With time,
the number of ringed seals in the western regions of the sea decreases
noticeably and, concomitantly, massive and active movements of herds
of ringed seals are noticed in the water (Freiman, 1936) in a reverse
direction; 1.е., to the northeast. The population of the ringed seal again
increases sharply from September in the extensive coastal expanse in the
region from Tauisk to Shelikhov bays where the bulk of the Okhotsk
ringed seals spend the winter. However, such a large cycle of migrations

188

246

does not evidently occur invariably (Fedoseev and Yablokov, 1965); even
in the years when migrations assume large dimensions, a part of the pop-
ulation remains there leading a semisettled, semiwandering way of life
all along the coastal belt from Penzhinsk Bay to Sakhalin. In any case,
in winter and spring the Okhotsk ringed seal leads a settled mode of life
(Fedoseev, 1971).

Migrations of one type ог the other are typical of other populations
too. Most of the ringed seals abandon the Bering Sea and even more
so the Chukchi Sea with the spring-summer break-up of the coastal
ice floes and their drifting. Deep in autumn, they return again to the
northern coasts of the Chukchi with the appearance of fresh ice floes
(Arsen’ev, 1935). In the Bering Sea the rest of the population is scat-
tered on the beaches; many animals transgress into the lower courses of
the rivers, including the Anadyr, and sometimes ascend to a considerable
distance (Gondatti, 1897; Portenko, 1941). Sometimes the ringed seals
are unsuccessful in entering the sea before the rivers freeze and, in such
cases (evidently tragic), once again seek their way to the sea on ice floes
and even on land. Some cases of simply unbelievable finds of ringed
seals in hilly and wooded locations, where they strayed while looking
for the sea, have been reported (Portenko, 1941; Ostroumov, 1960; and
others). Similar cases have been reported in the west too (Vrublevskii,
1959; Chentsov, 1959; Vishnyakov, 1961; and others).

It is quite possible that the ringed seals from the southern coasts of
the Chukchi move north through the Bering Strait in summer.

The ringed seals of the western seas of the Soviet arctic perform
fairly regular migrations. Novaya Zemlya hunters distinguished even in
the 1930s the “arriving” and local (“well-settled”) ringed seals. With the
onset of summer and the thawing of ice floes, the former abandon the
coastal strip on the southern part of Novaya Zemlya and migrate through
the Kara Strait into the Kara Sea; with the onset of autumn, they again
return by the same route. The autumn-winter arrivals of ringed seals
on the coasts of Novaya Zemlya, Timansk tundra of Kanin Peninsula,
Kolguev and Vaigach islands, and other regions of the Pechora Sea and
into the White Sea in the north as well as the Kara Sea, are associated
mainly with the en masse arrivals of spawning polar cods and at places
with the similar arrivals of smelt and navaga.

The Baltic ringed seals also perform local migrations. Thus the pop-
ulation reproducing in Riga Bay arrives there in autumn and, with the
thawing of ice floes in spring, goes elsewhere into the open sea for the
summer (Leis, 1960).

Some periodic variations are noticed even in the distribution of the
Ladoga ringed seal. From time to time it gathers in one or the other

247

part of the lake, these movements sometimes being associated with the
migrations of vendace or even whitefish and char (A. Smirnov, 1954).

Reproduction. The mating period among ringed seals, as among other
pagophilic seals, falls in the spring and sets in evidently at the end (or
in the second half) of the lactating period.°* In the absence of direct
observations, the mating season has to be judged from the state of the
gonads and other indirect features. Empty seminiferous tubules of large
diameter at the end of May and, contrarily, active spermatogenesis from
early February through early May, and some other data suggest that the
mating period of ringed seals in the Barents and Kara seas extends from
April end to at least the first 20 days of May (Chapskii, 1940). The intense
growth of follicles commences immediately after whelping, simultaneous
with the resorption of the corpus luteum of the preceding gestation. An
increase in.the total weight of the ovaries is also associated with this
feature. Similar data suggest that the mating period among the Okhotsk
ringed seals too falls in nearly the same period, i.e., in the second half of
April to the first half of May (Fedoseev, 1964a, b, 1965b; Tikhomirov,
1966d*). The mating of the Bering ringed seals evidently concludes in
May (sperm were not detected in the epididymis of the males in June;
Fedoseev, 1965c). According to other information (Razumovskii, 1933),
the Chukchi ringed seal presumably mates in March and April.

In the Baltics, corresponding to the prolonged whelping, mating
occurs mainly in April but could occur even in March, ceasing by April
end (Aul, Ling, and Paaver, 1957; Leis, 1960).

The mating season among the arctic ringed seals evidently proceeds
without serious conflicts between the males; this is due to the scattering
of the animals which do not form herds either in early spring, at its end or
in early summer. No traces of injury whatsoever, that could be attributed
to fights between competitors, have been noticed on the skin of males.
Nevertheless, due to the limited population of males and reproducing
females, fights do arise at some places. For example, in Riga Bay the
male resists any intruder attempting to approach a chosen female by
grasping his flipper (Leis, 1960).

Males capable of reproducing are usually not younger than six years
and most of them become productive for the first time only at seven
years of age.

Such a late maturation was reported among the eastern Canadian
and Far Eastern populations (McLaren, 1958; Fedoseev, 1965b;

58 An earlier view that the Okhotsk ringed seal mates in the second half of July and in
August after molting (Sleptsov, 1943) is based on an error arising from ignoring the lag in
the implantation of the blastocyst.

248

Tikhomirov, 19664*). It has been assumed that about 20% of the males
of the Okhotsk ringed seal mate for the first time even in the 8th year
(Fedoseev, 1964).

Maturation among females sets in somewhat earlier. They undergo
parturition at the earliest in the 5th year according to some authors
(Fedoseev, 1964a, b; Tikhomirov, 1966) and according to others
(McLaren, 1958), in the 6th year. However, the percentage of females
maturing early is not high. Quite a few females give birth in the 6th or
7th year. A small number produce the first pup in the 8th year. The

189 youngest mothers among the Barents ringed seals are 6 years old but

189

their number, about 3%, is very small (Nazarenko, 1965).

Embryonic growth is impeded right at the initial stage of fission
of the fertilized egg cell [delayed implantation]. The duration of this
period among the Barents and Kara ringed seals was originally regarded
to be about two months (Chapskii, 1940). It later became clear that the
implantation of the blastocyst among the Canadian ringed seals occurs
roughly 3.5 months after fertilization (McLaren, 1958). The duration
of the latent period among the Okhotsk ringed seals is 2.5-3 months
(Fedoseev, 1965b).

Fig. 116. Ringed seal on ice floes. Barents Sea, April, 1967 (photograph by
V.A. Potelov).

190

249

Even in the period of peak reproductive activity, not all ringed seals
undergo parturition every year. In the Canadian population the average
percentage of barren females is 5-10 (McLaren, 1958, 1960) although
according to recent calculations, it was assumed that only one-half of
the eligible population gave birth to pups (McLaren, 1967*). A fairly
significant percentage of barren animals was reported among the Far
Eastern ringed seals (20% on average among the Okhotsk ringed seals;
Fedoseev, 1964c*)°? and even more among the western arctic ringed
seals, of which 37.5% of the adults were without pups (Nazarenko, 1965);
V.A. Potelov puts this figure at as high as 50%.

Early spring is the period of whelping but it is quite protracted even
in a given region in the USSR part of the range as a whole, extending
for at least two months. In the Baltic basin, on the Estonian coasts and
more northwards, the ringed seals whelp mainly at February end to early
(more rarely end of) March (Lonnberg, 1899; Nordquist, 1899; Schubart,
1929; Freund, 1933; Ropelewski, 1952; Aul, Ling, and Paaver, 1957), and
in Lake Ladoga in early March.

In the White Sea, on Murman, southeastern parts of the Barents
Sea (close to the Novaya Zemlya coasts, in the southeastern parts of the
Pechora Sea, in Cheshsk Bay, close to Timansk coast, and in Yugorsk
Shar), the ringed seal whelps near about the same time, mainly from the
middle or end of March through the middle of April, and sometimes even
later (М. Smirnov, 1903; Knipovich, 1907; Ognev, 1935; А.М. Dubrovskii;
М.1. Vladimirskaya; Yu.I. Nazarenko; У.А. Potelov; and others). In the
Kara Sea this seal whelps evidently somewhat later, mainly in April,
though the data on this are contradictory and limited (Kirpichnikov,
1937; Kovalev, 1938; Tyulin, 1938).

In the northern part of the Sea of Okhotsk, in the region of Tauisk
Bay, pups arrive from March to mid-April (Tikhomirov, 1961), mainly in
the second half of March and the first half of April (Fedoseev, 1964b* ).
About the same time, ringed seals on the coasts of the Chukchi Penin-
sula also undergo parturition (Razumovskii, 1933; Fedoseev, 1965c). In
Tatar Strait, whelping extends from mid-February to probably mid-May
(Dorofeev, 1936; S. Naumov, 1941; Yu.A. Salmin).

_ Growth, development, and molt. Unlike most other species of seals,
the initial postnatal period has not been adequately studied with the
exception of the Okhotsk population. This is explained by the fact that
birth, lactation, and shedding of the embryonal pelage are all concealed
in the snow cover from the first appearance of the newborn almost until

53 According to Tikhomirov (1970), however, barren seals in the Far East constituted
only about 7%.

191

190

250

it becomes self-supporting. Only in the Sea of Okhotsk and in Tatar
Strait are pups born in the open; but even under these conditions, the
pup tends to remain among the hummocked ice or the icy ledges to
protect itself from the wind. Having selected such a site, it does not
leave it for long without purpose; a troughlike depression even forms in
the ice floe due to the prolonged resting of the pup at the same place
(Pikharev, 1941).

The embryonic coat of the newborn is creamy-white with a faintly
discernible greenish or grayish tone in the first few days. In some pop-
ulations (especially in the case of the Ladoga ringed seals), it is quite
often covered with a smoky-brown bloom dorsally. The embryonic coat
is almost as long and dense as in the newborn of other proximate species
but is inferior in fineness, abundance, and luster (Fig. 117).

How long the pup sports the embryonic coat has yet to be accurately
established but evidently it is longer than ten days. Since the pups of the
Okhotsk ringed seal are seen with an embryonic coat mainly from the
second half of March through the first half of April (Fedoseev, 1964b* )
and at the end of the first five days of May have only just begun to molt
(Pikharev, 1941), the firm hair coat apparently endures for at least two
weeks.

Fig. 117. Head of a ringed seal pup that has not shed its embryonic hair coat.
Bering Sea, June, 1964 (photograph by G.M. Kosygin).

251

The duration of molting of pups in the course of their suckling
evidently conforms to a common pattern. Among the normally grown
animals, molt commences from the snout while the bases of the fore
and hind flippers and the tail evidently molt later; the embryonic coat
lasts longest on the body flanks and also on the belly. Even before the
weakening of the embryonic coat, the growth of the hair characteristic
of the subsequent stages commences.

The body length of the newborn varies from 55 to 65 cm; more often
about 60 cm (Le, i.e., along the body surface up to tip of tail); weight
varies from 3.5 to 4.0 kg.

Lactation continues for not less than a month, evidently until the
pup has completely molted. The mammary glands of the Okhotsk ringed
seal function until mid-May; until this period, the intestines of the pups
contain no food other than milk (Fedoseev, 1964b*). During the suckling
period pups of the Okhotsk ringed seal grow to 64-72 cm (average up to
67 cm) and almost double their initial weight (raised to 6-9 kg, average
up to 7.5 kg). In the last seven months, by November, they add another
10 cm (average) in length and roughly 4.5 kg in weight, thus reaching
72-84 cm in length (Lc) (average 78 cm) and 10- 14 kg in weight (average
12 kg) (Fedoseev, 1964b*). Similar figures have also been recorded for
the pups of the Bering ringed seal (Fedoseev, 1965c).

The autumn yearlings of the Barents ringed seal are larger, with a
body length (Lc) varying from 74 to 103 cm, average 91 cm (the data are
not sufficiently representative) (Chapskii, 1940).

Under unfavorable growth conditions, when the pups lose suckling
mothers or become ill for example, they remain underfed but never-
theless do not die; they do not grow to the size normal for their age
and remain dwarfs. Such animals are encountered time and again on ice
floes in the White and Barents seas. The coastal people call such starved
animals “kavadei” or “telesai” (Fig. 118).

Further growth of the young before the onset of maturation is low,
as can be seen from the following changes in average body length (Lc)
and average weight of the Okhotsk ringed seal:

Length (cm) Weight (kg)

One-year-olds 84 14
Two-year-olds 92 19
Three-year-olds 98 24
Four-year-olds 102 27
Five-year-olds 106 29.5
Six-year-olds 110 32

Seven-year-olds 113 34.5

252

192 Fig. 118. Underfed pup (“Кауаде!” or starved) of a ringed seal in a state of
delayed molt. White Sea, April 12, 1967 (photograph by V.A. Potelov).

Growth and weight continue to rise in subsequent years also but stabilize
from 10 years of age at an average length of 119-122 cm and weight of
40-44 kg (Fedoseev, 1964b*).

The period of molt among juveniles, commencing from yearlings,
and among adults is highly protracted. In the western arctic seas of the
Old World, molting individuals are encountered even in May but June
and July represent the more normal molting period in the Barents and
Kara seas; the animals lagging behind complete molting as late as August.
On the western side of Novaya Zemlya, molting individuals constitute
a scattered herd far removed from each other on the smooth ice floes
among the shore ice.

192 In the Sea of Okhotsk ringed seals molt on drifting ice floes in
very dense groups from April end through mid-July. The adult males
begin to molt two to three weeks earlier than the gestating females and
hence molting of the latter is correspondingly more extended. Unmolted
juveniles are encountered not only in August, but presumably even in
September (Fedoseev, 1965f), although molting in the majority of these
seals ceases in the first half of July under normal conditions (S. Naumov,
1941; Tikhomirov, 1961).

Enemies, diseases, parasites, mortality, and competitors. In the arctic
zone the polar bear continues to date to be the number one enemy of the

193

253

ringed seal, even through the predator’s population in the twentieth cen-
tury has sharply decreased everywhere, especially in the western regions,
including those along the coasts of the southern island of Novaya Zemlya
and also in most of the regions of the Kara Sea. This predator of the
ringed seal has been exterminated almost wholly in the southeastern part
of the Barents Sea, in the southwestern part of the Kara Sea, and in its
Yamal-Obsk-Yenisey region. At places where the polar bear still con-
tinues to exist, it has survived mainly on the ringed seal. The Greenland
shark (Somniosus microcephalus) is the second enemy but mainly in the
Barents Sea part of the range, though it could attack the Far Eastern
ringed seals too. The same is true of the killer whale (Wilke, 1954).
Among the other probable enemies are the walrus; the rapacity of this
animal has been reported time and again (Chapskii, 1936). There is a
reference to the disappearance of the ringed seal from Rudder Bay (and
from this region in general) with the arrival there of walruses (Fedoseev,
1965c). Although the walrus does attack the ringed seal here and there,
it cannot be held responsible for any significant loss of seal reserves
since, firstly, the walrus has survived in the USSR only in the extreme
northeast and in comparatively small numbers in the Laptev Sea; in the
western arctic seas, however, it is almost absent. Secondly, the ringed
seal swims rather faster than the walrus and hence becomes its victim
only occasionally.

From among the land vertebrates, the fox is one of the direct ene-
mies of the ringed seal at places. Its tracks leading to the lairs of ringed
seals were encountered on the coasts of Alaska (Bailey and Hendee,
1926); such instances are reported sometimes even in the Ladoga region
where the ringed seal has its burrow close to the coast (S.M. Sorokin).
Sometimes the wolf, too, embarks on a hunt for the pups of the Ladoga
ringed seals (Andreev, 1875*; Bergman, 1956; S.M. Sorokin). In our
Baltic waters large predaceous birds, especially the sea eagle, attack the
young seals (Aul, Ling, and Paaver, 1957). Occasionally, ringed seals are
attacked by land carnivores (at times by man too) when for some rea-
son, although very rarely, the animals are compelled to traverse ice or
snowbound land for a long time in search of open waters (see p. 246).

From among the ectoparasites of the different subspecies of the
ringed seal, only one species of seal lice, Echinophthirius horridus, has
been detected.

The helminth fauna of the ringed seal® inhabiting the USSR waters
has been studied better than in any other zone of its range (Mozgovoi,

6 From the same sources (see p. 208).

254

1953; Delyamure, 1955; А. Skryabin, 1959; Delyamure and Alekseev,
1965; Delyamure, Zavaleeva, and Fedoseev, 1965; and others).

A study of the material (Treshchev, Potelov, and Zavaleeva,
1967; Treshchev and Serdyukov, 1965; and N.V. Yurakhno) from the
ringed seals of the White, Barents, Chukchi, Bering, and Okhotsk
seas eastablished 25 species and 5 larval forms of helminths in
this animal. Among the trematodes, Orthosplanchnus arcticus infects
the liver, gall bladder, and the pancreas, and is encountered more
often than any other species of trematodes; the liver of one 18-year-
old male from the Bering Sea revealed over a thousand specimens
of this trematode (N.V. Yurakhno). Pseudamphistomum truncatum
parasitizes the liver while Phocitrema fusiforme inhabits the intestine.
The following cestodes were detected in the intestine: Tetrabothrium
sp., Anophryocephalus anophrys, Trigonocotyle skrjabini, Trigonocotyle sp.,
Diphyllobothrium lanceolatum, D. hians, D. fasciatus, Diphyllobothrium
sp., Diplogonoporus tetrapterus, and Pyramicocephalus phocarum. The
nematodes infecting the stomach and intestine are Contracaecum
osculatum, Terranova decipiens, T. azarasi, and Phocascaris phocae; Ph.
netsiki and Anisakidae р. sp. infect only the intestine; Otostrongylus
circumlitus and Skrjabinaria spirocauda parasitize the heart, lungs,
and blood vessels while Parafilaroides arcticus and P. krascheninnikovi
parasitize only the lungs; Phocascaris phocae infects the ringed seal
more often than any other nematode. The following acanthocephalans
infect the intestine: Bolbosoma nipponicum, Corynosoma strumosum,
C. semerme, C. validum, C. hadveni, C. ventronudum, and Corynosoma sp.

A comparison of the helminth fauna of the subspecies Ph. (h.)
ochotensis and Ph. (h.) krascheninnikovi established that, in addition to
the common species of helminths, some are known to infect only one
of the subspecies. In particular, Phocitrema fusiforme, Diphyllobothrium
lanceolatum, Anisakis sp., Otostrongylus circumlitus, Corynosoma sp.,
and Bolbosoma nipponicum are not found in the Bering subspecies
(Ph. h. krascheninnikovi) while, on the other hand, Diplogonoporus
tetrapterus, Terranova sp., Skrjabinaria spirocauda, Parafilaroides arcticus,
P. kragcheninnikovi, Corynosoma hadveni, and C. ventronudum are not
found in the Okhotsk ringed seal.

The results of a study of 220 Bering ringed seals (N.V. Yurakhno)
revealed that helminths are found even among the yearlings, 90% among
one- and two-year-old animals, while 100% of the animals are infected
from the third year onward. Infection is particularly intense at 7 to 10
years of age. The colon (in 79.5% animals), rectum (70.4%), small intes-
tine (68.1%), rarely the stomach (9.1%), liver (9.1%), lungs (6.8%), and
heart (2.3%) are often infected.

194

255

The results of a study of 138 Okhotsk ringed seals (N.V. Yurakhno,
V.V. Treshchev, S.L. Delyamure, A.M. Serdyukov) revealed that
helminths attack animals of all ages, particularly and intensely those
in the first three years of age. The body parts more often infected are
the small intestine (66.6%) and colon (44.1%), and rarely the lungs
(10%) and stomach (9.1%). Apart from the other pathogenic species,
the nematode S. spirocauda, infecting the heart, blood vessels, and lungs,
deserves special attention. A large number of these nematodes infect
not only the adults, but also the one- and two-year-olds and cause severe
emaciation in them. Heavy infection probably leads to the death of ringed
seals (Delyamure, Zavaleeva, and Fedoseev, 1965).

There are undoubtedly other factors for the mortality of ringed seals:
diseases (not as yet understood), adverse birth conditions, and disrup-
tions of lactation, compression and hummocking of ice which can crush
not only the pups, but also older animals, washing away of the newborns
by waves or drowning of those on small ice floes in broad open water
pools. The total loss, including natural mortality and commercial killing
of various age groups, has been assessed only for the Okhotsk ringed seal
as follows (approximate): 40% of the pups are lost in the first year (3.3%
by commercial killing). In each of the subsequent 12- and 13-year-olds,
about 10-13% perish, in 14- and 15-year-olds roughly 30% each, in 16-
and 17-year-olds over 50% each, in 18- and 19-year-olds about 78%, in
20- to 25-year-olds over 90%, and in the older age groups wholly, 1.е.,
100% (Fedoseev, 1964c*, 1965f).

The ringed seal, in principle, has no serious competitors for food
from among the other seals. It is quite possible that in the autumn and
early winter months, the Greenland and bearded seals which thrive on
polar cod compete with the ringed seal at places in the Barents and White
seas. But the polar cod is so abundant that it leaves a large surplus after
meeting the requirements of not only these species, but also the white
whale and other animals. The common seal, 1.е., the larga, can hardly be
regarded as a serious competitor of the ringed seal in the Far Eastern
seas since the ringed seal consumes mainly the small varieties of fish
available in schools.

Population dynamics. This aspect has not been adequately studied
and some comments can only be made with regard to the Okhotsk ringed
seal and its intense hunting from the latter half of the 1950s and almost
throughout the 1960s. From the years of the postwar restoration of hunt-
ing using ships in the Sea of Okhotsk to the early 1950s, an average of
6,000 animals per annum was taken without disturbing the delicate bal-
ance of the herd. From the mid-1950s, the position underwent a sharp
change: the scales of hunting rose ten-fold to about 60,000 animals for

195

256

six years, touching a record of over 70,000 animals in 1960. This year
represents the turning point and the kill began to drop thereafter. The
smaller average number of animals caught per ship also served as proof
of the diminishing resources. While it was 6,500 in 1957, it had dropped
to 3,500 by 1963.

The unfavorable state of the population of the Okhotsk ringed seal
is also witnessed by the low average age of the animals caught (6.5 years)
and also the fall in the relative proportions of all generations aged
18 years or more to less than 1% (Fedoseev, 1966a). It should be remem-
bered that the total life span of the ringed seal, judging from the Cana-
dian populations (McLaren, 1958), exceeds 43 years. All of this viewed
together points to excessive killing of these seals.

The status of the ringed seal reserves in other regions of our range
suggests no such danger®! since hunting even at places where it is quite
stable does not threaten the herd with degradation.

Field characteristics. These are relatively small seals, not longer than
140 cm (Lcv), inhabiting the arctic seas, and also Lake Ladoga. They do
not form dense herds, remain on the shore ice in winter, and build snow
lairs and air holes in stationary ice floes. The skin usually has a predom-
inant pattern of fused light-colored rings with no dark-colored specks or
dabs. The snout is short (Fig. 119). The claws of the fore flippers are
relatively massive with a high dorsal longitudinal ridge. (K.Ch.)

Economic Importance

The economic importance of the ringed seal is quite significant and
diverse. Its skin at places even now is used in the local rural economy for
making nearly waterproof shoes (aviator boots, bags, slippers), mittens,
caps, and jackets. In the past the skins were cut into strips to meet var-
ious needs right up to weaving them into nets. Fox hunters caught the
ringed seal and even now use it as bait. It has immense importance as a
source of animal food and at places even as meat for human consump-
tion. At present, the skins are used for making superior and extremely
trendy furs in the natural state for caps, dress jackets, coats, and other
fur products. For this purpose animals with extremely short wool, i.e.,
growing or adult, are almost exclusively used.

The adverse economic implication of this seal as a consumer of com-
mercial fish is extremely insignificant since it feeds on crustaceans and
small fish which are of minor economic value (see above). Unfortunately,
however, the intake of salmon by the ringed seal has been exaggerated

61 Apart from Lake Ladoga in which the population is not high.

195

196

257

Fig. 119. Large adult Ladoga ringed seal, Phoca hispida ladogensis, on its back
(hunter’s catch).

or is even altogether baseless since this seal is usually not attracted to
salmon.

The main source of the ringed seal for fur and meat is the Sea of
Okhotsk. Here an average of 50,000 animals were caught annually in the
early 1960s (including catches by state ships and coastal collective and
state game farms). Hunting by the local population on the Chukchi coasts
occupied second place. From 1934 through 1940, the annual kill there
ranged from 19,000 to 37,000 animals (average 25,000). In the 1940s, it
varied from 14,000 to 29,000 (average 23,500), and in the 1950s, from
11,000 to 20,000 (average for the last five years of the 1950s, 15,500 per
annum) (P.G. Nikulin).

Accurate statistics of the catch of ringed seals are not available for
the White, Barents, and Kara seas. According to incomplete data, in
1933, these seas yielded a total of about 26,000 animals (nearly equal
numbers from the Kara and the other two seas taken together) while in
1962, the total (partial count) kill was only 17,000 animals. In the White
Sea alone, for the period 1928 - 1931, the incomplete figures of the kill
ranged from 4,800 to 7,400 animals (Yu.I. Nazarenko). In 1960-1967,
according to the approximate data based on average weight of skins with
blubber, the kill in the five hunting sections of the Cheshsk-Pechora -
region varied from 1,800 to 9,000, an average, of 5,000 ringed seals per
year (A.P. Golenchenko).

258

In the arctic seas of Siberia, hunting is not well developed; the ringed
seal is hardly caught in large numbers on the southwestern coasts of the
Kara Sea in Yamal, Yenisey Bay region, and at places in the east. In
the Kara Sea up to 2,000 ringed seals are caught annually to meet local
requirements (V.A. Potelov).

In the Lake Ladoga region, at the beginning of this century (from
1909 through 1918), 436 to 1,278 ringed seals were caught annually.
Hunting in subsequent years varied in a nearly similar range. Thus 677
animals were caught in 1928 and 1,262 in 1930 (Gottberg, 1927, 1930*;
Chapskii, 1932*). From 1941 right until now, no systematic hunting has
been carried out for the ringed seal. Small groups of hunters move on the
ice but the kill is extremely small. Animals are trapped in large numbers
in fishing nets or are caught by chance. On the whole, a few hundred
ringed seals at best have been caught in recent years.

The kill of the ringed seal on the southern coasts of the Gulfs of
Finland and Riga is slightly more; hunting here is undertaken by the
cooperatives of collective farms.

The hunting methods are quite diverse but essentially similar to the
methods used for catching other species of seals though there are some
local variations.

In the Sea of Okhotsk the vast majority of the ringed seals are caught
using boats dropped from hunting ships, which move among drifting
ice floes, mainly in the western regions of the sea. Brigades of hunters,
masked to some extent, approach the animals resting on the ice and
shoot them with rifles. The coastal collective and state farmers often
hunt for the ringed seal in this manner without a floating base, 1.е., а
hunting ship. At places, the hunters chase an animal sitting on the coast
and, after a successful shot, transport it quickly to shore in a light boat.

The winter-spring hunting on shore ice (in summer at places in the
arctic) is somewhat similar to the above method. The hunters go up to
the edge of the hard shore ice using dog or reindeer harnesses or simply
on skis, but when the shore ice is not wide enough, they reach there
on foot. In order to reach a killed seal on the shore ice, a light boat or
canoe is used or if the kill is close to hand, a harpoon. On the Chukchi
Peninsula typical tools are used for this purpose, such as a pear-shaped
stump lined peripherally with sharp hooks and tied to a branch or a
frame. The hunter casts this tool at the floating body, snags it, and draws
it toward himself.

In spring, when a ringed seal is resting on an ice floe near a hole, the
hunter, hidden in a concealing device, crawls toward it within shooting
range. He often uses various types of shields set up on a sledge or on skis.

197

259

In winter-spring, much before the ringed seal emerges onto the ice
floe surface, the hunters set out with dogs in various regions, most often
in the Baltics and in Lake Ladoga. The dogs help them locate the seal’s
air hole or lair with pups inside. Often, using the pups as bait, the hunters
attempt to catch the suckling mother.

There are many methods of trapping ringed seals using nets. Special
nets, with or without frames, often in the form of a sieve, are placed under
the ice floe near or under an air hole or in open water pools in the shore
ice. Trapping in nets (called in the western Soviet arctic “yundas” or
“kryuks”) is particularly widely adopted in autumn. The net, in the form

of a wall, is set up at an angle to the coast and is fixed at its farther end

with an additional crowd net with inward corners (or simply turned at the
end toward the shore and later bent once again toward the wall). Thus
a very good trap is formed; the seal encountering the net wall is snared
and cannot escape. In the White and Barents seas, and at places in the
Kara Sea, not only the ringed seal, but even the bearded and Greenland
seals, and sometimes even white whales get trapped in such nets.

The ringed seals of the western region of the Soviet arctic are
110-130 cm long and weigh 40-65 kg. They yield 20, or almost up to
30, kg of fat with the skin. The weight of such blubber varies in different
animals depending on their seasonal well-being, which is highest at the
beginning to the middle of winter. In the better fed animals the blubber
weighs 40 kg, more in exceptional cases. Among adults or near-adults,
the average weight of the “fat in the skin” goes up to 25-27 kg: the
average weight of the blubber among 130 animals caught in February
was 27 kg on the Timansk coast of the Pechora Sea (Moskalenko, 1945)
and 25 kg for the ringed seals of Cheshsk-Pechora region in the 1960s
(A.P. Golenchenko). The blubber accounts for 50% or more of the total
body weight. The skin of the ringed seal is now mainly valued as a raw
material for fur. Its unprocessed weight, without the flippers, for a young
animal 100-110 cm long (Lc) varies from 1.9 to 2.7 kg (average 2.3 kg)
and in adults about 3.0 kg. The eviscerated fleshy carcass weighs 10 to
15 kg in the case of a young seal (older than a year) to about 20 kg in the
case of an adult animal. The carcass without the viscera of the Pechora
ringed seals caught in winter weighs around 19 kg (Moskalenko, 1945).

Seal hunting should be developed in the following main directions:

1. Rationalization of hunting the Okhotsk ringed seal to obtain the
valuable fur and fat material in quantities that will not result in the dec-
imation of the animal resources. Simultaneously, ancillary products such
as the edible meat and so forth should be fully utilized (Tikhomirov,
1963).

198

260

2. Extending seal hunting into the White Sea and, in the western
regions of the Soviet arctic, into the Barents and Kara seas, and also
onto the Chukchi coasts and the eastern regions of the East Siberian
Sea.

3. Universal standardization of hunting norms everywhere and
prescribed hunting seasons (with hunting proscribed during the periods
of parturition, lactation, and molting).

4. Protecting the seal reserves from unproductive losses by banning
the shooting of the ringed seal in water in summer and autumn, encour-
aging the trapping of seals using nets, etc.

Extensive investigations on the ringed seal in all the important
regions of its habitat are essential. (K.Ch.)

CASPIAN SEAL
Phoca (Pusa) caspica Gmelin, 1788

1788. Phoca vitulina var. caspica. Gmelin. Systema Naturae. Ed. XIII,
1:64. Caspian Sea.

1929. Caspiopusa behningi. Dybowski. Bull. Intern. Acad. Polon. des Sc.
Cracov, 11, No. 8-10, p. 414. Caspian Sea (nom. nud.).

1929. Caspiopusa kisielewitschi. Dybowski. Ibid., р. 414. Caspian Sea
(nom. nud.).

1929. Caspiopusa dierzawini. Dybowski. Ibid., р. 414. Caspian Sea (nom.
nud.). (V.H.)

Diagnosis

The body length including the tail along a straight line (Lcv) does not
exceed 150 cm and along the dorsal side (Lc) up to 160 cm. The color is
mottled; dark spots are interspersed on a light-colored main background,
of which only narrow winding gaps remain at places (on the dorsal side).
The condylobasal length of the skull is not more than 190 mm and the
rostral part (from the anterior edge of the skull to the commencement
of the orbits) exceeds the length of the orbits (from the anterior edge
of the zygoma to the posterior wall). The tympanic bullae are small and
wide-set; the gap between them is more than their length, but not more
than 33 mm. The width of the bony lobe of the external auditory mea-
tus is less than the distance from its anterior edge to the crest of the
articular fossa. The anterior edge of the zygoma is usually considerably
wider than the posterior one and is forked. The molars and premolars
are small and set far apart; the accessory cusps on the lower jaw teeth,

261

commencing from the second, diverge from the main cusp. The greater
and lesser tubercles on the humerus (tuberculus majus and t. minus)
converge and fuse, forming a compact ring through which the tip of the
biceps muscle passes. (K.Ch.)

Description

The general build and the relative dimensions of the flippers are essen-
tially similar to the corresponding features among the closely related
species, 1.е., ringed and Baikal seals. The first two digits (or only the first)
of the fore flipper are longer than the remaining ones and become shorter
toward the fifth. The claws on them are well developed but not broad,
with a rounded dorsal ridge (rib), and not greatly elevated. The neck is
perceptible and does not narrow. The head is small, with a rostrum that
is extended somewhat more than in the other species of the subgenus.
The whiskers are flattened, with wavy edges. The labial whiskers are gen-
erally disposed in eight rows. In the first and the fifth row (counting from
below), there are more often nine whiskers each; from the second to the
fourth 11 to 13, and from the sixth to the eighth six (rarely seven) to
1-2. The total number frequently is 66-67 (on each side). The supraor-
bital whiskers number seven each and those around the nostrils one each
(Yablokov and Klevezal’, 1964).

The color of the hair coat varies, depending on age and sex, and also.
from animal to animal. Most adults are characterized by fairly intense
mottling, especially on the dorsal side of the body. Because of this, the
coat is considerably lighter on the belly side, more so in females. The
haphazard dark gray, brown, dark blue, and often nearly black spots of
various sizes and shapes, sometimes isolated and sometimes with fused
edges, sometimes overlapping each other, form diverse fanciful patterns
on the back of the neck, on the shoulders, and on the back. In many
cases the spots are so profuse that only narrow and winding light-colored
streaks remain of the main background, sometimes resembling longitu-
dinal half-closed cz closed ringlets. Spots are perceptibly rarer on the
flanks (see color Plate II).

The spots on the belly are usually smaller and paler, soft brownish-
gray, and their outlines less sharp and indistinct, as a result of which the
underside of such animals is perceptibly lighter in color than the upper
side. In most adult males, even the belly side of the body is variegated
with contrasting dark brown fanciful spots. The flanks usually sport a
‘ transitory type of coloring; the dark color gradually turns lighter and
descending from the back fuses with the whitish coloration of the belly
running from the opposite side.

262

The fore flippers are sometimes more, sometimes less dark gray on
the outside with minute dark gray (almost black among adult males)
spots and dabs; their underside (lower side) is the color of the belly,

199 usually without spots. The outer side of the hind flippers is steel-gray,
usually the same as the back or even darker, sometimes with faint minute
spots; their inner sides, turned toward each other, are light gray. The tail
on the upper side is dark gray, blackish, usually with a light-colored
fringe along the sides; the underside is ashy.

Although color differences between the hair coats of males and
females is disturbed by individual variations, it is nevertheless quite
noticeable (Figs. 120 and 121). The females have very dull tones and are
similar in this respect to the immature specimens of both sexes. The dor-
sal side in most of the animals is a dark gray, with an olive shade, usually
with an indistinct, slightly contrasting, spotted pattern, but is sometimes
quite mottled. The ventral side, however, is very light-colored, grayish,
with a few rather small pale brown spots or altogether without them.
The spots on the fore flippers of females are either very few or absent.

S we ЗЕ:
За:

Saas

Be ZEEE ae TE
277

ЕЕ: => >= пы
22 Е igs ==) й et
Me EL EEE LIFE, 9, OX Ra
И a =
77 LE gat Z >
Zz fgg he
„РРР, Я

iy. р ar
bi SO th 7, т
Я РИ,

199 Fig. 120. Caspian seal, Phoca caspica. A—adult male (with variegated colors);
B—adult female (mother) with pup (figure by N.N. Kondakov).

‘(Aoyepuoy ‘мм Аа soinsiy)
эпиэлп! „рэзпоиз-рэл,, & $1 148 dy) UO ‘зэпиэлп! рэло[оэ-Улер OM] эле э[еш5] э41 pulyoY ‘(sajtuaanf ‘sayetway pue sayew) зрециие sunod
‘punossyorq ay} ш ‘(1эцзош) эешэ} паре ие и puryag ‘(,, JAS]UOY,,) эеш ро ие $1 puUNOIBIIOJ эц1 и] “WH poidsno (psnq) vo0yd еэ$ ие $)

‘TI 21а

foro:
Bee

200

200

263

Fig. 121. Juvenile Caspian seal, Phoca caspica (figure by М.М. Kondakov).

The males have a more complex and contrasting pattern on the dor-
sal side (which is why they are called “animals with variegated coloring’’);
further, the spots are more often of diverse colors, from brown to almost
black. The flanks and the ventral side are also mottled with spots of var-
ious sizes; small numbers of them usually fuse together though not as
densely as on the back.

The individual color variation in all age groups of both sexes is also
significant. It is difficult to find two skins, especially of adult males or
females, which are identical in all the design features. The form, size,
number, and even the color of the spots on the ventral and dorsal sides
of the body vary in the males; the color of the ventral side is particularly
variable among the females. Light-colored streaks on the upper side of
the body (when they are present), speckles on the flippers, and also the
spotted pattern on the occiput and neck vary extremely in both sexes.
Insofar as seasonal color variation is concerned, the color after molt
turns brighter and is more lustrous. After summer the color becomes
somewhat dull and the spots lose their sharpness.

The skull (Fig. 123) is more elongated and relatively narrower than
that of the ringed seal, with a small and rather narrow cranium whose
width above the mastoid processes is nearly equal to its length measured
up to the orbits, but more often somewhat less. The zygomatic arches
are moderately separated, hardly exceeding the width at the mastoid pro-
cesses and forming not more than 110% of the latter in adults. The width
of the skull at the level of the upper canines in adults is 27-29% of the

264

201 Fig. 122. Fetus of a Caspian seal at the end of embryonal growth (figure by
N.N. Kondakov).

LE NACE
AN, “WX

4,
4,

201 Fig. 123. Skull of Caspian seal, Phoca caspica (figure by N.N. Kondakov).

mastoid width; the length of the tympanic bullae is roughly 18% of the
condylobasal length and varies in the range of 27-30 mm. The nasals are

201

265

long (20-25% of the condylobasal length) and narrow (width at the base
of the apex 15-18% of the total length), their anterior edge terminat-
ing in three angular projections, of which the middle one is sometimes
barely bifurcated and shorter than the lateral ones in most cases. They
wedge into the frontal bones to one-fourth to one-third their length. The
uncinate processes of the pterygoid bone are highly elongated and rather
low. They terminate anteriorly with an elongated pointed projection. The
posterior edge of the bony palate bears a deep arcuate notch, quite often
assuming an angular form at the apex. The compact longitudinal bony
septum in the choanae is faintly visible and runs posteriorly not beyond
the very beginning of the palatine bones.

The upper premolars, commencing from the second, have a highly
prominent main cusp and two weakly developed additional cusps; some-
times the anterior cusp is totally lacking and the posterior one better
developed; one more cusp is faintly visible behind the posterior cusp. The
upper molar has two cusps; it is usually separated from the fourth premo-
lar by a prominent diastema, larger than the gap between the rest of the
teeth. The corresponding lower premolars have better developed acces-
sory cusps than in the upper teeth; one of these accessory cusps is ante-
rior and usually two posterior. The true lower molar bears three cusps.

Sex-related differences in the skull are manifest mainly in the slightly
wider rostrum in males at the level of the upper canines and in the
ratios between the length and width of the tympanic bullae: in males
these are slightly longer than wide (length averages 29.7 mm and width
28.8 mm) while in females these values are identical (average 28 mm)
(Yu.K. Timoshenko). Moreover, in males the upper maxillary bones form
a more prominent bulge anterior to the orbits.

Indices of the width of the cranium and the facial length, width of
the rostrum and zygoma, the first molar, uncinate processes, as also the
height of the face, increase with advancing age. On the other hand, the
indices of the width of the mastoid and cranium as also the height of the
latter decrease (Smirnov and Chapskii, 1932).

Individual variability of the skull is fairly large both with respect to
dimensions and structural features. Among these, for example, the shape
of the anterior margin of the nasals is variable. Although their lateral
projections are usually longer than the middle one, the latter is some-
times slightly longer than the lateral or equal to them. The form of the
median process of the nasals is also variable: it is sometimes symmetri-
cally made up of both the bones while sometimes only one bone plays a
major role in its development; sometimes, it is made up wholly of one
or the other bone; quite often, it is bifurcated; and in exceptional с ases
it may even be absent (Gadzhiev, 1957). As in many other species, the

202

266

length of the contact line of the maxillary bones with the nasal bones,
the depth of wedging of the latter into the frontals, the base width of the
apex and the anterior margin of the nasals, the shape and depth of the
palatine notch, the length of the diastema between the fourth premolar
and the first molar in the upper jaw, the number and degree of develop-
ment of the accessory cusps on the crown of the premolars and molars,
etc. also vary.

The body length of adults measured along the body surface from the
nostrils to the tip of the tail (Lc) is generally 130-140 cm, sometimes
going up to 150 cm (or even more) in the largest animals.

The body length measured in a straight line averages 10 cm less.

Sex-related differences in the body length are seen only in averages.
Thus the average length (Lc) of mature (older than nine years) males is
134.2 + 0.77 cm and of females 131.7 + 0.47 cm (Yu.K. Timoshenko).

The total weight of the adults in.the period of their utmost well-being
varies from 50 to 85 kg (average 70 kg), the subcutaneous fat with the
skin accounting for up to 40-50 kg. The animals attain maximum weight
late in autumn and early in winter. They are impoverished by spring and
their weight falls, on average, to 40-45 kg (their maximum weight at this
time does not exceed 65 kg); the weight of the blubber (with the skin)
decreases, on average, up to 20-25 kg (more than 30-32 kg of fat in the
skin is unusual in this season).

The condylobasal length of adults aged 10 or more years is
170-190 mm, the width of the cranium at the mammiform (mastoid)
processes 85-100 mm, and the width at the zygoma 89-150 mm.

The size difference of the skull between males and females is not
much. Among males, the condylobasal length varies from 171.6 - 190.3 mm
(x = 182.1,) width at the mastoid 88.2 -97.8 mm (x = 93.4), width at the
zygoma 91.6 - 104.6 mm (x = 98.7), width above the canines 24.1 - 29.2 mm
(x = 26.4), and interorbital width 4.5 -7.7 mm (& = 5.9).

The condylobasal length among the females varies from
171.8- 182.0 mm (х = 175.1), width at the mastoid 85.0-95.5 mm ( =
89.6), width at the zygoma 89.3-98.7 mm (х = 93.6), width above the
canines 23.2 -27.0 mm (x = 24.8), and interorbital width 4.0-6.7 mm (М
5.6) (Yu.K. Timoshenko). (K.Ch.)

Taxonomy

Genetic relations with the closely related species are defined by grouping
this species in the same subgenus (or genus) with the ringed seal (Ph.
hispida) and Baikal seal (Ph. sibirica). These are no doubt extremely close
but the interpretation of the Caspian seal (like the Baikal) as a subspecies

203

267

of the ringed seal (Phoca hispida Schreb.) has not been supported by
contemporary scientists.

The direct evolution of the Caspian seal has not yet been conclu-
sively established. In fact, recent descriptions of some paleontological
finds, a review of former finds, and also an analysis of the morphological
features have brought this problem closer to a solution. Nevertheless,
there are two main hypotheses explaining the evolution of this species.
According to one, the seals colonized the Caspian Sea in the Quaternary
period having somehow entered there from the north under the influence
of the glaciers which displaced some part of the population of the ances-
tors of the ringed seal to the south. Having entered a new water body,
they were ultimately transformed into the present-day Caspian seal.

With regard to the Caspian seal’s entry into the Caspian Sea, some
researchers assign a decisive role to the extensive freshwater body formed
as a result of the significant head of river waters over the vast expanse of
northwestern Siberia (Pirozhnikov, 1937; Davies, 1958). This water body,
extending up to present-day northeastern Kazakhstan, encompassed the
present-day Aral Sea and came close to the Caspian, with which it could
have come into contact through the river streams, and came close even
to Lake Baikal.

According to the second hypothesis, based on paleontological data
for the Pliocene and the Miocene, the Caspian seal is regarded as a
descendant of the seals of the subgenus Pusa inhabiting the successively
superseded Sarmatsk, Meotichesk, Pontichesk, and other basins, lead-
ing ultimately to the formation of the present-day Caspian Sea. Several
fossil finds of Upper Tertiary and Early Quaternary seals from along
the periphery of the Caspian and in other southern regions reveal fea-
tures of close genetic affinity with the present-day Caspian seal. The
second point of view received fresh substantiation in the last decades

(Chapskii, 1948, 1955; Gadzhiev, 1959; Kirpichnikov, 1964). Evidently
there is greater justification for the hypothesis of the autochthonous evo-
lution of the Caspian seal than for considering it as having arrived from
the north.

All the three branches of seals of the subgenus Pusa, i.e., the Caspian,
Baikal, and ringed seals, became isolated presumably at the end of the
Tertiary and embarked on a long course of independent development.
(K.Ch.)

Geographic Distribution

The geographic distribution is limited exclusively to the Caspian Sea,
from which some rare strays into the Volga and Ural have been recorded.

268

А good number of seals are encountered in some seasons throughout
the sea, from the coastal regions of the northern Caspian to the Iranian
coasts. They are widely distributed throughout the sea in summer, at the
end of the breeding and molting periods. Then they are found simulta-
neously in various regions of the northern shallow waters, right up to the
northeastern extremity of the sea, in the region of Tyulen’ archipelago
(Kulaly, Morsk, Rybachii, Podgornyi, Novye, etc. islands north of
Mangyshlak Peninsula), on the western side of the sea (commencing
from the prodelta of the Volga to Lenkoran and farther south), and
along the eastern edge of the sea from Mangyshlak to Atrek and more
southward (except in Karabogazgol), and also in its central parts.

The nature of the summer-autumn distribution of the seals is depen-
dent on food conditions. Concentrations of schools of fish and other edi-
ble items can bring about fairly intense concentrations of seals at times,
although they are thinly dispersed in the summer season. In the northern
part of the sea, sporadic concentrations are seen on Zhemchuzhnykh,
Rakushechnaya, and Kulalinsk banks and additionally between Cape
Urdyuk and Kendirli Bay during massive concentrations of sprat there
(Badamshin, 1948). By autumn, they are confined preferentially to the
northern Caspian at places of feeding as well as in regions of island rook-
eries. Autumn concentrations are particularly regularly seen on small
dry islands with shell and sandy shores (shalyg and “pleshina”) in the
northeastern corner of the sea at Karatonsk, Suendykovsk, Balashovsk,
Kolkhoznye, and Borozdinsk, and also on the Zyuidvestov Islands in the
fore-estuary of the Ural, Utaespinsk, and Kulalinsk Rivers, and on Malyi
Zhemchuzhnyi Island (В.Г. Badamshin). The seals gather in the southern
Caspian on the islands of the Apsheron archipelago and on Ogurchinsk.
They live in relatively small numbers on all the above islands in sum-
mer also.

The period of autumn concentrations of seals in the region of bald
patches extends usually from September to the time of complete freezing.
In winter most of the population is concentrated in the icebound north-
ern part of the sea but occasionally stray animals and small groups are
encountered at this time even in the more southern regions of the sea.
From spring,.on the contrary, most of the population migrates southward
and is dispersed over wide expanses in the central and southern Caspian
(see p. 270).

Geographic Range outside the USSR

Covers only a narrow strip of the southern Caspian in the territorial
waters of Пап. (K.Ch.)

204

269
Geographic Variation
Not reported.
Biology

Population. It was impossible to make a direct count of the population
of the Caspian seal until the early 1970s and only indirect indices were
used for estimations. These indices are the stability or dynamics of the
annual catch, areas of the nurseries, counting from ships the number of
animals sighted in the water, etc.

In the past, when hunting exceeded an average of 100,000 animals
per annum for 90 years (1824 through 1915), the population of this seal
was large and evidently exceeded a million.

It has not yet become possible to count the present-day population.
A maximum of 750,000 animals was cited for the end of the 1950s and
470,000 for the mid-1960s (Badamshin, 1960, 1966). Even a figure of
600,000 was advanced for this period (Chapskii, 1966).°? The latter two
figures represent the main reserve, i.e., the population before breeding
(without pups of the current year). One of the fairly reliable bases for
these calculations was the position prevailing in the hunting season at
the beginning of 1966 when, supposedly, 88,500 pups were killed. But
even then, the population was calculated largely on arbitrary assump-
tions of the percentages of the rest of the herd constituents: quantitative
proportion of immature animals of both sexes and productive animals.
The figure of 500,000 - 600,000 is perhaps quite close to the present-day
size of the herd.

Habitat. The Caspian seal is not very particular about its habitat
conditions (aquatic). It is found in very shallow water regions and in the
zone of extremely large depths along the coasts as well as in the pelagic
sections of the sea. It is biologically associated with ice floes on which
it breeds and suckles the pups and also spends much of the molting
period. It has no special requirements for the ice floes except that they
be quite stable. Land is also used for setting up rookeries: bald patches as
well as sandy-pebbly and rocky coasts of islands. These seals avoid coasts
overgrown with reeds and other vegetation. They are not very fond of the
rivers though they ascend the Volga in rare cases up to Astrakhan and
even up to Volgograd (Badamshin, 1966). They are obviously not very
timid and do not avoid waters with fairly active shipping; for example,

62 Some foreign publications (Scheffer, 1958; King, 1964) put the Caspian seal population
at 1.5 million, a figure communicated at one time by these authors to S.V. Dorofeev.

205

270

they enter Tyub-Karagansk Вау (in Port Bautino). They, however, select

deserted, isolated, and uninhabited islands, and bald patches for forming
coastal rookeries.

Food. These seals feed throughout the year evidently without pro-
longed or total interruptions but with varying intensity in different sea-
sons. Feeding is not very intensive in late autumn, winter, or early
spring nor in summer although this phenomenon is manifest to different
degrees in different groups. Animals undergoing parturition and suckling
mothers feed less regularly and sometimes probably remain half-starved.
On large-scale autopsy of adult seals caught in the nurseries in Febru-
ary, food was found in nearly all the stomachs (Dorofeev and Freiman,
1928) and only a few (Roganov, 1931) contained a small amount (five
of 30 animals autopsied). Similar observations were reported even later
(Badamshin, 1948; Kurochkin, 1958) and hence it is hardly an exaggera-
tion that, after winter, the half-starved seals “starved in the winter, rush
to feed” in the spring (Kurochkin, 1958). However, in early spring, during
the molting period, feeding is even further weakened due to prolonged
resting of the animals on floating ice floes, ice mounds, and later even
on bald patches. At this time, food remnants were detected in 8 to 16%
(Badamshin, 1948) and sometimes in 30% (Roganov, 1930) of the seals
caught. From mid-April, the seals abandon the ice floes and bald patches
in the northern part of the sea and take to even more intensive feeding,
especially along the coasts of southern Dagestan in the second half of
April and in early May.

In the autumn-winter period, in Mangyshlak region, even in years of
fish abundance, the seals fed almost exclusively on trash fish or fish of low
commercial value, and also on crustaceans and some other invertebrates.
Of the 150 seals with food in the stomach, the majority were found
to have fed on goby followed by sprat and sand smelt, and only 2%
contained the remnants of pike-perch (Samofalov, 1931* ). In the stomach
of nearly 400 seals caught in nets in the same region and in the same
months but in much later years (1939 through 1946), goby occupied first
place (in 29 to 43% of the animals), sprat accounted for 3 to 30% in
different years, while shrimps of different types played a major role (18 to
33%); further, 11 to 15% of the animals contained mollusks; sand smelt
was found in two seasons in 1.5 to 11.0% of the animals; and herring
was found in only two seals (Badamshin, 1948).

In winter, in the northern part of the sea covered by ice floes, the
seals subsist exclusively on fish of no commercial value, mainly goby and
various invertebrates. At the end of the 1920s, goby and amphipods pre-
dominated in the food of seals in this season; crustaceans and mollusks
were consumed very rarely while commercial fishes held no attraction

206

271

Fig. 124. Adult female Caspian seal, February, 1953 (photograph by Yu.V. Kurochkin).

at all (Dorofeev and Freiman, 1928; Terebenin, 1930; Roganov, 1931;
Samofalov, 1931*; Kurochkin, 1958).

The food of these seals in spring has not been adequately studied. In
earlier years the food in April consisted mainly of sprat, some sand smelt,
and herring to an even lesser extent. The latter was detected in only 14%
of the seals whose stomach contained food. The remaining items (small
Caspian roach, sabrefish, and amphipods) formed 2% (Roganov, 1930).
In the more southern regions in spring the nutritive value of commercial
fishes rose somewhat, especially when their arrivals coincided with the
residence of the seals. In the past this was promoted to some extent by
the use of sweep nets to catch herring. In early May, 1929, on the coasts
of southern Dagestan, the seals rushed to the nets, jumped into them,
and quite often were trapped inside them along with the fishes (Chapskii,
1930).

The summer food of these seals has been studied even less. However,
it is known that in summer as well as in early autumn, they are regularly
seen in the regions of sprat concentrations (Badamshin, 1966).

Experiments on the prolonged rearing of seals were carried out in
the basins of a fishing station in Bautino village and on Kulaly Island.
The seals there had a choice from diverse fishes; goby, sprat, herring,
pike-perch, Caspian roach, sand smelt, as also crustaceans—shrimps and

272

crayfishes. The first preference of the seals was goby and sprat and, when
these were abundantly available, the animals did not turn to other fishes’
or crustaceans. When the goby and sprat were inadequate, the seals addi-
tionally took to herring and Caspian roach; the last preferences were sand
smelt, shrimp, and crayfish (Badamshin, 1948, 1960). Evidently goby and
sprat could be regarded as the predominant food of the Caspian seal. The
animals initially consumed live fish exclusively but later became accus-
tomed to dead fish and in time could be hand-fed. A young seal 105 cm
long consumed in one sitting up to 100-120 sprat, i.e., 250-280 g; an
hour later, it consumed an additional helping of 30-40 more. The daily
ration of the seals of different ages held in the basin varied from 1.2 to
1.8 kg (x = 1.4) in the case of yearlings and 3.2 kg (up to 1.2 kg in one
helping) in older juveniles; an adult male consumed up to 1.7 kg in one
sitting and up to 4.5 kg in one day (see p. 276).

Home range. In the conventional sense of the term, a home range
is not characteristic of either single or groups of these seals. In differ-
ent seasons of the year, under different environmental conditions, and
to some extent also depending on the region, the extent of seal con-
centration is extremely variable; on a solid substratum—land or ice—it
is usually far greater than in water. The spring and autumn haunts are
most concentrated on bald patches, especially when a section of such
projecting from the water is small. Then the animals are often packed
densely. On the more extensive bald patches, 100 or more seals can be
found in an area of 100 m?. Somewhat more rarely than in the autumn
rookeries, but no less densely, the seals are disposed even in spring at
the time of molt on icy hummocks and drifting ice floes. On the latter
the animals rest only on the extreme edge, usually in single file, head
facing the water, in the form of a typical border extending for tens and
sometimes hundreds of meters.

The lactating females (mothers with pups, Fig 125) on the ice floes
of the northern Caspian are usually much less compact. Pairs disposed
at intervals of 5-6 to a few tens of meters and sometimes even farther
apart on the wide expanse of the ice floes, create the impression of a
typically large, highly scattered, herd. But the herd is nowhere united; the
animals are not concentrated in any definite section of the ice floes; on
the contrary, they are divided into sections on the wide expanse at fairly
large intervals (see p. 280). It is this feature that renders the application
of aerial photography for a quantitative estimation of the mothers or
pups extremely difficult.

The densest concentrations of seals are seen in the water firstly in
the period immediately preceding whelping, when the females ready to
birth are concentrated on the more suitable ice floes; secondly, when the

207

273

Fig. 125. Female (mother) Caspian seal with white pup. Caspian Sea, February,
1958 (photograph by Yu.V. Kurochkin).

molting rookeries are disbanded; and thirdly in the autumn, in antici-
pation of the drying up of the bald patches on which the animals can
gather. In all other cases the seals are distributed sparsely, mainly singly
or in small groups in which one animal is separated from another by
tens or even hundreds of meters.

Hideouts and shelters. Like some other seals with a biological affinity
for ice floes, the Caspian seal living in water among ice floes is compelled
to ensure for itself access to air for respiration and also some means for
crawling onto the ice and re-entering the water. Air holes, open water
pools, and cracks in the ice floes are inadequate for this purpose and the
animal is thus compelled to make openings in the ice and to maintain
them. They resort to such, however, only when the sections of open
water surface among the ice floes is covered by a continuous crust of
recently formed thin ice. When its thickness is 1-2 cm, the seals can
easily pierce it from below with their head but do not necessarily use
such holes (Fig. 126). If, however, the ice thickens rapidly, they use the
holes made earlier. With time, depending on the thickness of the ice,
these openings (air holes), like those made by the ringed and harp seals,
acquire the form of a truncated cone due to constant use, narrowing
toward the top and opening on the surface of the ice floe or covered by

209

208

Fig. 126. Caspian seal in a wide hole. February, 1958 (photograph by
Yu.V. Kurochkin).

a thin icy arch. This arch is formed gradually, partly due to the freezing
of the water splashed during diving, and partly by the snowdrifts thawing
inside under the influence of the warm exhaled air. At the center or side
of this arch is a narrow throughway 5-10 cm in diameter (Badamshin,
1948). If or when the ice cover is broken and open water pools form in
the vicinity, the seals abandon their former air holes and use the open
water for respiration until it is covered by an ice crust, compelling a new
air hole to be made.

Later, as the whelping period approaches, gestating females scrape
the air holes with their claws, widening the upper part of the cone in the
same manner, and transform the initial air hole into a lair which is used
later for crawling onto the ice and re-entering the water throughout
the lactation period. The same lair is often used by several animals.
The Caspian seal makes no other hole or lair in the snow or in the
icy hummocks for the pup, preferring to undergo parturition directly on
thick stable ice floes with hummocks that can protect the newborn from
the winds.

Daily activity and behavior. These aspects are even less studied. The
daily activity and behavior are closely dependent on the seasonal peri-
odic phenomena which, in the life of the Caspian seal, proceed in the

275

same sequence as in the case of other species. The commencement of
the calendar year marks the period of high activity. The seals have to
acclimatize to the formation of ice floes; the adult animals are concen-
trated in the breeding zone and the females on the eve of whelping look
out for appropriate sites. The “nursing” period involves the care and
suckling of the pup, which conforms to no daily rhythm of activity as at
other times. The mating period follows thereafter.

With the conclusion of these stages of the annual cycle, a relatively
more quiet molting period sets in. The animals rest on the ice floes for
long periods and later on the bald patches in the same region, lost in
sleep and drowsiness that is interrupted rather rarely and at irregular
intervals for food intake.

The origin of early rookeries on the northern bald patches is evi-
dently related to weather conditions. They are formed by the same groups
of seals which have not been able to complete molt by the time the ice
floes have thawed (see p. 282). After severe and prolonged winters, these
rookeries are formed only in May while, after normal or mild winters and
in early spring, they are formed even by April. The spring rookeries are
usually active for not more than a month and comprise three-fourths
juveniles of both sexes and one-fourth adult females; the adult males in
them do not exceed 0.5%. The animals are found on bald patches even
in summer but their concentrations at that time are unstable and less
numerous and consist predominantly of juveniles; they gather at night
and usually go into the water in the morning (Badamshin, 1948a, 1950).

Thus these contingents, like the other seals which have completed
molt early, again enter a period of high activity in the second half of
spring. Throughout the rest of summer, up to mid-September, they are
seen in the water scattered singly or in small but sparse groups. They
can sleep in water as well as on land. Their activity at this time depends
mainly on their appetite, which continues to be voracious until they are
fully satiated. Summer and part of autumn represent the most important
period of feeding, when the animals not only recoup their fat reserves,
but also accumulate them to face the oncoming winter.

In the basin satiated animals sleep for hours quietly, only sometimes
turning from one side to the other or from the back onto the belly. Hun-
gry animals, however, are restless and very active (Badamshin, 19486*,
1960).

In autumn, having fed themselves fully, the. seals once again turn
to land, particularly to bald patches in the northeastern part of the sea,
but their activity has sharply decreased by this time. As long as the bald
patches are covered by high water, the seals swim leisurely in their prox-
imity or lie immobile on the surface in anticipation of their exposure by

209

276

а surging wind. As soon as such conditions set in, the animals crawl onto
the shoals, which are quite often covered completely with their bodies.
On quiet nights, discordant sounds can be heard occasionally from this
crowd and various minor skirmishes occur here and there (inevitable in
close living quarters). The seals sleep most often on the side, but quite
often on the back or belly. The neck is foreshortened, the hind flippers
extended, and the fore flippers pressed close to the body. One of the
hind flippers often forms a fist and is concealed by the other. From time
to time, for short intervals, the hind flippers are opened out fanlike,
then dropped limply but extended to the ground. Then the seal raises its
head and hind flippers high, as though stretching, bowing its body, then
once again resumes its original posture. On clear, sunny days the seals
rest a long time on the bald patches; in cloudy weather, especially in
rainy weather, they become restless or leave the bald patches altogether;
a thunderstorm drives them into the water (Badamshin, 1948).

The autumn rookeries on the northeastern bald patches comprise
males and females of different age groups but productive males and
immature animals of both sexes predominate in them. Mature females
are relatively few.

Like the autumn rookeries, the spring-summer island rookeries
formed in the 1930s in the south, especially in the region of the Apsheron
archipelago, also consisted almost exclusively of adult males with a
negligible admixture of juveniles of different age groups. Adult females
appeared only as solitary individuals (Kleinenberg, 1939).

The seals are naturally afraid of man, especially in the rookeries
on bald patches. However, by assuming the posture of a lying animal
and moving quietly, one can crawl close to them. The self-preservation
reflexes are poorly developed among the young compared to the
adult. Instances of the inexplicable gullibility of the animals have been
recorded. Thus in the autumn of 1959, one yearling that had evidently
strayed from the herd onto a bald patch, repeatedly attempted to get onto
a drifting ship, apparently because of its overpowering need to find land.

Seasonal migrations and transgressions. The Caspian seal undertakes
extensive migrations annually and with high regularity. In spring, with
the thawing of drifting ice floes in the northern Caspian, the bulk of the
seals concentrated there even from the end of autumn to early winter,
constituting barely 0.9% of the total population, begin to move gradu-
ally southward. The first to leave are the females that have undergone
parturition, followed by their suckled and molted pups. Later, having
completed molt, the adult males too move in the same direction; the
last to abandon the northern waters are immature animals of both sexes.

7/1)

This process is protracted since the movement southward coincides

210 with intensive feeding. By the end of spring to the commencement of

summer, much of the population has moved into the central and southern

Caspian Sea, where the animals, wandering from place to place, remain
until the end of summer before moving northward subsequently.

These seals migrate mostly in very sparse, small groups, forming no
close-knit herds. They do not usually move continuously in the same
direction, especially during their southward migration, but nonetheless
reach the south although the migrations appear disoriented at first
glance. There are presumably no strictly fixed routes for journeying south
and returning north. While feeding on the way, the animals move in a
broad front, some closer to the western coasts of the sea, others strictly
along the eastern rim, and some others quite far away from the coasts.
Nevertheless, the seals visit the same regions en route year after year.

The intense summer warming of the water, which evidently the seals
cannot withstand, is usually regarded as the most important factor caus-
ing their southward migration in spring from the shallow northern parts
of the sea. In the central and southern parts of the sea, because of the
greater depths and constant mixing of the water body, there is no warm-
ing up of the surface waters and hence more favorable temperature con-
ditions are created for the seals. Without doubt, the food factor. also
plays an important role.

In the summer months most of the seals are confined to water, usu-
ally sparsely, and it is difficult to say where they are more numerous
at that time. According to recent observations (В.Г. Badamshin), much
of the population is distributed in summer throughout the expanse of
the central and southern Caspian. Evidently the seals do not require a
solid substratum in summer. Therefore, as in the past, now too seals
are encountered in summer irregularly and, too, in tens or rarely in
hundreds in the rookeries surviving in the southern Caspian: on sev-
eral islets (oil stones) along the Apsheron Peninsula, nameless islets at
the inlet of Krasnovodsk Bay, Ogurchinsk Island, and others. With the
onset of autumn, much larger rookeries are formed at these places but
they nowhere are equal in size to those characteristic of the northern
Caspian.

With the approach of autumn, almost all the seals that have fed or
are continuing to feed begin to gradually migrate northward again. The
seals moving in this direction are abundant in the Mangyshlak region,
Tyulen’ archipelago, and the extreme northeastern, highly shallow part
of the sea. Much of the population is concentrated in the northern and
northeastern Caspian even before the formation of ice floes. It is in such
places on the northeastern bald patches that the largest prewinter seal

211

278

rookeries are formed regularly at present. With the approach of winter
and the first appearance of ice floes, the rookeries on the bald patches
break up and the whole seal population approaching the north moves
slightly southwest and is later confined mainly to. the edges of the ice
floes that have formed over a large area.

Concurrent with these local movements of the animals that have
already arrived in the north, animals continue to arrive from the south,
from the regions of the central Caspian. Judging from seals caught in
nets, formerly practiced at Mangyshlak, even mothers and productive
males continue to migrate northward in December and January (Samo-
falov, 1930).

Some seals, contrarily, join no group traveling north in autumn but
remain in the south for winter. These are mainly immature animals and
an insignificant number of adult males. The latter have lost the urge
to migrate for some reason (probably diminution of the reproductive
function) and their herd instinct (or other ecological factors caused by
subordinate relations) holds them to their young. |

With the approaching whelping period, gestating females enter deep
into the icy environment by any possible means, search out the most
stable large ice floes and, when the time for parturition approaches,
crawl onto the ice. Since they migrate in herds and the season of births
is not very prolonged, a large number of animals are seen on the ice
almost simultaneously and quite close to each other. The first rook-
eries on ice floes thus initially consist almost exclusively of gestating
females, with pups appearing later. The adult males remain until this
time with the immature animals of both sexes near the edge; they begin
to crawl on the ice slightly later and form individual, very dense, compact
rookeries.

Reproduction. Like other species of the subfamily, the Caspian seal
too does not form pairs for long. Because pairs are short-lived, encoun-
ters between competitors are not very serious. The natural population
ratio of adult males to females is close to one but due to unilateral
hunting in recent years (see p. 289), there is evidently some surplus
of productive males even now (Badamshin, 1966a). However, it would
be more appropriate to view the situation as a shortage of productive
females.

Among females that have attained maturity but have not yet under-
gone parturition, though ovulating (corpus luteum present in one ovary
in autumn), the uterus varies from 25 to 50 g in weight and the ovaries
from 2.0-3.5 cm in length and 1.0-3.8 g in weight.

The testes of mature males are 7.5 -8.0 cm in length and 20-35 g in
weight (with the appendage).

212

279

The mating season commences roughly mid-February, 1.е., even
before lactation ceases, and ends in the last few days of the same
month or in the first few days of March (Dorofeev and Freiman, 1928;
Yu.V. Kurochkin and B.I. Badamshin) or even extends up to March
20 (Smirnov, 1930*). The total duration of gestation has been roughly
determined as 11 months. However, the duration of active embryogeny
(after implantation of the blastocyst) and that of the latent stage have
yet to be fully established. Evidently they do not differ significantly from
the corresponding values for other closely related species.

The whelping of most of the females occurs in a short time span
of 10 days, from the end of January to about February 5. In very rare
exceptions, the pups are born slightly earlier, or more often, later. In any
case, the total period of whelping, with occasional exceptions, extends to
no more than 20 to 25 days and ceases by February 10. Much later solitary
cases of births have been reported: on February 27 (Samofalov, 1930)
and even in the first few days of March (Roganov, 1931). Even greater
but extremely rare deviations have likewise been reported (Badamshin,
1948). One gestating female was found on May 10, 1942, on Suendykov
bald patch. The embryo was 53 cm long and, theoretically, this female
should have undergone parturition roughly 1.5-2 months later, 1.е., at
the end of June or in the first half of July. A pup 79 cm long was caught
with its mother on July 28, 1941, on Balashov bald patch. Judging from
its state of intense molting, the pup was no more than 2-2.5 weeks old.

The litter of the Caspian seal, like that of the other species of Pin-
nipedia, consists of a single pup, two being extremely rare. At birth, the
body length of the pup varies roughly from 70 to 75 cm (Lc) while the
weight varies around 3-4 kg. The newborn is almost devoid of subcuta-
neous fat layer. The hair coat is usually of the embryonal type: dense,
long, silky, almost pure white (with a creamy-pistachio tinge in the first
few days), often with a smoky-gray bloom on the dorsal side.

The period of lactation extends evidently for about four weeks. In
the first few days after birth, the mother almost does not lose sight of
her pup, suckling it repeatedly at different times round the clock. Later,
she leaves the pup for a long duration, thus suspending suckling for
prolonged periods. The pup spends much of the time sleeping on the
ice floe, and waking up, begins to look for its mother. Often, not finding
her, it crawls to an air hole and peers into the water as though expecting
its mother to emerge. When the wait is long, it calls in a voice resembling
the wail of a child (Badamshin, 1949). To suckle the pup, the mother
lies on her side or almost on her back, exposing the teats.

Until recently, there were no rational data on the age at which matu-
rity sets in among these seals. It was assumed (N.A. Smirnov, 1931) that

280

the female attains maturity at two years of age and the male а year
later. Similar views were expressed even more categorically (Roganov,
1931). Later, based on raising seals in an artificial basin on Kulaly
Island, the view was expressed that they attain maturity at five years
of age (Badamshin, 1960). Finally, by studying the reproductive organs
in relation to different age-related criteria, it was demonstrated that the
females commence undergoing parturition mainly in the sixth year of age
(Chapskii, 1965а*).

Not all the females that have attained maturity reproduce every year
but there are as yet no reliable data on the extent of barrenness. Pre-
sumably, it covers 15 to 30% of the eligible females (Badamshin, 1950,
1960).

The seals gather for breeding on ice floes in the northern part of
the sea mainly in the Gur’evsk Channel region and west and southwest
of it (relatively close to the edge on a Strip extending roughly from
the meridian of Kulaly Island to Rakushechnaya and Zhemchuzhnaya
banks). Depending on weather conditions, the maximum concentration
of mothers with pups is seen on all sides. Further, when the winter is
mild and late and hence the expanse bound by ice floes is relatively small,
the seals reproduce far away in the northeast. Because the area of ice
cover is small, the region inhabited by the seals is greatly reduced and
their concentration perceptibly increases. A reverse picture is observed in
abnormally severe winters. The area of ice cover then becomes markedly
enlarged, the rim of the ice floes extends far southwest and the bulk of
the animals move in the same direction as the ice floes.

For whelping, the most stable ice floes with fairly level sections inter-
spersed with hummocks are chosen. The females prefer the rims of such
ice floes but, in their absence, are compelled to give birth even on very
smooth young ice that is strong enough not only to sustain the weight of
the gestating mother, but also to withstand some thrust of the ice floes
in a fresh wind.

The herd instinct, characteristic of the Caspian seal, combined with
the need to select the most appropriate ice floes for whelping, results in a
fairly high concentration of reproducing females. Their disposition with
pups on an ice floe largely resembles the nurseries of the harp seal but
of smaller size and density. The animals lie widely scattered with highly
dispersed groups (so-called “spots”) varying in size and concentration
alternating with floes uninhabited by animals.

Some spots consist of several tens to hundreds and even a thousand
animals. In the gaps between the spots, often running into several kilo-
meters, there are no animals at all or they are encountered very rarely.
Sometimes, however, the spots are quite close to each other.

214

281

Growth, development, and molt. The young pup grows and devel-
ops as rapidly as any other earless seal. After a brief lactation period,
the pup grows roughly 20% longer (compared to its original length) to
85-90 cm (Lc) while the original weight increases more than four times.
It accumulates up to 8-12 kg or more of subcutaneous fat.

The following data describe the growth tempo of the young female:
body length (Lc) at about nine months varies from 86 to 104 cm, average
94.5 cm; in autumn of the second year, the average is 107 cm; in the third
autumn 114 to 119 cm (average 117 cm); and by the fourth autumn the
average is 124.5 cm. Further growth is highly retarded (Chapskii, 1965а*).

In the normal course of lactation, 15-20 days after birth the belek
[= white pup] begins to molt. Initially molting is barely perceptible but
with each successive day, its furry white hair is increasingly shed. The
coat becomes less even and shaggy and is interspersed with dark-colored
bald patches and covered with the new short and bristly hairs typical of
seals. In this phase of intense molting, extending roughly for two weeks,
the pup is called a “tulupka” [highly molting pup] (Figs. 127, 128). En
masse molting of pups usually occurs around the 10th of February and
is completed in the majority of them by the end of February or the first
few days of March. The pup that has completely shed its juvenile hair
coat is called а “sivar”. Lactation quite often ceases in the last stages of
molt but often the mother suckles even the “sivar”.

The upper side of the hair coat of a “sivar” is almost monochro-
matic dark gray; spots, even when visible, are not always very distinct.
The underside is light-colored, whitish, and without spots in most ani-
mals. From year to year, with every successive molt, the spotted pattern
becomes increasingly evident on the back. Spottiness is seen more sharply
in males and less so in females. The color on the ventral side is also as
variable: in males, with time, the number and brightness of spots increase
ever more but in females the increased spottiness is usually faint, dull,
and even altogether absent in very rare cases.

Molt is protracted in the first year and subsequently in all animals
commencing from yearlings. In some animals it apparently continues
for a little longer than a month. The total duration of molt, however,
has been put variously at 1.5 months (Dorofeev and Freiman, 1928) to
2 months (N. Smirnov, 1931) and even up to 3-4.5 months (Badamshin,
1948, 1965*). Molting commences first in the females that have given
birth (in many cases even during lactation); it commences somewhat later
among adult males and then among immature animals of both sexes (in
the so-called “zheltyaks”’).

Not all the seals succeed in molting on drifting ice floes; many of
them are forced to continue it on icy hummocks and when these also

282

213 Fig. 127. Intensely molting young Caspian seal (“tulupka”). February, 1958 (pho-
tograph by Yu.V. Kurochkin).

break up, molting continues on bald patches and islands. Immature ani-
mals, together with a fairly good admixture of adult males, form late
molting colonies. Adult females however complete molting on the drift-
ing ice and only small numbers of them are encountered on bald patches
in spring. Weather conditions play no mean role in the disposition of
molting animals. When the ice floes hold for long in the northern part
of the sea (which happens in very severe winters in which, evidently, the
ice floes break up slowly), the seals molt on them; otherwise, when the
winter is mild and the floes thaw rapidly, many animals are compelled
to complete molting on land (on the northern and southern Caspian
islands).

Enemies, diseases, mortality, and competitors. The factors and magni-
tude of natural mortality of the Caspian seal are not yet fully understood.
There are no natural enemies of the seal in water, with the exception,
perhaps, of large white whales which may catch very young seals from
time to time.

The main enemies from among the land vertebrates are the long-
tailed [pallas’] sea eagle (Haliaeétus leucoryphus), white-tailed [gray]
eagle (H. albicilla), and partly the golden eagle (Aquila chrysaetus), but

213

215

Fig. 128. Young Caspian seal (“tulupka”). End of February, 1958 (photograph
by Yu.V. Kurochkin).

their role as enemies of the newborn is hardly significant. Foxes and
wolves rarely pursue the seals on ice and the damage caused by these
predators is perhaps extremely small, though it has been mentioned in
the literature (Khastatov, 1894*; Badamshin, 1949).

A comparison of the helminth fauna of the Caspian seal with that
of the northern seals (from which, according to some, the Caspian seal
has evolved) established that the Caspian seal “rid itself of almost all the
parasites characteristic of its northern kin but has become the host to
large numbers of two or three new species of parasites” (Dogel, 1947*;
Delyamure, 1955). Excluding the unidentified larval forms, 12 species
have been established among the helminth fauna of the Caspian seal
(Kurochkin, 1958, 1961,* 1962; Kurochkin and Zabolotskii, 1958; Delya-
mure, Kurochkin and A. Skryabin, 1964). From among the trematodes,
Criptocotyle lingua is a facultative parasite of the intestine; Сшгеапа
badamschini infects all the animals, with a few to half a million spec-
imens being found in the intestine of a single seal depending on the
severity of the infection; Maritrema sobolevi infects the intestine of 35%
of the seals with up to 300,000 specimens present when the attack is
severe; Mesorchis advena infects the intestine of 95% of the seals with up

284

to 300,000 specimens present in the case of severe invasion (Kurochkin,
1962); Opisthorchis felineus, a dangerous parasite of man and carnivo-
rous mammals, infects the liver of 25% of the Caspian seals®?: and Pseu-
damphistomum truncatum infects the liver rather infrequently. Among
the cestodes, Ligula colymbi is a facultative parasite of the stomach and
Diphyllobothrium phocarum a parasite of the intestine. The nematode
Anisakis schupakovi infects the stomach and is sometimes encountered
among the seagulls but does not attain maturity in them. The larvae
of this nematode infect almost all the fish found in the Caspian Sea
(Kurochkin, 1961*), Eustrongylides excisus is detected in the small intes-
tine, and Parafilaroides caspicus parasitizes the lungs of 20% of the seals
with an invasion intensity of a few hundred specimens. The acantho-
cephalan Corynosoma strumosum infects the intestine of 70% of these
seals (Kurochkin, 1962).

The helminth fauna of the Caspian seal offers justification to con-
clude that “the transition of the host to relict conditions of existence
was marked, on the one hand, by a spurt in the speciation of parasites
and, on the other, by a sharp modification of its parasite fauna” (Dogel’,
1958). Thus the characteristics of the helminth fauna of the Caspian seal
can serve as a reliable proof of the relict nature of this animal.™

Specific data on diseases and epizooty are very scarce though it has
long been known that a large number of seal carcasses, the so-called
“plyvuns” or “plavurs,” are annually washed ashore in the Caspian Sea
in summer and autumn. The reason for such a loss of seals is not yet
fully clear. An inspection of dozens of carcasses in November, 1957, did
not reveal a single death that could be ascribed to helminth infection
(Kurochkin and Zabolotskii, 1958). Probably, a considerable proportion
of these floating bodies represents the wounded and killed animals that
were not collected during the winter-spring hunting season (Chapskii,
1931; Badamshin, 1948; Kurochkin, 1958). The unfavorable ice condi-
tions and storms which are particularly fatal to the young on ice floes
play no mean role in natural mortality (Smirnov, 1931). At the same
time, instances of the en masse death of seals during the second half

63 «Tt could be said with certainty, for example, that a large independent focus of
opisthorchiasis exists in the Caspian Sea. Suffering intensely from it during residence in
the freshwater zone (especially in the prodelta), the seals spread the eggs of O. felineus
with their excrement, thus infecting mollusks and later the fish through them” (Kurochkin,
1961*).

4 1 should be borne in mind that none of the three species of helminths that are
common to the Caspian seal and the northern species “1$ specific to the pinnipeds”. On the
other hand, the species of parasitic worms characteristic of the northern seals, especially
of Phoca hispida, are altogether absent in the helminth fauna of the Caspian seal. (K.Ch.)

216

216

285

Fig. 129. Caspian seal in open water pool. Gur’evsk Channel, February 1, 1966
(photograph by С. Nesterov).

of summer and in the autumn not only in the northern, but also in
the southern and central Caspian, compel us to look for some other
lethal factors. Pathological factors are also possible, including diplococ-
cus which attacks not only the liver, lungs, and intestines, but also the
muscles, joints, and skin. The percentage of diseases in some age groups
is quite high (Vilezhanin,* 1965).

Population dynamics. The population dynamics of seals in the
very early times when there was no hunting, in spite of the above
factors (helminths and diseases), could hardly have been significant.
After intense hunting began, it became the sole factor responsible for
population fluctuations, in the background of which natural changes
became imperceptible. It is also difficult to establish the effect of
hunting since direct population data are not available. Indirect indices,
primarily the extent of hunting, have to be relied upon (Badamshin, 1960,
1961).

_ The intensity of hunting in the first quarter of the nineteenth century
(see below) was quite stable and prolonged at 150,000 - 160,000 animals
(Sklabinskii, 1891) and this in itself points to the abundant population

217

286

of the animal at that time. However, some population reduction did
occur even then judging from the reduced average number of animals
killed annually, roughly at 110,000 - 120,000 in the next decade. This level
of hunting continued right up to World War I. Evidently the popula-
tion was still quite high and revealed‘no sharp drop although reduction
nevertheless continued.

During the years of war, revolution, and civil war, Caspian seal hunt-
ing decreased markedly, the fall in population was arrested, and the herd
multiplied. From the early 1920s, hunting recommenced and by the end
of that decade had almost reached the prewar level, at which it remained
until the mid-1930s. This restrained population growth and the herd
began to dwindle in proportion to the intensity of hunting. Large-scale
killing of the animals for several years in the 1930s severely affected the
reserves (a maximum of 227,500 animals were killed in 1935). The high
degree of hunting of mother seals and, concomitantly, pups, exerted a
particularly adverse influence. The process of herd stabilization, although
at a much lower level, was noticeable only in the 1940s while the pop-
ulation began to increase somewhat from the early 1950s (Badamshin,
1958*, 1960, 1961). However, by the mid-1960s, the Caspian seal popu-
lation was far less than what it was more than a hundred years ago and
this aspect has to be considered while planning the present-day hunting
levels (see p. 289).

Field characteristics. Recognition of the Caspian seal from its external
characteristics has never posed any difficulty since it is the only species of
seal in the Caspian Sea (and in the lower courses of some rivers entering
it). It is distinguished from the other species of the subgenus by color
characteristics, the growing young and the adult sport innumerable spots
of different shades, varying from gray (more rarely brown) to black in a
light-colored general background; the clear spaces between the spots are
usually not ring-shaped. (K.Ch.)

Economic Importance

As an important object of marine animal hunting, this seal occupies a
foremost place in the number of seals caught in the USSR. Hunting
(Fig. 130) has been practiced by the local coastal populations from very
ancient times; the valuable raw material, 1.е., hide, fur, and blubber, are
used in various industries.

The antiquity of seal hunting is supported by references to the
Caspian seal by Herodotus (N. Smirnov, 1931). The hunting of seals
and partly the processing of raw material are of vital importance to the

218

287

Fig. 130. Autumn congregation of seals surrounded by hunters on the shelly
shoals in the northeastern part of the Caspian Sea. October, 1958 (photograph
by КК. Chapskii).

economy of the Gur’evsk region of the Kazakh Soviet Socialist Republic
and Dagestan Autonomous Soviet Socialist Republic.

During the first quarter of the nineteenth century, 150,000 - 160,000
animals were killed annually (Sklabinskii, 1891). In the next nearly 50
years (up to 1867), the annual catch varied around 105,000. The maxi-
mum number, 290,000, was caught in 1844. From the end of the 1860s to
the beginning of World War I, an average of 115,000 seals were caught
annually.

In the decade from 1927 through 1936, the average annual catch
remained almost at the same level as in the prerevolution period, 1.е.,
at 115,600 (Badamshin, 1960). At the end of the 1930s (1937-1940), it
varied from 110,000 to 160,000, on average 147,500. In the 1940s, a much
smaller number, 33,000.(1945) to 88,000 (1948), on average 64,400, were
caught. In the 1950s, the hunting figures continued to drop: the average
annual catch decreased to 47,000, ranging from 23,700 to 71,500; the
average for the first half of the 1960s was 93,000 a year.

Until comparatively recently, the seals were caught in the Caspian
mainly for their subcutaneous fat and hide, the latter mainly used as a
raw material in the leather industry.

218

219

288

Among adult animals, during their maximum well-being (late
autumn and winter immediately before whelping), the skin with blubber
recovered by commercial methods (without flippers) weighs 40-50 kg,
even 60-70 kg in some individuals. The thickness of the skin with the
fat layer goes up to 7 cm at the breast level. In spring, however, after
lactation and mating, and in recently molted animals, the weight of the
blubber is reduced by half to 20 kg or less. A skin (without fat) weighs
3.5-4.0 kg. After summer fattening, toward autumn, the animals again
recover spent reserves. Among the normally fattened and completely
molted pups (‘“‘Sivar’’), the blubber weighs 10-13 kg and the skin alone
without fat about 1.2 kg.

The seals are presently hunted for their fur and therefore only in the
snowy period; the exclusive target is the newborn pup with a firm, furry
white hair coat or a fully molted pup in which this wool is shed and a
smooth and short fur has formed.

Wooden motor fishing boats with a load-carrying capacity of 35
to 40 tons are used for seal hunting. The aerial surveyor, who com-
mences his work 2-3 weeks before the hunting season, plays a signifi-
cant role. He usually arrives January end or early February. Additionally,
2-3 icebreaker-type vessels patrol the coasts and are available to those
seeking assistance.

In the recent past, when the seals were mainly caught for their blub-
ber and the fur was less coveted than at present, hunting continued
almost year round. In winter hunting proceeded among the broken ice
floes from fishing boats; when the ice floes became stable and firm, horses
and sledges were used to reach the Gur’evsk Channel (from Dzhambai
village to the northern coast). Early in spring, when the ice thawed in
the Volga, hunters set out in fishing boats and very small sail boats
(“reyushkas”) to kill seals molting on the drifting ice and icy hummocks.
After the ice floes disappeared, island rookeries were raided and the ani-
mals caught together with sturgeons using fishing gear. At places (mainly
in the region of Tyulen’ archipelago), fishermen in oar boats hunted for
the seals in water using firearms.

From spring until late autumn, hunting was carried out on the
Apsheron archipelago islands; from September until total freezing, in
the island rookeries in the northern Caspian; and from October through
April, by using nets in the region west of Mangyshlak Peninsula. Females
were the predominant targets.

To conserve the population, the hunting of adult females on ice
floes was banned in 1966 and the killing of all seals except beleks and
sivars was banned year round in 1967. Hunting of pups was restricted.

289

At present, although the total population has not yet been exactly ascer-
tained, it does not exceed 500,000 and only a catch of 70,000 pups can
be endorsed at the present time. Even this is an overestimate and in no
way ensures the restoration of the population to its former maximum
proportions.

With the prevailing state of the population, the number of animals
killed should not exceed 40,000 a year (V.D. Rumyantsev).

Since 1973 aerial surveys have been undertaken of seals in the north-
ern Caspian during the lactation period; such surveys will probably pro-
vide a new basis for an accurate census of the mothers and thus a rational
ground for determining the hunting quota.

The adverse role of this seal as a killer of economic fishes has been
discussed time and again. The reported instances of fish consumption,
including white salmon, by these seals from fishing nets at various places
(Varpakhovskii, 1891; Khlebtsov, 1902; and others) served as a basis for
derogating this animal as an enemy of the fishing industry (Grimm, 1907;
Averkov, 1914; and others). The general view thus created, consciously
or otherwise, served as an extremely convenient rationale for unrestricted
seal hunting, namely, destruction of a predator that killed large quantities
of valuable commercial fish.

The objective data that has become available over time has helped
to draw a more correct and complete picture, albeit not entirely com-
prehensive, of the food of seals, and also enabled a fresh assessment of
their role in the fishing industry.

Even at the beginning of this century (N.A. Smirnov, 1907) it was
found that items of very little commercial importance, especially goby,
were very significant in the winter diet of seals. Subsequent observa-
tions (Dorofeev and Freiman, 1928; Roganov, 1930, 1931; Samofalov,
1931*; Terebenin, 1930; Badamshin, 1948, 1960) convincingly demon-
strated that the Caspian seal was undeservedly branded an enemy of the
fishing industry.

Even in years when the Caspian abounded in herring, Caspian roach,
pike-perch, and others, not to mention the sturgeon, seals fed mainly on
extremely low value and noncommercial fishes as well as invertebrates.
Thus, in the spring of 1929, in the northern Caspian herring constituted
only 14% of the food of the seal while the small Caspian roach and sabre-
fish played such an insignificant role that, together with the amphipods,
they constituted only 2%; the seal subsisted mainly on sprat and to a
small extent on sand smelt (Roganov, 1930).

Rearing these seals under artificial conditions has revealed their defi-
nite food selectivity. In all cases, when the seals had a wide choice of food
items, they invariably preferred goby and sprat (Badamshin, 1948, 1960).

220

221

290

These seals have never had а predilection for the sturgeon. The
reported damage by the Caspian seal to the fishing industry in the
Caspian Sea is entirely baseless. Thus the Caspian seal is not responsible
for the depletion of the Caspian fish reserves. (K.Ch.)

BAIKAL SEAL or BAIKAL RINGED SEAL®
Phoca (Pusa) sibirica Gmelin, 1788

1788. Phoca vitulina var. sibirica. Gmelin. In: Linnaeus, Systema Naturae,
Ed. XIII, I, p. 64. Lakes Baikal and Oron (the latter reference is
erroneous).

1872. Phoca baicalensis. Dybowski (Dybovskii). Izv. Sib. Otd. I. Russk.

_ Geogr. Obshch., 3, по. 2, р. 86. Baikal.

1922. Phoca oronensis. Dybowski (Dybovskii). Arch. Tow. Nauk. Lwow,
I, p. 352, Nom. nudum. Lake Oron (there were never any seals in
Oron). (V.H.)

Diagnosis

The color of the hair coat at all ages is monochromatic, without spots
(Fig. 131). The fore flippers are totally covered by the hair; all the digits
on the hind flippers are covered from outside, and only the two extreme
ones are covered from inside. The claws on the fore flippers are long (up
to 5 cm) and strong, and have a high triangular outline in cross section.
The anterior edge of the nasal bones has no median projection.

Description

These seals are of small dimensions.

The hair coat is dense, short (up to 2 cm). The subcutaneous fat layer
is thick (12-14 cm). The bones of the shoulder and forearm, and of the
thigh and shank, are encased in a common skin cover. While moving, the
forelimbs elongate to almost double their length due to the stretching
of the forearm bones. The edge of the web extends beyond the tips of
the claws of the hind flippers on which the longest digit is the first.
The upper lip bears regular rows (usually eight) of 120 semi-transparent

6° Belyak—white pup that has not shed the embryonic hair coat; khubun or nerpyash-
molted underyearling; chernysh—one-, two-, and three-year-old seals; yalovka—unfertilized
female; matka-gestating or whelping female; and argal or sekach—mature male.

291

220 Fig. 131. Adult Baikal seal, Phoca sibirica (figure by М.М. Kondakov).

221 Fig. 132. Flippers with claws and cross section of claw of Baikal seal, Phoca
sibirica (figure by N.N. Kondakov).

whiskers varying in length from 10 to 100 mm; the longest are seen at
the corners of the mouth. The diameter of the eyeball may reach 47 mm
(Figs. 133, 134).

22

221

292

Fig. 133. Head of adult male Baikal seal, Phoca sibirica (photograph by V.D. Pastukhov).

Fig. 134. Whiskers of the Baikal seal, Phoca sibirica (figure by N.N. Kondakov).

The color of the adults is silvery-gray dorsally, lighter on the flanks,
and light gray, sometimes with an admixture of yellow, on the breast and
belly (especially in the arm pits). Age-related color variations are not
significant. Spottiness is seen as an extremely rare exception.

The skull is thin-walled; the zygomatic width is much more
than the mastoid width and considerably more than one-half the
condylobasal length (Fig. 135). The zygomatic arches are distinctly
seen when the skull is viewed from behind. The anterior edge of
the nasal bones forms only two lateral processes. The posterior edge
of the bony palate is usually in the form of braces (Fig. 136).
The tympanic bullae are not large and the gap between them
is usually equal to the length of each. The interorbital space is
narrow, not more than 4 mm in adults. The molars and premolars
have additional cusps, set without gaps; their accessory cusps extend
roughly parallel to the main one. The teeth in the upper jaw are

293

=)

wy

GEO oot а

ret yf),

ое
С ye > =
=

:
> ¥ ww
te" ‘he att

00

Fo ae

® 2,

`. А =
в МИ Bis
eo: by И : 5,
SS. a SUS и о с
= У ди. ау is Se АЙ ia S $ ЕС
ет ЗА S (ts BSS ES 5
X, iit: ae i WSS к „9 @ PF wave
\ ' Aes “
yes й

222

ard

тя “=

ita

2227

“>:
~

~~,

д = с к
=.
22777

777

77
as

Ages
<a

222 Fig. 136. Bony palate of the Baikal seal (figure by N.N. Kondakov).

usually more spaced than the lower ones (Ognev, 1935; Ivanov,
1938; Chapskii, 1955, 1963) (Fig. 137). The body length from tip
of nose to tip of tail is 110-142 cm in a straight line. The

222

223

294

Fig. 137. Teeth of the Baikal seal, anterior view (figure by N.N. Kondakov).

largest male measured was 141 cm long and the female 142 cm.
The length differences between sexes is insignificant. The weight of
the adults is 45-55 kg but some individual animals can reach 100 kg.
The weight of the blubber is 25-29 kg. The internal organs weigh
(g) on average: heart 300-400 (heart index 6.66 - 7.27), liver 900 - 1400,
lungs 400-500, kidneys 300, spleen 400, stomach 600, and intestine
400.

The condylobasal length of the skull is 173.5-204.4 mm, width at
zygoma 108.3 - 121.2 mm, and maximum width of skull 95.8- 102.9 mm
(Ivanov, 1938). Age-related changes in the structure of the skull are as
follows.

1. The cranial capsule of the young seal is relatively larger than that
of an aged seal. Measured from the posterior edge of the orbits (along the
posterior edge of the interorbital constriction) to the posterior edge of
the occipital bone, its length is roughly equal to the entire remaining half
of the skull, from the posterio edge of the orbit to the anterior section of
the intermaxillary bones. Among adults, the length of the cranial capsule
is roughly equal to the distance from the posterior edge of the orbits to
the end of the nasals.

2. The central portion of the nasal bones among young seals is con-
siderably wider than in adults. The width of these bones is usually notice-
ably more than the longitudinal diameter of the alveolus of each of the
upper canines.

3. The bony lobes of the lower outer edges of the auditory meatus
among young seals are relatively wider than in adults (Qgnev, 1935).

A small number of seals of very small dimensions, some with pre-
dominantly ocherous shades, have been encountered. It is suggested
that these do not represent individual variations but “starvelings,” 1.е.,
pups which have lost their mothers in the period of lactation but yet
somehow managed to survive; they are termed dwarfs (Ivanov, 1938).
(V.A.)

295
Taxonomy

The Baikal seal represents an independent advanced species of a group
that is well adapted to living in fresh water.

In craniological features the Baikal seal is closely related to the
Caspian but its evolution is evidently not directly related to the lat-
ter. There are two hypotheses of the evolution of the Baikal seal: its
entry into the Baikal during the Quaternary along the rivers of Yenisey-
Angarsk or the Lena system from the Polar Basin, and the arrival of the
ancestors of this seal in the pre-Quaternary period from the Sarmatsk-
Pontichesk Basin. The first of these hypotheses is more widely accepted
and has been confirmed by fairly weighty arguments from helminthol-
ogists. The louse parasitizing the Baikal seal belongs to the species
Echinophthirius horridus, found on the seals of the northern seas, while
the nematode, Contracaecum osculatum, detected in the Baikal seal is a
widely distributed parasite of the seals inhabiting polar waters. Evidently
the view of the northern origin of the Baikal seal is more correct (Ass,
1935; Mozgovoi and Ryzhikov, 1950; Chapskii, 1963; Lomakin, 1964).
(V.A.)

Geographic Distribution

These seals are not found outside the boundaries of Lake Baikal. They
inhabit mostly the northern half of the lake where extremely significant
beach rookeries are formed in the summer months. In winter they are
distributed more uniformly, but are fewer in the southern half of the.
lake (Ivanov, 1938; Pastukhov, 1961). (V.A.)

Geographic Variation

Not reported. (V.A.)
Biology

Population. In the spring of 1953, an aerial survey was made of the seals
resting on ice floes and of their air holes. The data recorded for the
surveyed sections were extrapolated to the entire area of the lake, taking
into account the density of disposition of the animals. The total pop-
ulation was thus determined at 20,000-25,000 (Sviridov, 1954). On the
basis of a census of the seals on ice floes in autumn in the main regions
of the concentration of the animal together with the assumed number of
seals in the other (less important) sites of autumn concentrations, the

224

224

296

Fig. 138. Haunt of seals on the ice. Northern Baikal (photograph by V.D. Pastukhov).

population later was put at 35,000 - 40,000 (Pastukhov, 1967). All these
figures should be regarded as approximate.

Habitat. In the winter months the seals inhabit mainly floating
ice under a continuous ice sheet, using air holes in the hummocky
ice floes for respiration. From the time of thawing of the ice floes
to their total disappearance, the animals form rookeries on the ice
(Fig. 138). In summer and autumn, beach rookeries are found mainly
on the northeastern coast of St. Nos and Ushkan’ Islands in the south.
Some “extinct” (Ivanov, 1938) rookeries which were regularly inhabited

297)

sometime in the past, are seen in the southern part of the Baikal. The
summer rookeries are distributed on the coastal flat stones jutting out
of the water, slightly away from the coast.

Food. Examination of the stomach and intestinal tract of some 500
seals provided information about their food for almost the entire year
in different parts of the Baikal (Pastukhov, 1965b, 1967). Most of the
stomachs of the Baikal seal investigated were found to be empty, which
is characteristic of the other species of Pinnipedia too. Semidigested
food remains were detected only in three stomachs but the intestine
and especially the rectum invariably contained the remnants that were
difficult to digest (otoliths, eye lenses, etc.). These provided a basis for
identifying the species of fishes consumed (Table 13).

In addition to the 17 species of fishes listed in Table 13, the food of
the Baikal seal included some species of invertebrates: gammarids of the
genera Odontogammarus, Macrochectopus branickii (pelagic), Garjajewia
(deep water), and Acanthogammarus. Mollusks of the genus Baicalia
were detected twice. Sand, pebbles, and mica were found quite often in
the alimentary canal.

Table 13. Species composition of fishes consumed by the Baikal seal (Pastukhov, 1956b)

Species of fish No. of seals Frequency,
in which this %
species was
detected
Baikal omul, Coregonus autumnalis migratorius Georgi 8 6.6
Stone sculpin, Paracottus kneri Dyb. 3 2:5
Sand sculpin, P. kessleri Dyb. 22 18.0
Big-heated sculpin, Batrachocottus baicalensis Dyb. 3 2.5
Spotfin Baikal sculpin, В. multiradiatus Berg. 4 3.3
Yellowfin Baikal sculpin, Cottocomephorus grewigki Dyb. 70 57.4
Longfin Baikal sculpin, C. inermis Jakowl. 70 57.4
Red Baikal sculpin, Procottus jettelesi Dyb. 9 7.4
Humped sculpin, Asprocottus megalops Gratzian 73 1.6
Big red Baikal sculpin, Procottus jettelesi major Tal. 1 0.8
Big Baikal oil-fish (big golomanka), Comephorus 60 49.2
baicalensis Pall.
Lesser Baikal oil-fish (lesser golomanka), C. dybowskii 92 75.4
Korotn.
Common perch, Perca fluviatilis L. 5 4.1
Burbot, Lota lota L. 2 1.6
Ide, Leuciscus idus L. 1 0.8
Siberian dace, Leuciscus leuciscus baicalensis Dyb. 1 0.8
Roach, Кии; rutilus Pall. 6 4.9

225

226

298

Throughout the spring and summer the food of the seal evidently
remains the same and is restricted mainly to four species of fish: yel-
lowfin Baikal sculpin, longfin Baikal sculpin, big Baikal oil-fish, and
lesser Baikal oil-fish. “Other species of sculpin” occupy a very insignif-
icant position. The autumn food of the seal is somewhat more diverse
though the number of predominantly consumed fish species even at this
time does not exceed eight. The list of fishes consumed by the seal in
winter is similarly restricted to four species. Some food differences are
seen in different regions of Baikal but since the list of food items itself
is small, the differences too are not large.

Gammarids were found in the stomach of most of the seals in. the
year of their birth. The seals take to fish with advancing age. There are
no other convincing data on age-related food variations.

The number of commercial fishes consumed by the Baikal seal
(mainly in the autumn) is so insignificant that this animal cannot be
regarded as a threat to the Baikal fishing industry. The main food consists
of the Baikal oil-fish and goby, which are of little use to man (Pastukhov,
1965b).

Home range. The Baikal seals have distinct home ranges only in the
winter since most of them live close to air holes at this time (Fig. 139).
Barren females, immature seals of both sexes, and adult males live singly
around one or several air holes; the animals sometimes use common
air holes. The females and the newborn during lactation have their own
home ranges among caves under the snow.

Hideouts and shelters. Holes in the ice floe can be regarded as such
structures only by stretching the interpretation. The simplest of them are
the air holes of barren females, adult males, and immature seals. These
are in the form of a cone enlarging downward and ending in a small
opening at the top. When making the air holes, the seals quite often
take advantage of the natural openings formed during the hummocking
of ice floes and keep them open throughout the winter. In other cases
the animal has to make such holes by breaking the ice when it is still
thin or even “biting” it with their teeth and scratching it with their claws.

The nesting holes of the female represent genuine shelters. They
consist of a large opening 40-80 cm in diameter in an ice floe through
which the female can freely emerge from the water and the lair. The latter
is made in the snowdrifts adjoining the hummocks and is invariably on
the leeward side of the prevailing winds. The lair is in the form of a
snow cave with its arch and walls covered with an ice crust formed by
the respiration of the animals. The ice cover imparts adequate strength to
the cave. The smallest of the lairs measured was 47 cm high, 110 cm long,
and 108 cm wide (Ivanov, 1938). Some were much larger. Sometimes the

225

299

Fig. 139. Air hole in an ice floe made by an adult male Baikal seal (photograph
by V.D. Pastukhov).

roof of the lair thaws and an air hole is formed. The female undergoes
parturition in the lair and suckles the pup; molting of the juvenile hair
coat of the pup is probably completed in the lair also.

Daily activity and behavior. Data on these aspects are extremely scant.
In winter, from the moment of ice formation, most of the seals spend
continuous time in water, rising in the air holes only for respiration,
hence it is impossible to study their behavior.

From early April the seals form spring rookeries on the snow. The
first to arrive are the immature animals (one-, two-, and three-year-olds).
Sometimes some mature males are also seen among them. The en masse
emergence of pups on the ice occurs in mid-April, at which time the
changeover from suckling to independent feeding takes place. The next
to arrive are the barren females and adult males, followed finally by
mothers that have undergone parturition in the current year.

227

300

The seals bask in the sun in different postures: on the back, side-
ways, and on the belly. A characteristic posture of nearly all the animals
of every age is the head turned toward the air hole and in its immediate
proximity. Initially the animals are extremely restless and rapidly “disap-
pear” in the water at the slightest hint of danger. A “reassured” animal
is quieter but often raises its head and may suddenly disappear in the
water at any moment.

The summer rookeries are almost exclusively colonized by the
females. The animal population in the rookery changes round the clock
somewhat systematically. The seals are usually not found in the rookery
until 7.00 a.m. They start approaching from 7:00-8:00 a.m. and arrive in
the lair at 8:00-9:00 a.m. Later, the number of seals increases gradually
and reaches maximum by 11:00 a.m. The number then begins to decrease
and approaches minimum by 1:00-2:00 p.m., but once again increases
by 3:00 to 4:00 pm. Maximum numbers are seen at 6:00 p.m. but by 8:00
to 9:00 p.m. the rookery is almost empty. The periodic departures of
the seals for feeding explain these population variations in the rookery
(Ivanov, 1938).

In spring the seals feed more intensively at night and in the early
hours of the day (the stomach of seals caught toward evening never
contained food remnants while only some caught in the morning had
an empty stomach). The period of intensive feeding exhibits seasonal
variations (Pastukhov, 1965b).

Seasonal migrations and transgressions. The Baikal seals perform reg-
ular but relatively small migrations in the Baikal water body. Whelped
and barren females and part of the mature males pass the winter on the
eastern coasts while the immature seals and part of the adult males do
so on the western coasts. Early in spring large (“‘polyn’yas”) air holes are
formed in the ice at the same places every year along the eastern and
western shores of the Baikal. The first of the spring rookeries of seals
are formed along these shores. With the formation of a large number of
fissures and tears close to the western shore, a large number of mature
males and barren females migrate there from the eastern shores and, as
a result, fairly large congregations of seals are formed at times on the
western shores.

At the end of May, in the southern half of the lake, the ice begins
to break up and floating ice floes are gradually carried northward by the
prevailing southwestern winds. Along with these drifting ice floes, almost
the entire southern herd of seals reaches the north and forms numerous
rookeries there on them. Beach rookeries are formed after the ice floes
thaw. Only a small number of seals, regarded as “locals”, remain in the
southern part of the lake.

228

301

The beach rookeries begin to break up early in autumn. The seals
finally abandon them by September-October and concentrate on the east-
ern shore of the lake in Proval, Barguzinsk, and Chivyrkuisk bays, and in
the estuarine sections of the Upper Angara, i.e., along much of the east-
ern shore. Such rookeries are not seen only in the extreme southeastern
part of the shore. All of these shallow-water regions are protected from
the autumn winds and hence covered with ice floes earlier than in other
zones. As soon as the shore ice is strong enough to support the weight
of the animals, the seals form sometimes fairly large (hundreds of ani-
mals) rookeries on it. Animals of both sexes and of different age groups
are seen in the rookeries, which remain in force for 1.5 to 3 months
depending on the time of total freezing.

As the ice zone enlarges in the shallow-water regions, the seals hold
onto its edges and gradually move toward the unfrozen deeper part of the
lake; not a single animal remains in the shallow-water zone in November-
December (period of total freezing). The edge of the ice zone, running
comparatively close to the eastern shore, is regularly split by wind and
waves and the broken floes enter the open Baikal carrying the seals with `
them. Here the animals no longer form large groups but divide into
small ones spread over a large area. From the time of the final freezing
of the lake, the seals take to living under the ice until the lake opens up
in May. Spring hunting is concentrated mainly in these regions (Ivanov,
1938; Pastukhov, 1961). Thus the Baikal seal performs regular though.
short-distance seasonal migrations.

Some stray transgressions of the Baikal seal occur in the rivers enter-
ing or emerging from the Baikal, Salenga to Selenzhinsk (280 km) and
even up to Ust’-Kyakhty (400 km), Barguzin to Ust’-Barguzin, and down-
ward along Angara to Irkutsk (not exactly) and up to Olonka village
(150 km) (Ivanov, 1938; Lomakin, 1964).

Reproduction. No direct observations of the mating of the Baikal seal
have been reported. Presumably, mating occurs in water under the ice
cover at March end to the first half of April, soon after the birth of pups
(Pastukhov, 1966). Most of the females undergo parturition in February
and March and only some stray births occur earlier or later. Even at the
beginning of April, females in the last stages of gestation or newborn
pups (with the umbilical cord not yet dry) are very rare. Thus the pups
are born while the females are still on stationary ice floes. Usually, the
female undergoes parturition in a snow cave (see above). The entire
duration of whelping extends for 30-40 days.

A study of the embryos and an analysis of the sex ratio of the seals
caught showed that the male to female ratio is close to 1:1 (Ivanov,
1938).

229

302

А study of the female genitalia revealed that the period of sexual
maturity and reproductive capability extends to several years. At the age
of 2-5 years (from the state of the genitalia), the females become poten-
tially mature but do not actually participate in reproduction. At the age
of 3-6 years they attain productive maturity and the first fertilization has
been observed among the four-to-seven-year-old females. All the seven-
year-olds and older females investigated were already mothers. Later,
they whelp almost every year since barrenness affects only 12% of them.
Gestation extends for 11 months, of which the first three represent the
latent period of fetal growth (Pastukhov, 1966, 1969a).

Growth, development, and molt. The pups of the Baikal seal are very
large at birth and measure one-half the length of the mother (Fig. 140).
Lactation extends for 2-2.5 months and the pups grow very rapidly. Of
the eight pups weighed in April (1-1.5 months after birth), the smallest
weighed 22 kg and the largest 44 kg; average weight 31.3 kg (Ivanov,
1938).

The newborn pup is covered with a dense yellowish-white hair coat
3-4 cm long. In 1.5-2 months, quite often even during the period of res-
idence in ice caves, i.e., from February end to early April, the embryonic
hair coat is completely shed and the pup acquires the color of the adult
(Fig. 141). The molting period of the adults is considerably prolonged

227

Fig. 140. Newborn Baikal seal pup (photograph by V.D. Pastukhov).

303

and its duration depends to some extent on the well-being of the animal.
Some seals begin to molt even in early May. At May end and in early
June, the number of molting seals is already significant but molting ani-
mals may be encountered throughout June and even up to mid-August.
It is assumed that the better fed the herd in a given year, the earlier
the ouset of molt; the faster molt proceeds, the sooner it is completed
(Ivanov, 1938).

Enemies, diseases, parasites, mortality, and competitors. This herd suf-
fers the most damage from bears, which prowl the beach rookeries. There
are no other enemies.

Among the ectoparasites, the louse Echinophthirius horridus (Olfers,
1816) Fahrenholz, also known among other true seals, Steller’s sea lion,
and fur seals, has been reported on the skin of these seals. This parasite
of the Baikal seal has been isolated as a distinct subspecies, E. horridus
var. baicalensis Ass.

The helminth fauna is represented by a single species of nematode,
Contracaecum osculatum (Rud., 1802) Baylis. A distinct subspecies of this
species, C. o. baicalensis Mosgovoy and Ryjikov (Ass, 1936; Delyamure,
1955), is present among Baikal seals.

The diseases and causes of mortality of the Baikal seal have not
been studied. Probably, some seals perish every year in winter due to
unfavourable conditions of the Baikal ice crust.

228

Fig. 141. Molted pups of the Baikal seal (underyearlings); unmolted pup seen in
the foreground (photograph by V.D. Pastukhov).

230

304

Population dynamics. Noticeable natural changes in the population
of the Baikal seal have not been reported. The population varies only in
relation to the hunting activity. Further, the number of seals caught also
reflects to some extent the variation in their total population. Although
the total catch rose sometimes to 9,000- 10,000 per annum in the pre-
revolution years, hunting activity subsequently decreased. It again rose in
the 1930s when the total catch reached 6,000 - 6,500 per year. In subse-
quent years, catches steadily decreased, which is explained by a reduction
in total population. This downward trend continued and the catch never
exceeded 1,500 per annum. The population dynamics were strikingly
dependent on hunting, which ultimately led to a significant reduction
of the entire seal population (Pastukhov, 1965a).

Field characteristics. The Baikal seal is characterized by a monochro-
matic fur free of spots. It may be further noted that no other species of
seals exists in the Baikal region. (V.A.)

Economic Importance

Hunting data in the pre-revolution period are scant and contradictory;
the annual catch of seals has been put at 3,500 to 9,000- 10,000 by var-
ious authorities. Evidently the significant proportion of the catch fully
utilized by the hunters themselves has not always been included in catch
records. At the end of the 1920s, the annual catch was 3,500 - 4,000 seals.
Commencing from 1931, the volume of hunting rose and comprised an
average of 5,700 for six years (maximum 6,466); later hunting gradually
declined. From 1940 through 1950, the average annual catch was 1,484
seals but fell to a mere 847 from 1950 through 1960. The annual catch
in this period never exceeded 1,500 seals. It should be borne in mind
that a certain quantum of catch does not always figure in the accounts
as it is used privately; this figure in some years could be almost equal
to the official figure. The actual annual catch [in this decade] was about
2,000 (Pastukhov, 1967, 1969b).

The Baikal seal is of no importance except to the local people for
whom its hunting plays а growirig role with increased production of raw
furs.

The technique of catching these seals is as typical as it is diverse.
Formerly, hunting was carried out year round but experience showed
that successful hunting on the spring ice floes accounted for almost all
of the annual catch. Therefore, from 1935, hunting in the summer and
autumn beach rookeries was banned and later, hunting in the southern
half of the Baikal was also prohibited. These measures did not reduce the
annual kill. Subsequently, hunting from boats was banned. Thereafter,

231

305

hunting on sledges became the main method and the hunting season
declared open from April 25. A group of some 20 hunters with horses
harnessed to sledges set up a hunting camp on a Baikal ice floe. Usu-
ally each hunter has his own horse-drawn sledge but sometimes two
hunters use the same sledge. Early in the morning, the hunters set out
in different directions in search of the seals. On sighting resting seals
through binoculars, the hunters leave their horses 2-2.5 km away from
the animals. Then, sporting little white caps and pushing their small
sledges adorned with white sails in front of them, the hunters endeavor
to approach as close as possible to the seals (Fig. 142). The sail has two
openings, one for observation and the other for shooting through. Con-
cealment against the sun and invariably against the wind is also necessary
as Otherwise a seal can sense a hunter’s presence far beyond the shooting
Tange.

The Baikal seal is so sensitive that, even under favorable conditions,
it is impossible to approach it within less than a hundred meters. Having
come within the required range, the hunter shoots the seal with a rifle
and then runs headlong toward it with a hook since even a mortally
injured animal can, with its dying breath, still dive into the water; the
hunter has to catch it on the ice. In a successful hunt the horse sledge
is brought to the kill, the seal loaded on it, and the chase for the next
animal begun. The hunters return to camp in the evening with one or
two and sometimes even as many as ten seals.

230 Fig. 142. Device for concealment from seals. Baikal (photograph by V.D. Pastukhov).

306

When the ice is thin and dangerous for sledges, the hunters set out
on foot, which greatly reduces the hunting range. However, even when
using sledges, some hunters operate on foot (Ivanov, 1938).

Hunting by means of boats (now banned) used to commence after the
ice floes had broken up. The hunters negotiated the channels using small
boats with the prow masked by a white sail. On sighting an animal, the
hunter turned the boat toward it so that the vessel remained concealed
behind the sail. The animal was shot with a rifle, the kill loaded on the
boat, and brought to the camp in the evening, where all the hunters
gathered.

In recent years, hunting by sledges has undergone some mechaniza-
tion. In the 1960s, hunting on foot almost ceased and most of the hunters
chased the seals on horseback but some began using light motorcycles
for towing the hunting sledges. Finally, in 1969, any type of hunting of
the adult animal was totally banned. At present, the use of rifles is per-
mitted on the spring ice from April 25 for shooting seals only below the
age of one year.

Much effort has been expended in the preservation and restoration
of the population of the Baikal seal. Gradually, hunting in the summer,
autumn, and winter, and in the southern half of Lake Baikal, as well as
of all adult animals was prohibited. Finally, a norm was fixed for the
annual kill. The most effective measure has been the banning of killing
of adults and allowing the catch of only pups.

Hunting of unmolted pups in snow caves, where they shed the embry-
onic pelage, is practically impossible. It is therefore recommended that
hunting of molted underyearlings be permitted. These are far bigger
than the unmolted pups (average weight 20 kg and weight of blub-
ber 12 kg) and provide a no less valuable fur. Shooting with rifles
could then be totally prohibited as this results in inevitable losses in
the form of injured animals and damaged fur. The young seals should
be caught in nets laid under the ice. The right time for catching them in
this manner is the end of April (not before April 25) since by then
all the pups have molted and lactation has ceased. Test catching in
nets has shown that adults are not trapped in them (Pastukhov, 1967,
1969b).

At the present level of population, the annual kill can be set in the
range of 2,000-3,000 with no loss to the herd (approved limit 2,500).
This change in the system of utilizing the herd can promote its popula-
tion rise. The organization of census work can provide a base for rational
regulation of hunting. A catch of young “fur-bearing” animals is far more
economical than hunting for “skinny” adults. (V.A.)

232

1758.
1811.
1811.
1820.
1823.

1824.
1828.

1828.
1828.
1844.
1864.

1902.

1902.

1902.

1902.

1936.

1941.

307

Subgenus of True Seals
Subgenus Phoca Linnaeus, 1758

COMMON SEAL, LARGA®
Phoca (Phoca) vitulina Linnaeus, 1758

Phoca vitulina. Linnaeus. Syst. Nat. Ed. X, 1:38. Northern part of
the Baltic Sea.

Phoca largha. Pallas. Zoogr. Rosso-Asiatica, 113. Eastern coast
of Kamchatka.

Phoca canina. Pallas. Zoogr. Rosso-Asiatica, 1:114. Atlantic (?).
Phoca variagata. Nilsson. Skand. Fauna, 1:359. Atlantic (?).
Phoca scopulicola. Thienemann. Reise im Norden Europas, 1:59.
Iceland.

Phoca littorea. Thienemann. Jbid. Northern Russia (?).

Phoca linnaei. Lesson. Diction. classique d’Histoire Natur.,
13:415. European waters.

Phoca thienemannii. Lesson. Ibid., 13:414. New name for Phoca
scopulicola Thienem.

Phoca chorisi. Lesson. Ibid., 13:417. Kamchatka.

Phoca nummularis. Temminck. Fauna Japon, р. 3. Japan.°’
Halicyon richardii. Gray. Proc. Zoolog. Soc. Lond., р. 28. Van-
couver Island.

Phoca ochotensis. J. Allen. Bull. Amer. Mus. Nat. Hist., 16:480.
Gizhiga Estuary, Sea of Okhotsk. Nec Phoca ochotensis Pallas,
1811. |

Phoca ochotensis macrodens. J. Allen. Ibid., 16, p. 483. Avachinsk
Bay, Kamchatka.

Phoca steinegeri. J. Allen. Ibid., 16, р. 485. Commander Islands
(Bering Island).

Phoca richardi pribilofensis. J. Allen. Ibid., 16, р. 495. St. Paul
Island, Pribilov Islands.

Phoca vitulina largha natio pallasit. $. Naumov and М. Smirnov.
Tr. Vses. n-i. in-ta rybnogo khozyaistva i okeanografii (VNIRO),
3:177. Sea of Okhotsk.

Phoca petersi. Mohr. Zoolog. Anzeiger, 133, р. 49. Coasts of the
Korean Peninsula.

Also mottled seal (Far East), rock seal (sometimes in Murman).

67 According to King (1961), this form has been described by Temminck from material
of the ringed seal, Phoca (Pusa) hispida Schreb., and hence should be excluded from the
synonyms of Phoca (Phoca) vitulina.

233

308

1942. Phoca ochotensis kurilensis. Jnukai. Shokobutsu Dobutsu, 10,
no. 10, p. 930. Southern Kuril Islands.

1964. Phoca insularis. Belkin. Dokl. AN SSSR, 158, no. 5, p. 1217.
Iturup Islands (Kuril Islands) Cape Dokuchaev. (V.H.)

Diagnosis

The larga is a large, well-proportioned seal. The body length measured
along the dorsal surface (Lc) exceeds 1.5 m; the neck and the snout are
somewhat elongated. The color of the hair coat varies from a bright mot-
tled and contrasting (with a predominance of light-colored background,
speckled with small gray and black spots) to intensely dark color with
Clear spaces in the form of oval rings or streaks (Fig. 143).

The skull is relatively massive and its length not less than 190 mm;
the interorbital width usually exceeds at least 1.5 times, and the total
width of the nasal bones at the base of their apex (along the frontal-
maxillary suture) at least two times the smallest diameter of the sub-
orbital aperture. The longitudinal width of the crown of the second to
the fourth upper premolars usually exceeds the height of the crown. The
accessory cusps of the corresponding lower teeth are short, usually less
wide-set, and close to the main cusp as though adjoining it (Fig. 144).
(K.Ch.)

Description®

In terms of body proportions, the seals of this species can be regarded as
typical of the genus. The fore flippers are relatively small, somewhat
shorter than the hind flippers. The third digit on the fore flippers
together with its claw is shorter than the second and the first while
the second is often somewhat longer than the first or equal to it. The
whiskers are fairly well flattened, with wavy edges, and set in six rows
(only one whisker in the seventh row) with the maximum number (8
or 9) in the second and third rows from below; the total number of
whiskers on each side of the snout varies from 39 to 48. The supraorbital
whiskers number five each, rarely four each; one whisker each occurs
near the nostrils. The nares are bordered by a narrow strip of bare
skin.

The color of the hair coat after the first (infantile) molt exhibits
extreme variation not only individually, but also in relation to the

68 The larga described here is the arctic Far Eastern form (for more details, see
“Geographic Variation”).

309

rh a WN

Rey
ays

BS SE 2
7... «т
РАЯ 9S, А 72 5 ith vi
727. 4 О ДАН Е
ies SRLS rvs Moy =
Saez И РА erg TAL oe
: .. ПАСЕ 7. -.

ff О >
Л.

2 bo
A рее _

=>

17755. = я и . :-
=~ SALE Fe “8 й 5

=== Е
т Me eee Se SG

233

Br nth

A ` и Е ИА: я г iu
к Seine ed’ tins. Gon
haa Qe Apelor

у

235 Fig. 144. Skull of the common seal, Phoca у. vitulina (figure by М.М. Kondakov).

234

310

geographic distribution of the population and its affinity to the
pagophilic or pagophobic group.

These seals can be divided into two main types—very dark and very
light-colored. The dark type is most often encountered in the group of
Atlantic common seals and also among the pagophobic™ populations of
the Pacific Ocean part of the range. The light-colored type is common
among the pagophilic Pacific populations, 1.е., the larga.

Among the dark-colored animals, the main background is either
very highly pigmented or very densely covered with additional, very large
spots. One way or the other, there is little space for the light-colored
sections on the dorsal side of the body. Such sections are seen sometimes
only in the form of narrow and sinuous light-colored streaks, more often
as short, as if broken streaks in the form of light-colored dabs on a dark-
colored background or as indistinct rings. Dark-colored animals are also
encountered with large contrasting rings and an abundance of minute
ringlets interspersed with spots of diverse shapes, sizes, and colors (usu-
ally with an indistinct contour and often superposed on each other).
These are colored different shades of brown-olive, cinnamon, dark gray,
and black. The ventral side of such specimens is also usually dark-colored
although the color of the spots on it is somewhat fainter (see Plate III).

The main color of the light-colored animals, on the contrary, forms
a light-colored background on which dark gray spots admixed with black
ones are scattered quite densely. The black spots appear as though super-
posed on the dark gray spots and rarely as dabs. This significant diversity
of color acquires a very definite and distinct character in the background
of geographic variation (See pp. 323 -330).

The age-related color changes have not been adequately traced
(Millais, 1904; Havinga, 1933; and others) due to extensive individual
variation of all the elements constituting the skin pattern. Among the
under-yearlings of the European common seal, a broad, dark, and more
monochromatic (brownish) band extends along the middle of the back
from the head to the tail. This band is only slightly interrupted by dark-
colored specks and gaps which are usually smaller than those among
older animals. This is also a characteristic of the Pacific Ocean forms,
including the larga. The ventral side is silvery-white, with extremely
rare spots in most cases. Sometimes, however, even young animals are
encountered with innumerable spots on the belly, as noticed among the
pagophobic form (Fisher, 1952) and the arctic larga (Chapskii, 1967).
The spots on the body flanks are more diverse in number, brightness, and
configuration. The color of the head is lighter than among older animals.

6 Breeding not associated with ice floes.

(лохериох ‘мм
Aq ainsi) (1еэл эц1 ип) ‘Teg 2840] вийпил (0э20ца) DIOY_ ‘eBIe] рие чехии] 55иэйииху вицпил (DIOY_) DIOY_ Чеэ$ плау 1O pues]

Ш эа

we

311

With age, the dark-colored dorsal band gradually becomes mottled
with white gaps, often in the form of fairly distinct oval rings. As a result,
the median longitudinal band on the back, which is distinctly monochro-
matic among juveniles, ultimately disappears altogether. Further, such
a process of age-related color changes is evidently not seen among all
underyearlings. Animals are encountered among them (Moore, 1955*;
Heinroth, 1956*; and others), which are almost indistinguishable from
the adults either in differences in mottled clear spaces medially on the
back, or in the development of innumerable light-colored oval rings on
the dorsal side of the body, or in the color of the head.

Young animals, which have yet to reach adult size, are generally sim-
ilar in color to the latter but nevertheless differ in a more monochro-
matic color of the back (Ognev, 1935). Fully adult and older animals
have a more vivid and contrasting spottiness in which the dark and light
components of coloration are fairly evenly represented and uniformly
alternated, creating a bright mottled, speckled pattern usually lost on
the light-colored ventral side. This is relevant, however, only to a def-
inite type of spottiness that is especially characteristic of the Atlantic
forms and the arctic larga. Coloration is highly diverse and very light-
colored animals are also encountered together with brightly mottled and
very dark, sometimes almost wholly black animals. The dry skin of the
light-colored animals appears almost white from a distance.

The hair coat of the under-yearlings is softer, somewhat denser, with
a better developed “layer” of thin, tender, and extremely short hairs,
almost like underfur. The coat of adults is considerably coarser; the seta-
ceous guard hair (base of the coat) is thicker and dominates sharply over
the underfur, which is difficult to distinguish among this coarse guard
hair (Havinga, 1933; Fisher, 1952).

The color differences between males and females have not been thor-
oughly distinguished; some regard the spotted pattern as more intensely
manifest among males (Havinga, 1933); others (Millais, 1904) consider
the back of the female to have more dark spots while the ventral side is
covered less densely with spots than in the male and the color of females
is therefore considerably paler. The ringed pattern is better developed
among male largas than in females and the color in general is brighter
and the pattern better contrasted (Chapskii, 1967).

Information on the seasonal color variation requires verification. It
was pointed out (Millais, 1914*) that before every molt the general color
shade, as well as the intensity of spots, turns much lighter and duller as
though faded. According to other authors (Havinga, 1933), the color of
the hair coat turns a dirty yellow before molt.

235

236

312

Two main types are noticed in the structure of the skull (Fig. 144)
as also in the color of the hair coat. These types are associated with
the ecological and taxonomic grouping of the species into two forms
as pagophilic and pagophobic.’? Among the animals of the pagophobic
form represented in the Atlantic Ocean, the nasal processes of the
maxillary bones usually do not reach the nasal bones or sometimes only
slightly contact the latter without wedging deeply posteriorly between
their outer edge and the maxillae. The length of the anterior part
(rostrum) of the nasal bones in most cases somewhat exceeds one-half
their total length. The uncinate processes of the pterygoid bones are in
the form of club-shaped thickenings or hooks slightly flattened laterally
and not bent outwardly. The bony nasal septum in the choanae does
not extend posteriorly farther than the anterior edge of the palatine
bones. The posterior edge of the bony palate is distinctly notched,
often with slightly curved sides and usually with an additional angular
notch medially. The occipital foramen [foramen magnum] usually has a
high angular notch on the upper edge. The premolars, except the first,
are large; the second and the third, with rare exceptions, are disposed
obliquely in relation to the tooth row in such a way that the anterior edge
of the last tooth runs somewhat inward beyond the posterior edge of the
preceding tooth. The anterior articular edge of the zygomatic bones is
somewhat longer than the total of their posterior edge measured between
the most prominent points.

The body length of adult seals’! caught on the Dutch coasts and
measured in a Straight line (Lcv) was around 150 cm in most cases, the
largest of them reaching 160-165 cm” (Havinga, 1933). The maximum
length evidently not measured in a straight line, but along the dorsal
surface (Lc) among the Norwegian seals is 180 cm (Collett, 1911 - 1912).
On the German coasts the length of the largest animals (measured
evidently along the dorsal curvature) does not exceed 175 cm (Moore,
1955*).

According to one view in the literature, the Pacific Ocean seals of
this species are larger than their Atlantic counterparts (N. Smirnov, 1929;
Ognev, 1935; Bobrinskii, 1944*, 1965*). Such an assumption, however,
is justified only on an indiscriminate comparison with the latter of all

The skull description is based on specimens from the North and Barents seas (nominal
form). For the skull characteristics of the Far Eastern form, see under “Geographic
Variation”.

71 Various authors cite the sizes of seals in their own way and do not always clearly state
how the animal was measured—in a straight line or along the dorsal body curvature.

72 The values cited by Havinga expressing the body length along a straight line but up
to the tip of the hind flippers have been converted using his own conversion factor (13%).

313

the seals belonging to the various Pacific Ocean forms, although their
taxonomic structure is quite varied. In fact, the pagophobic seals of the
Far East, for example from the Kuril Islands, are perceptibly larger: the
body length (Lc) of the largest animals is around 200 cm or even more
(Belkin, 1964). The larga, however, belongs to the pagophilic forms and
its maximum dimensions are no different from those of the Atlantic seal
at maximum length (Lc) 175 cm (S. Naumov, 1941) to 182 cm (Chapskii,
1967).

Under the conditions of intense oppression by man, the small size
of the European seal is perhaps wholly to be expected; at the same time,
natural factors too could have been responsible for the size differences.”

The length of the adult male larga averages 173 cm (Lc), of the
female 162 cm. Two very large female adults caught on the southern
coast of the North Sea weighed 76 and 105 kg; the maximum weight of
two males was 100 and 114 kg (Moore, 1955*). Two male Okhotsk largas
in the spring weighed 92.5 and 96 kg while an adult female also caught
there weighed 82.5 kg (Wilke, 1954); the weight of male Bering largas
in the spring reached 100 kg and that of females 88 kg.

The condylobasal length of the skull of adult animals of both sexes
from various parts of the range varies from 190 to 255 mm. Among adult
males, the skull on average is 10 mm longer than that of adult females.
The width at the zygomatic arches exceeds the width at the mastoids:
among adult largas, the former measured 110-138 mm and the latter
112-125 mm; the rostral width varied from 31-45 mm.

The relative weight indices of the internal organs among male largas
in the spring at an average body weight of 73.5 kg are as follows: heart
average 6.3%., lungs (with trachea and larynx) 13.3%., liver 26.2%.,
kidney 2%., and stomach 9.2%. (Sokolov, Kosygin and Tikhomirov,
1966). The length of the os penis among adults is 13.5-14.0 cm (for
more detailed body and skull dimensions, see “Geographic Variation”).
(K.Ch.)

Taxonomy

The seals: of the subgenus Phoca are usually compared with those of
the subgenus Pusa, and especially with the ringed seal (Phoca hispida),
from which they have inherited many features. However, the structure
of the cheek teeth reveals significant differences and these along with

3 Data on the Atlantic seals are extremely scant and fragmentary and there is no accurate
information on their actual ages. Moreover, the oldest arfd the largest rarely attract hunters.

237

314

some other differences probably justify the placement of these species in
different but nevertheless very close genera.

The taxonomic structure of this species in its broader interpretation,
as adopted in this publication, has been reviewed several times and even
today evidently cannot be regarded as conclusively established. In spite
of describing several forms from the Pacific Ocean (see under synonyms),
its taxonomic structure in the waters of the USSR has long led to two
subspecies: Atlantic—Phoca v. vitulina L. and Pacific—Phoca v. largha
Pall. (N. Smirnov, 1929; Ognev, 1935). However, the ecological popula-
tion and morphological heterogeneity of the Pacific population of the
common seal (Chapskii, 1960) detected after the 1930s necessitated a
review (see p. 158). As a result, the efforts of researchers attempting to
reflect the diversity of the Pacific Ocean seals by describing new species
and subspecies had to be evaluated from a new viewpoint. Of particular
interest at present are the attempts of Allen (1902) who described a new
species of seals from the Commander Islands (Ph. steinegeri) and Inukai
(1942) who pointed out that it be regarded as a special subspecies of the
larga (Phoca kurilensis). This tendency again intensified in the 1960s fol-
lowing the collection of extensive new data on the ecology, morphology,
and distribution of the Pacific Ocean seals. As a result, suggestions were
made to regard the Kuril “island” seal (Belkin, 1964; McLaren, 1966)
and the pagophilic form of the larga (Chapskii, 1966, 1967; McLaren,
1966) as independent species.

It is possible to interpret differently the ranks of these seals but
the following premises in any case are beyond doubt. Firstly, the Pacific
Ocean members of the subgenus Phoca s. str. cannot be regarded as
a single form whatever the level be, species or subspecies, assigned to
it. The ecological and morphological features of the larga are so sharp
that they run beyond the limits of even the so-called good subspecies.
Secondly, there is greater relative similarity between all the pagophobic
populations of the Pacific and Atlantic than between the Pacific Ocean
forms of different ecological types (i.e., between the pagophilic larga on
the one hand and the pagophobic, island or Kuril, seals, and Richard’s
seal on the other). As a result, the species of the common seal, Phoca
vitulina, according to some authors (Chapskii, 1966a, b, 1967; McLaren,
1966) should be divided into two species: 1) common seal—Phoca vit-
ulina L., and 2) larga—Phoca largha Pall. The composition of the second
species (larga) thus does not include the pagophobic Pacific seals, possi-
bly deserving in turn the rank of an independent species. In the present
work, however, according to the note on p. 158 and the general attempt
to resist extreme division of the species, the larga has not been regarded
as an independent species. (K.Ch.)

239

315
Geographic Distribution

This includes the waters of the continental shelf of the temperate and
subarctic belts of the Atlantic and Pacific oceans with the congruent lim-
ited regions of the North Arctic Ocean. The overall range, interrupted by
the arctic seas and land fringes of Asia and America is distinctly divided
into two isolated sections: North Atlantic and North Pacific. These rep-
resent a typical-example of interrupted amphiboreal distribution.

Geographic Range in the USSR

Commensurate with the above character of overall distribution, the seals
belonging to this group inhabit, on the one hand, our western marine
boundaries and, on the other, the Far East (Fig. 145).

On the western coasts of the USSR the Atlantic seal is encountered
almost only in the Murman region in the east up to the inlet and in rare
cases up to the isthmus of the White Sea. It is not found in the central
basin and the White Sea bays. In the middle of the last century (Ber,
1862*), it was reported in Novaya Zemlya but no one found it there
subsequently.

In our territorial waters of the Baltic, the distribution has not been
established with certainty either on the southwestern continental coasts
of the Estonian Soviet Socialist Republic, on the Khiuma and Sarema
islands (Aul, Ling, and Paaver, 1957), or in the coastal waters of the Lat-
vian and Lithuanian Soviet Socialist Republics. References to the rather
frequent encounter of this seal on the Baltic coasts including even Esto-
nia (Lewis, 1885*; Grosse and Transehe, 1929) should be regarded as
erroneous. In general the correct view of the distribution of the com-
mon seal in the Baltic Sea was established even in the 1930s, when it
was regarded as an inhabitant of only the western and southern fringes
of the sea (Freund, 1933). However, in the adjoining parts of the sea,
especially in Gdan’sk Bay, the common seal was without doubt encoun-
tered in the recent past (Ropelewskii, 1952). Its incidental find there-
fore is wholly possible on the coasts of Kaliningrad region although it
may not be a regular inhabitant there. Its presence even more north-
ward along the Lithuanian and the adjoining sections of the Latvian
coasts is also possible. There is no doubt of its absence in the Gulf of
Finland.

Thus in the USSR waters of the Atlantic portion of the range the
seals of this species are distributed almost exclusively at places where
there is no formation whatsoever of a stable ice crust. This pattern of
distribution wholly corresponds to the ecological nature of the European

` ~

“30

Fig. 145. Distribution of the common seal, Phoca vitulina, in the USSR (К.К. Chapskii).

238

317

populations of the species pertaining to the pagophobic form though
some animals and groups can be encountered on ice floes.”

In the USSR waters of the Far East the seal, essentially belonging
to a different ecological-taxonomic (pagophilic) form, is distributed very
widely from the USSR boundary with the People’s Democratic Republic
of Korea to the coasts of the Chukchi Peninsula and the southermost
part of the Chukchi Sea adjoining the Bering Strait.’> In the Sea of Japan
the larga’ is common in the Peter the Great Gulf and is encountered
north of it along the coast up to Tatar Strait where its concentrations
are considerable on the continental coasts, Sakhalin coasts, and on the
ice floes away from the coasts.

In the Sea of Okhotsk this seal is distributed all along its periphery
but unevenly and not uniformly in different seasons. In spite of adapta-
tion to breeding on ice floes, in winter the larga does not remain close
to the continental coasts and islands surrounded by dense, massive, gen-
erally stationary ice floes, but prefers instead the more pelagic regions
of the sea with drifting ice floes abounding in many pools of open water
and washed-out holes. The main regions in which the larga is concen-
trated in the snowy period of the year in the Sea of Okhotsk, accord-
ing to the latest data (Fedoseev, 1970; С.А. Fedoseev) are: the western
part of the sea (opposite the mid-northern part of Sakhalin Island and
also in Terpeniya Bay), northwestern coast of Kamchatka, in Yamsk and
Tauisk Bays, and in the region from Cape P’yagina to the Kola Решп-
sula. In summer, with the thawing of ice floes, the entire population of
the Okhotsk larga is wholly concentrated in the coastal belt of the sea.
The animals are not dispersed uniformly all over the territory, however,
but gather mostly in the regions that provide the most favorable food
conditions. The estuarine sections of innumerable rivers are extensively
used and the larga transgresses quite far from the estuary into many such
rivers. The uneven distribution becomes even more perceptible closer to
autumn as a result of seasonal concentrations of the population in cer-
tain fixed sites year after year where coastal rookeries are formed. They
are concentrated in three very important regions: (1) in the western part
of the sea (Shantarsk Islands), (2) in its northeastern fringe (including
Shelikhov Gulf and Taigonos Peninsula), and (3) in the coastal belt of

74 Three specimens of this species were caught in such an environment on the Murman
coasts, between Kil’din Island and Nokuev Bay, in the spring of 1901 (Smirnov, 1903).

75 The larga hardly spreads along the polar coast of Eastern Siberia into the west beyond
Long Strait although there are references (Rutilevskii, 1962) that it perhaps reaches even
the estuary of the Indigirka River.

76 For accurate definitions of “larga” and “Kuril or island seal,” see under “Geographic
Variation”.

240

318

western Kamchatka. In the first of these zones rookeries are formed on
the mainland coast where they are generally few, as well as on the islands.

In the western part of the Gulf of Sakhalin small rookeries and stray
haunts are encountered on individual reefs of Capes Litke and Mofet
(S. Naumov, 1941). Some rookeries are seen in Konstantin Bay (western-
most corner of Akademii Bay). A rookery exists in Nikolai Bay. The exis-
tence of rookeries has not been confirmed in Ul’bansk Bay although the
larga is found there in autumn. In Tugursk Bay rookeries are known in
Mamga and Kumchai bays and also near the isthmus separating Tugursk
Bay from Konstantin Bay. In the Shantarsk archipelago rookeries are
found exclusively on the small islands: Sivuch’i Kamni (northern and
southern), Utichii, Ptichii, Srednii, and on the reefs close to Bol’shoi
Shantar Island (Pikharev, 1941; S. Naumov, 1941).

In the region of Taigonos Peninsula, where the larga is preferentially
confined in summer to the estuary sections of the more southern rivers
of Penzhinsk and Gizhiginsk gulfs, beach rookeries were noticed in the
early 1930s mainly in the proximity of Capes Verkholamsk, Naklonnyi,
Krainii, Povorotnyi, and Pupyr’; on Tretii and Krainii, Khalpili, Rechnaya
Matuga, and Uikana islands; and also in Taigonos and Dorozh’ya bays.
Farther to the southwest, the larga was similarly distributed all over the
region and its rookeries were found in the same places year after year,
the most important of which are Cape Ostrovnaya, estuarine sections of
the Berezovka and Ireta rivers, and several points on the P’yagina and
Kona peninsulas (Freiman, 1936).

In Tauisk Bay rookeries were found mainly in Motyklei, Tokhar’,
and Stanyukovich bays, and on Nedorazumeniya Island; they were also
seen in Nagaeva Bay and on the open coast from it to Kholkhotlya
(Freiman, 1936). In the early 1960s, rookeries in Tauisk Bay were seen on
Capes Polonsk and Amakhtonsk, in Melkovodnoi Bay, and at some other
places (Tikhomirov, 1966). Recently, Babushkina, Ushki, Shel’tinga, and
Penzhinsk bays, and also sections on the western coast of Kamchatka
have also been cited (G.A. Fedoseev).

In the Tigil’sk region of western Kamchatka, right in the early
1930s, rookeries were detected (Lun’, 1936) at 10 places on the coast:
(1) 20-25 km north of Lesnaya River, (2) 10-13 km more to the north,
(3) оп Cape Kinkil’sk, (4) 40-50 km north of Amashino River, (5) on
Cape Babushkina, (6) on Tal’nichnaya Islet (15 to 20 km south of Cape
Babushkina), (7) near Moroshechnaya River, (8) near Cape Utkholoksk
(Kavachinsk rookery), (9) on Cape Yuzhnyi (Utkholoksk rookery), and
(10) in the other Moroshchechnaya River (south of Cape Khar’yuzov)
(Lun’, 1936).

241

S19

Six rookeries were detected in 1958 roughly in the same section of
the western Kamchatka coast: two in the lower courses of Moroshchech-
naya River, one on Cape Khar’yuzovo, one on Cape Yuzhnyi, and the
rest near Cape Utkholoksk and Cape Babushkina (Tikhomirov, 1966a).

No definite data whatsoever are available on the concentrations of
larga on the continental coast of the Sea of Okhotsk to the southwest
of Okhotsk; however, as at other places, here too the larga is found in
the summer-autumn season but more isolatedly, and evidently without
forming regular rookeries.

On Sakhalin, it is common not only along the western coast, espe-
cially in the northern part commencing roughly from the threshold of
Nevel’sk Strait to the Gulf of Sakhalin inclusive (Tyk Strait, Cape Lak,
etc.; Gakichko, 1931), but also on the eastern side. In the northern third
of Sakhalin largas are confined in larger numbers than in the more south-
ern regions, on the open beaches as well as in Nyisk, Daga, Chaivo bays,
and are particularly abundant in Pil’tun Bay, forming here and there
fairly regular rookeries (Ambroz, 1931).

The Kuril range is inhabited almost wholly but the true larga inhabits
mainly the northern and southern islands, being almost wholly absent in
the central part of the range inhabited by the pagophobic island or Kuril
seal, which is more widely distributed there, on no less than 28 islands
including the Malaya Kuril range (Belkin, 1964; Marakov, 1968).

The coastal strips of the southern half of Kamchatka, western as well
as eastern, fall in the normal range of the larga; the island seal, however,
inhabits predominantly the sea coast. Both forms are encountered on the
Commander Islands, predominantly on Mednyi Island in Zabobrovaya
and Zapalata bays in the northwestern extremity (Marakov,* 1966) form-
ing regular rookeries there. The majority represent the pagophobic form
of seal; the larga is encountered there comparatively rarely.

The summer-autumn rookeries on the coasts of southern Kamchatka
are generally few. They are noticed particularly near the estuary of the
Mutnaya River, on Capes Khodzhelaika and Senyavina, and also in the
region of Vakhil’ River; about a hundred seals were counted in each
area (Ostroumov, 1966). In Karaginsk Gulf, the larga is common and
even abundant at times in winter and summer and there is even a rook-
ery along the coasts of Karaginsk Island (N.N. Gerasimov). The animals
inhabit other points too, including Avachinsk Bay (V.F. Muzhchinkin).
Groups of them are found in the region of Kronotsk Bay from July
through October in the estuarine sections of the Kronotskaya, Tikhaya,
and Mutnaya rivers (in water as well as on sandy shoals) (R.M. Vik-
torovskii). The range in the Bering Sea encompasses the entire coastal

320

belt from Kamchatka to the Bering Strait. The larga is evidently encoun-
tered almost exclusively (arctic form of the subgenus) in all of this
extensive expanse, covering the region of Parapol’sk Dolo and Kara-
ginsk Island, Olyutorsk Gulf, Koryak coast, and Gulf of Anadyr, as well
as the Chukchi Peninsula coast. This coastal zone is unfavorable to the
habitation of the island pagophobic seal as it is blocked by the winter ice
floes with the exception of only the southernmost part; this very factor
is responsible for the seasonal distribution of the larga. As in the Sea
of Okhotsk, it is confined here in winter to the areas far away from the
coasts in the sea beyond the limits of the shore ice and highly cohesive
but broken ice practically outside the continental zone. Only at places
along the Chukchi Peninsula coast where the shore ice is not much and
open pools are formed, is it seen there from time to time even in winter,
thus providing a basis for affirming that it is confined there year round
(Freiman, 1936b).

In the winter-spring season the range of the larga in the northern ©
part of the Bering Sea enlarges roughly up to the edges of drifting ice
floes whose disposition is unstable. The range can extend even beyond the
limits of the continental shelf by the time of maximum ice floe formation.
The main collections of the larga at this time are seen along the extensive
edges of the ice floes (at some distance from the very fringe deep onto
the ice floes), in the region of St. Matthew Island in the northeastern
part of the sea, and almost up to the Pribilov Islands. Another part of
the population at this time is usually found in the zone of drifting ice
floes in the Gulf of Anadyr.

With spring thawing of the ice floes and also to some extent with
the movement of their masses under the influence of the prevailing
southeastern current, the “pelagic” part of the range gradually recedes
northward and by June end (when ice floe remnants are seen only in
the northwestern corner of the Gulf of Anadyr) is confined to a nar-
row strip fringing the contour of the mainland. In this summer-autumn
period isolated stray sections of the range inhabited temporarily by small
local populations are formed around the St. Lawrence, St. Matthew,
and Pribilov islands. This situation, however, is extremely hypotheti-
cal.

On the southern Chukchi coasts, although rare, the larga is encoun-
tered everywhere. In the Chukchi Sea west of the Bering Strait, distribu-
tion of the larga extends in the form of a small tongue into the coastal
belt, evidently only to Kolyuchinsk Bay (P.G. Nikulin; Tikhomirov, 1966b),
which thus represents the western arctic boundary of the Pacific Ocean part
of the range (see footnote on p. 317).

321
243 Geographic Range outside the USSR

Coasts of Japan, Korean Peninsula, and China up to the Yangtse. In
the territorial waters of the Korean People’s Democratic Republic and
South Korea, some stray or a few animals are seen here and there, and
probably not every year; they penetrate on the west coast up to Yonilman
Bay (Nishiwaki and Nagasaki, 1960)77 (Fig. 146).

On the Japanese coasts the larga is more common on Hokkaido
Island. It is carried on drifting ice to the Okhotsk coast and the pago-
phobic form enters from the Kuril Islands. It is also encountered on the
Pacific Ocean side and on the coasts of the Sea of Japan. Evidently it
transgresses south of Sangarsk Strait, along the coast of Honshu Island
and up to Cape Inubo and possibly even more southward (Nishiwaki
and Nagasaki, 1960). The range for the western side of this island has
been shown in extremely general terms (Moore, 1965*). Sometimes the
animal enters even up to Kyushu and Osumi islands.

Young animals are seldom seen every year in spring on the Chinese
coasts of the Yellow Sea (on Shantung Peninsula) (Leroi, 1940*; Schef-
fer, 1958). Yet the seals enter even the Yangtse estuary (Allen, 1938).

On the American side of the Pacific Ocean and the Chukchi Sea the
seal is distributed along the entire coast from the northern Alaskan coasts
through the Bering Strait and the eastern coastal section of the Bering
Sea (including the Aleutian Islands). It then extends in a narrow belt
all along the Pacific Ocean coast including the Alexander archipelago,
and south to the Californian Peninsula (Mexico) at 28°12’ N lat. (Cedros
Island).

Along the northern coast of Alaska the seal reaches east beyond
Point Barrow, entering the estuary of the Colville River (Bee and Hall,
1956) although it is extremely rare there; it was noticed even on Herschell
Island (near the estuary of the McKenzie River).

In the European part of the northern Atlantic it extends to the
extreme southwestern regions of the Baltic Sea (along the Swedish coasts
in the north not farther than Gotland Island), in the Denmark Strait,
and the adjoining regions of the North Sea. From here, one branch
of the range extends north and northeast along the Norwegian coasts
up to Murman; another branch runs along the western side of Jutland,
along the coasts of the Federal Republic of Germany and Holland up
to Pa-de-Kale. Here the range branches again: the smaller and weaker

П The authors undoubtedly committed an error by assuming that the ringed seal (Phoca
hispida) inhabits within the boundaries outlined by them. Equally.erroneously, they point
to the distribution of this species in the south and along the eastern coast of Honshu Island.

‘(ysdeyD УМ) рийпла о20ц4 Чеэ$ чош@оэ эцз до э8ие1 рэ12п1150099] “OPT “BIJ 2

091

323

branch runs along the French coasts, encircles Brittany, and is lost in
Biscay Bay. Rarely, stray animals sometimes reach the Spanish coasts and
extremely rarely even the Portuguese coasts (41° М lat.). In the north,
from La Manche, the range includes the coastal waters of the British
Isles and Ireland and also the west and north of Scotland: Hebrides
archipelago, Orkney, Shetland, and Feeroe islands. The coastal waters of
Iceland represent an isolated section of the Atlantic part of the range;
here the common seal is distributed almost everywhere and is regarded
as the most abundant representative of the Pinnipedia (Semundsson,
1939*). Along the eastern coastal waters of Greenland, the range extends
north up to the Polar Circle and along its western coasts moves roughly
up to 73° N lat. up to Upernavik (Mansfield, 1967) although some stray
animals are encountered from time to time even more northward (Vibe,
1950).

The American part of the North Atlantic range mainly encompasses
the Canadian coastal waters north of Mann Strait including Nova Scotia,
Gulf of St. Lawrence, Newfoundland Island, and the Labrador coast. Far-
ther north and west, the seal is distributed along the eastern, southern,
and southwestern coasts of Baffin Island and along the entire coast of
Hudson Bay. Lancaster Strait can be regarded as the westernmost point
of records of this seal in the Canadian archipelago (Mansfield, 1967).

South of Canada, along the eastern coast of the United States, it is
encountered rarely and only some stray animals reach the state of North
Carolina at times. (K.Ch.)

Geographic Variation

Early in this century concepts regarding the species and subspecies of
seals of the subgenus Рйоса $. str. were quite confusing because of the
extremely incomplete descriptions that were often difficult to compare.
At least three species and three subspecies belonging to the same
subgenus were proposed for the Pacific part of the range alone
(Allen, 1902). Later, attempts to recognize the infraspecific differences
(N. Smirnov, 1908; Ognev, 1935; S. Naumov and N. Smirnov, 1936)
tended to return to the viewpoint of Nordquist (1882*), according to
whom the subgenus has only one species, Ph. vitulina, and only one
subspecies, Ph. у. largha, vicariating with the European Ph. у. vitilina,
which inhabit the entire Pacific Ocean Basin.

Even the latest revision of the seals of the genus (subgenus) Phoca
adopted by Doutt (1942) did not alter the prevailing situation. Moreover,
Doutt affirmed that he was not able to detect any characteristics which
could distinguish the animals on the American coast of the northern part

324

of the Pacific Ocean from their counterparts on the Asian coast. It was
equally impossible to distinguish the Atlantic European common seals
from the American seals.

Doutt notwithstanding, differences between some populations of the
Pacific and Atlantic members of the subgenus Phoca were once again
confirmed while concomitantly the differences noticed within the Pacific
populations remained unexplained (Chapskii, 1955). With further study
of the systematics of the latter, the reason for the heterogeneity of the
Pacific populations of the subgenus was found in their ecological dif-
ferences (Chapskii, 1960). With the growing collection of specimens and
greater information, the taxonomic heterogeneity was demonstrated more
fully (Belkin, 1964; Moore, 1965*; Chapskii, 1965, 1966, 1967; McLaren,
1966). It was possible to differentiate these seals into two groups based
on ecological as well as morphological features: pagophilic seals breeding
on ice floes and aigialoid or pagophobic seals breeding on land.

According to some contemporary scientists (McLaren, 1966; Chap-
skii, 1966, 1967), who assign an independent species status to Phoca
largha Pall., the larga is almost the only form belonging to the first group
of pagophilic seals. Seals of the second group, of which five forms have
been described, wholly represent a different proximate species, Phoca
vitulina L. As pointed out above, in this publication the larga has been
regarded as a subspecies.

Three subspecies of one or the other types are encountered in Soviet
waters. In general, however, the geographic variation of the common
seals in our waters has not yet been adequately studied.

1. Larga, mottled seal, Phoca (Phoca) vitulina largha Pallas, 1811 (syns.
ochotensis, macrodens, pallasii, petersi).

Body length (along the dorsal curvature, Lc) varies from 150- 180 cm
and condylobasal length of the skull, 185-230 mm.

The color (Fig. 147) is relatively light, mottled: the main background
on the underside of the body is whitish, light silver, while the upper part
is mostly dark with a fairly dense network of rather small whitish oval
streaks or rings and with small but vividly colored spots (brownish to
black) scattered over the body. The newborn sports a white furry coat
that lasts through the lactation period.

The transverse profile of the tympanic bullae reveals a steep and
high drop to the base of the shortened and rounded lobe of the external
auditory meatus (Fig. 90). The less stable skull features are as follows: the
nasal processes of the maxillaries extend far posteriorly along the nasal
bones, while the frontal part of the nasal bones is shorter than one-half
their total length (Fig. 92); the posterior edge of the zygomatic bone has

245

245

325

Fig. 147. Typical coloration of an adult pagophilic larga, Ph. у. largha. Verkho-
turov Island, Kuril range, August, 1971 (photograph by S.V. Marakov).

an arcuate notch (Fig. 91); the bony nasal septum in the choanae runs
posteriorly beyond the anterior edge of the palatine bones to the middle
and farther than the latter; the posterior edge of the bony palate has an
oval outline or even an additional arcuate notch at the center, and the
uncinate processes are usually flattened and turned outward (Fig. 93); all
the premolars are located directly one behind the other and not inclined
relative to the general line of the tooth row.

This subspecies inhabits the Far Eastern seas from Peter the Great
Gulf and Pos’et Bay to the limits of the range in the Chukchi Sea, includ-
ing Tatar Strait, Sea of Okhotsk, northern and southern islands of the
Kuril range, the Bering Sea, and partly the Commander Islands.

Outside the USSR it is found in the coastal waters of Alaska (in
the Chukchi Sea in the east to the limits of its range), Aleutian, Pribilov
and other islands in the eastern part of the Bering Sea, south to Bristol
Bay. South of the USSR border, it is found up to the northern coasts of
Japan and the Yellow Sea inclusive.

Two races (natio) were identified within this form (S. Naumov and
М. Smirnov, 1936). One of them, п. pallasii, distributed in the Sea of
Okhotsk and in the northern part of the Sea of Japan, is a very small
form according to S. Naumov and N. Smirnov (condylobasal length
188-228 mm). The other race, п. /argha, inhabiting the Bering Sea,

246

326

differs from the preceding one in much larger craniometric dimen-
sions’ (condylobasal length 191-238 mm). The differences between these
races are artificial since the Bering larga, according to the above, repre-
sents a mixture of the typical larga with the pagophobic populations
(Kuril or island seal). It is possible that the Okhotsk largas in fact
are somewhat smaller than the Bering counterparts and differ from
them in other features, but-they should be compared exclusively with
the pagophilic kin. These “races” are not consistent in the proposed
form.

2. European common seal, Phoca (Phoca) v. vitulina Linnaeus, 1758
(syns. canina, variegata, scopulicola, littorea, Птпае, thienemannit).

Does not differ much from the larga in size. The body length (along
the dorsal curvature, Lc) varies from 175-180 cm; the condylobasal
length of the skull of adult males varies from 203-217 mm and that
of females 190-205 mm, average 205 mm (Ognev, 1935).

The color is more or less dark. The main background on the upper
side is dark, quite often almost black, interrupted by white, uneven cel-
lular streaks; the underside is lighter; dark-colored spots on the general
dark-colored background are less noticeable and less contrasting. Pups do
not sport a white fur coat as it is shed before or at the moment of birth.

The transverse profile of the tympanic bullae is flattened with an
insignificant short step-like drop to the base of the elongated lobe of the
external auditory meatus which is almost straight anteriorly and asym-
metrically pointed at the tip. The posterior edge of the zygomatic bones
has an angular notch; further, the upper portion of the fork has a slightly
convex contour (Fig. 91). The rest of the skull features likewise stand in
contrast to the corresponding ones of the larga (See its characteristics
оп pp. 323-326).

This subspecies inhabits the southernmost part of the Baltic Sea and
from St. Nos in the west to the state boundary in Murman.

Outside the USSR it is encountered throughout the rest of the
European section of the range.

3. Island or Kuril seal,’® Ph. (Phoca) у. curilensis Inukai, 19457? (syns.
chorisii (?), stejnegeri, nummularis,®° macrodens, and insularis; the name
richardi was also used sometimes).

78 Known on the Kamchatka and Commander Islands from the end of the nineteenth
century under the local name “antrus” or “апиг”.

79 The first name for this form should be stejnegeri J. Allen, 1902 (editor’s note). (У.Н.)

80 See the note on р. 307 (synonyms).

247

327

This is the largest form of the species with a body length (along the
dorsal curvature) ranging from 140-204 cm; the condylobasal length of
the skull varies from 208-247 mm (Belkin, 1964).

Color (Figs. 148-151) varies intensely but an extremely dark, often
almost black, main background is a characteristic feature. The ventral
side is often very light in color and interrupted by numerous ring-like
clear spaces, mostly isolated and dispersed unevenly. Among other ani-
mals, the body underside is as dark in color as the upper portion. A broad
and obtuse snout is a characteristic feature. Newborns, as in the case of
the European common seal, shed the embryonic coat in the mother’s
womb.

The transverse profile of the tympanic bulla has the same short
projection to the base of the lobes of the external auditory meatus as in
the common seal, the bony lobe itself being long, set sideways and often
pointed. The posterior edge of the zygomatic bones has an angular notch;
the longitudinal bony septum in the choanae is usually very poorly devel-
oped; a characteristic feature is the angular flexure of the upper contour
of the profile in the zone of interorbital constriction. The second and
third premolars are usually set obliquely relative to the tooth row. The
articular fossae are broadly exposed, flattened. In other skull features,

Fig. 148. Dark-colored Far Eastern island seal (“antur’’), Ph. у. kurilensis. Iturup
Island, Kuril, July, 1966 (photograph by S.V. Marakov).

328

247 Fig. 149. Color details on the vent:al side of the female island seal, Ph. у. kurilen-
sis, Iturup Island, July, 1966 (photograph by S.V. Marakov).

248 Fig. 150. Variation in dark coloration of the adult island seal, Ph. у. kurilensis,
Iturup Island, July, 1966 (photograph by S.V. Marakov).

248

249

329

Fig. 151. Island seal (“antur”), РИ. у. kurilensis, Mednyi Island, July, 1972 (pho-
tograph by S.V. Marakov).

there is only a shift in the specific features or they stand between the
characteristics of the larga and the Atlantic common seal.

This seal is found on the Kuril and Commander Islands and on the
eastern coast of Kamchatka.

Outside the USSR it is found on the Aleutian and Pribilov islands
and Japan and probably on the coasts of southern Alaska.

The tendency to combine the island seal with the subspecies Phoca
vitulina richardi found on the American coasts (Moore, 1965*) cannot
yet be regarded as substantiated for the simple reason that the skin
coloration of these seals differs sharply.

Outside the USSR three more pagophobic subspecies are recognized
(Scheffer, 1958; King, 1964): (1) Ph. v. richardi Gray, 1864—American
waters of the Pacific Ocean from the Alaskan Peninsula to California,
and eastern part of the Bering Sea, including the Aleutian and Pribilov
islands; (2) Ph. у. concolor De Kay, 1842—western Atlantic American

330

and Greenland waters; and (3) Ph. v. теПопае Doutt, 1942—Lower and
Upper Seal Lake on Ungava Peninsula (eastern Canada).
Craniologically, Ph. v. richardi is somewhat closer to the island
(Kuril) seal and correspondingly differs sharply from the larga, but the
skin coloration of the adults and semiadults of the two is often similar..
The range of this seal evidently does not extend in the north beyond
Bristol Bay and should thus be limited to the regions free of ice floes
(Moore, 1966*; McLaren, 1967*). Insofar as the other two forms are
concerned, their morphological features require further study. (K.Ch.)

Biology

Population. It has not yet been possible to determine precisely the pop-
ulation of the common seal (including the larga) in the USSR waters in
spite of several efforts in this direction. In some sections of the Sea of
Okhotsk a visual count has been made in the rookeries but this has not
helped to assess the total reserves of even the larga in the Sea of Okhotsk.
Some local populations have been estimated more accurately. Thus in
the rookeries on the Kuril Islands the population was put at 5,250, of
which 2,000 - 2,500 constituted island (pagophobic) seals (Belkin, 1964).
Very similar figures were recorded even later but without differentiation
into island and larga seals: 6,000 on the Kuril Islands, 1,000 on Bering
Island, 1,000 on Mednyi Island, and 100 on Seal Island (Marakov, 1970).
Insofar as the total population of seals of the subgenus Phoca s. str. in the
Pacific Ocean is concerned, it has very approximately been estimated in
the range of 20,000 - 50,000 (Scheffer, 1958) to almost 400,000 - 450,000
(Chapskii, 1966).

Attempts were made toward a more rational census of the seal pop-
ulation. Based on the average area of the ice floes inhabited by lactating
animals and their average density (1 to 3 newborn/km’, area of “rookery”
150-200 km?, and their number in the Sea of Okhotsk and the Bering
Sea over 2081), the maximum number of pups is approximately 15,000.
This figure is evidently much less than the actual number.

Different figures were arrived at by the end of the 1960s to the early
1970s. According to aero-visual counts on the spring ice floes, the total
larga population in the Sea of Okhotsk was put at 170,000 (Fedoseev,
1971). A similar method had been used sometime before to estimate the
larga population in the Sakhalin section of the range. The results were:
about 10,000 (with pups) in the region east of the mid-northern section
of Sakhalin, about 4,000 in Terpeniya Bay, and 8,000- 11,000 in Tatar
Strait (Fedoseev, 1970; G.A. Fedoseev).

81 This calculation was based on the data of Tikhomirov (1966b).

250

331

А very large proportion of the total population is concentrated in
the Sea of Okhotsk. There is a view that the larga is second to the ringed
seal in “natural level of population” (Pikharev, 1940*; Fedoseev, 1966)
while its total reserves in this sea are not less than 20-25% of the total
population of all species of seals (Tikhomirov, 1966a).

The larga is most abundant in the following areas in one season or
the other: (1) southwestern Sakhalin part of the sea north of the southern
Kuril Islands; (2) western corner of the sea from the Gulf of Sakhalin to
Uda Bay, including the Shantarsk archipelago; (3) northeastern part of
the sea including Shelikhov Gulf; (4) the region adjoining northwestern
Kamchatka; and (5) the coastal belt west of P’yagin Peninsula up to
the estuary of the Tauya River. The larga population elsewhere in the
continental expanse between Tauisk Bay and the Shantarsk archipelago
is evidently much less than in all the regions cited above.

Such a large number of summer-autumn coastal rookeries as found
along the coasts of the Sea of Okhotsk is not found anywhere in other
parts of the Soviet Far East. The maximum number is concentrated there
in the western corner of the sea (mostly in the Shantarsk archipelago
and Yamsk-Siglansk region) and along the western coast of Kamchatka.
The most detailed information on the distribution and population of the
rookeries is available for that part of the sea from the Gulf of Sakhalin
to Uda Bay, including Shantarsk Island. The concentrations of the larga
here along the mainland beaches are few and rather small, with the
exception of Konstantin Bay; at one time there in the 1930s, up to eight
rookeries with a total population of 900 to 4,000 seals were reckoned
(S. Naumov, 1941; Pikharev, 1941).

The population of the larga is not high in the western part of the
Gulf of Sakhalin (S. Naumov, 1941). In Nikolai Bay in 1929, one source
put it at 100 or a little more and another source in 1938 at 1,000. The
larga is not very rare in Ul’ban and Tugursk bays in autumn. In Uda
Bay there are obviously no large congregations. The population is some-
what more on the island rookeries in the Shantarsk archipelago, espe-
cially on the Sivuch’i Кати, Srednii, Ptichii, Utichii, and also Мау
Shantar islands. In 1929, the total population of this region was put at
2,000 (S.P. Naumov, 1941) and in 1938 and 1939, at 16,000 to 18,000
(Pikharev, 1941). The population was the same in 1932 but later dou-
bled (27,500 -35,000) (P.G. Nikulin). In some very large rookeries оп
the Shantarsk Islands, especially on the Sivuch’i Кати! (Shantarsk archi-
pelago), 6,500-7,000 (Pikharev, 1941) or even more (P.G. Nikulin) were
reported in the 1930s. In 1958, the total population of the larga in these
rookeries was put at 1,000- 1,600 (Tikhomirov, 1966).

250

251

332

Fig. 152. Гагра, Ph. у. largha (left) and island seal (“antur’’), Ph. у. kurilensis,
Mednyi Island, June, 1969 (photograph by S.V. Marakov).

Considerable concentrations of the larga were noticed in 1929 - 1930
in the Yamsk-Siglansk region in the fore-estuary expanses at Ireti and
Yama and very significant rookeries at Ostrovno, Kamyl, and the Bero-
zovka River estuary. Not even a rough estimate of the population there
was ventured. The largas were fewer in Tauisk Bay than in the preceding
region; coastal rookeries were few and small in the 1930s (albeit large
numbers occurred in the western part of the bay) (Freiman, 1936). In
the 1960s, the number of largas in Tauisk Bay was small, barely 2,000.
The low population is explained by the relatively high coastal habitation
and possibly low food availability (Tikhomirov, 1966).

In terms of larga population, Shelikhov Gulf is next after the Yama
region and the western coast of Kamchatka (Fig. 154) but the rookeries
there, as already mentioned, are few, especially in the region of Taigonos
Peninsula although the data on the former population have been given
for only two—Uikan rookery (up to 500) and Cape Verkholamsk rookery
(up to 400) (Freiman, 1936).

No less than 12,000 seals were counted in the 1930s in 10 rookeries
in the Tigil’sk region on the western coast of Kamchatka (from the
Moroshchechnaya River to the Tigil’ River). The population in some

333

\

251 Fig. 153. A herd of largas, Ph. v. largha, Karaginsk Island, Bering Sea, July, 1968
(photograph by N.N. Gerasimov).

252 Fig. 154. A section of the rookery of the larga, Ph. v. largha, western coast of
Kamchatka in the estuary of the Utka River, September, 1967 (photograph by
D.I. Chugunkov).

rookeries went up to a few thousand (Tal’nichnoe, 15-20 km south
of Cape Babushkina, 3,000-4,000 as also on Cape Utkholoksk) (Lun’,
1936). In 1958, six rookeries with a total population of about 6,000 were
found in the same region (Tikhomirov, 1966).

The total population of the larga in the western part of the Bering
Sea has not yet been determined. It is known thai it is not rare along
eastern Kamchatka and Koryak Land (from Cape Olyutorsk to Cape

252

334

Navarin). It is quite abundant also in the coastal waters of the western
and southern regions of the Gulf of Ападуг; by the early 1930s, the
proportion of the larga there constituted up to 15% in the local hunters’
catch but dropped to a few percent on the Chukchi coasts where this seal
inhabits in maximum numbers the Pinkegnei, Tkain, and Privideniya bays
(region of rivulets on bald patches) (Freiman, 1935а*).

The population on the- Commander Islands, as already noted, is
extremely small; it is usually represented by the settled pagophobic form
of island (Kuril) seal; the larga is, however, encountered there as a rare
find. While only one rookery was known there in the early 1930s with
a hundred seals or slightly more (Barabash-Nikiforov, 1936), evidently
there are now no less than 15 rookeries (although no exact figure is
available). On the Kuril Islands, the island seals have been reported on
28 islands, with pups included on 11 of them (Velizhanin, 1967).

During their residence on drifting ice floes, the largas are much more
numerous in the central and eastern parts of the sea, especially in the
region of St. Matthew Island, in the zone of ice floes south almost up
to the Pribilov Islands, and northeast of the latter. Here, in the spring
of 1963, over 90% of the seals encountered on ice floes were largas. A
nearly similar picture was noticed even in the preceding season; in the
more northern regions of the sea, between St. Matthew and St. Lawrence
islands, the larga accounted for only a little above 20% (Kosygin, 1966a*;
Tikhomirov, 1966). Larga in the Bering Sea has been reported as con-
stituting not less than 30% of the total seal population inhabiting there
in the zone of drifting ice floes far from the coasts, and occupies sec-
ond place after the ribbon seal in terms of population (Tikhomirov,
1966b).

In the other regions of the Far East, considerable concentrations of
larga are seen in Tatar Strait although these groups are much smaller
than in the Sea of Okhotsk and the Bering Sea. In Tatar Strait the larga
predominates markedly over all the other species of seals. The reserves
in the Sea of Japan are small; it is evidently maximum on the threshold
of Tatar Strait and in Peter the Great Gulf. It has, however, not been
possible to express the population numerically.

The population of the common seal in our waters of the Atlantic
Basin is extremely small. These seals are particularly rare in the south-
eastern Baltic Sea and relatively few on the Murman coasts. It is signifi-
cant that the coastal people so far have not given their own name to this
species.*? Only some stray animals were caught in the Murman region
after several years, at the very end of the last century (N. Smirnov, 1903).

82 At the beginning of this century, this seal was known in Murman as “kamenka”.

253

253

335

The seals of this species were not caught in nets, which was the form of
hunting in vogue in the 1930s.

Habitat. The Atlantic common seal and the Pacific pagophobic forms
inhabit selected sections of the coastal zone in a fairly settled manner.
Within the southern Baltic, on the Polish and German coasts, the seals
emerge onto land in the uninhabited sandy or rocky coasts, small islands,
spits, and bald patches in the river estuaries and bays. On Murman, they
are evidently confined to the bays and estuarine sections of rivers, espe-
cially of the Voron’ya; they probably take advantage of the rocky coasts
and islets. The Far Eastern pagophobic populations (island seals) select
for their rookeries reef ranges and groups of individual rocks, as well
as sandy-pebbly coastal sections which offer them protection from the
surf (Fig. 155). Such conditions on the Kuril Islands are provided by
reefs, small coastal islets, sections of very large uninhabited islands, low
rocky ledges, niches between rocks, and coastal shoals (Belkin, 1964а*;
Velizhanin, 1967). Some animals (males) at places, for example on the
Lovushki Islands, rest on small isolated rocks almost fully submerged in
the strait and overgrown with algae (Belkin, 1964b*).

Fig. 155. Typical habitat of the island seal (“antur”), Ph. у. kurilensis, on Mednyi
Island. Commander Islands, Zapalata Bay, June 6, 1962. Sea otters also live here
(photograph by S.V. Marakov).

254

336

The island seals on the Commander Islands (especially on Mednyi
Island) select diverse biotopes for rookeries. These may be sections of
the low coast, pebbly or rocky talus zones at the foot of a precipitous
cliff, scarps, sloping projections of rocks, etc., especially under reefs pro-
tected from the surf. For temporary residence, these seals inhabit large
exposed rocky mounds that have been rounded by intense wave action
and smoothened by surf abrasion (Marakov, 1968, 1969). The Bering
Island population is confined mainly to the region of the Barrier Reef
(Marakov, 1967b).

The pagophilic seals, 1.е., the largas, associated with the coastal land
are not permanent settlers since they abandon the beaches jammed by ice
floes in winter and spend the winter-spring season on drifting ice floes.
There is a distinct difference in the site selection of the larga for whelp-
ing, lactation, and molt. Nevertheless, it is important for it that the ice
floes be quite stable, clean, without large hummocks, and at the same
time not excessively compacted, with an abundance of water pools to
provide access to the water. Largas avoid stationary shore ice or a highly
compacted mass of broken ice floes. According to some observers, they
exhibit a distinct preference for ice floes along the edges (Tikhomirov,
1964) since they are very sparse; according to others, they are encoun-
tered not only close to the edge but also far from it, inside drifting ice
floes that are highly compacted (Kosygin, 1966). The larga begins to be
seen directly on the coasts immediately after the thinning of the ice cover
and access to them becomes available. Evidently it appears earliest and in
some numbers close to the river estuaries where the ice floes disintegrate
usually more rapidly and where the food conditions are more favorable.

Immediately after coming onto the beaches, initially in small

numbers and later en masse, the seals begin to move on land to

form temporary rookeries. At the beginning of summer, these rookeries
appear disorganized and tentative and the animals rest here and there
without confinement to any one particular site for long. The islets
and spits in the river estuaries that dry up in low tide often serve
as rookery sites; where such sites are not available, they are confined
right on the gentle beaches in the lower courses of the river itself
(Tikhomirov, 1966b). When attracted to the fish migrating for spawning,
they themselves enter the estuaries and the lower courses of rivers and
even ascend tens of kilometers up the estuary of some rivers.

Transgression into rivers is a regular phenomenon. Evidently there
is no single sufficiently deep river abundant in fish in the spawning
period which has escaped the notice of the larga. Its long transgres-
sions, sometimes hundreds of kilometers, into rivers such as the Amur
and the Anadyr, have long been known. In the former river the larga

254

255

98

Fig. 156. Largas on a drifting ice floe in Litka Strait, Karaginsk Island, Bering
- Sea, early June, 1968 (photograph by D.I. Chugunkov).

was seen even 400 km away from the estuary (Nikol’skii, 1889). In the
past the larga ascended 60 km away from the sea along the Sakhalin
rivers Poronai and Тута. Transgressions for 10-15 km were noticed in
the Tyma River comparatively recently (Ambroz, 1931). The larga trans-
gresses regularly into many rivers of western Kamchatka such as the
Voyampolka, Khairyuzov, Belogolovaya, Tigil’, Kavacha, Utkholok, and
Sopochnaya, ascending 5-35 km along them (Lun’, 1936; Ostroumov,
1966). These seals also enter some rivers on the southwestern coast of
Kamchatka. —

The larga transgresses for considerable distances into the rivers of
eastern Kamchatka. In the 1960s, they were found 35 km from the estuary
of the Kamchatka River; a few decades before that they transgressed even
farther, up to Klyuchi village and even up to Kozyrevsk, 250 km from
the estuary. The larga transgresses for distances of 25-30 km into Zhu-
panov, Vyvenka, Kultuchnaya, and Apuka. In many other rivers, however,
these seals are concentrated close to the estuary, often a whole herd, or
transgress only into the lowest courses (Ostroumov, 1966).

Instances of distant transgressions of the common seal, episodic in
nature and regarded as exceptional events, have been known in the
remote past even in the rivers of Western Europe, especially in the Elba,

338

in which they were sighted at a distance of 646 km, almost 700 km, and
even 757 km from the estuary (Moore, 1955*).

By autumn, sightings in the rivers become increasingly rare and the
bulk of the well-fed largas concentrate on the sea coasts close to their
favorite haunts, where they form regular (permanent) rookeries.

Groups of isolated rocks smoothened by water and close to the coast
or the rocky shore protected from the surf represent preferred sites in the
Sea of Okhotsk. In any case there is a distinct preference for land sections
submerged in high tide. Sections with an intensely rugged coastline and
abounding in reefs and tiny islets meet these conditions to a very large
extent. Some dependence could even be established, other conditions
remaining comparable, between the degree of ruggedness of the coast
and the nature of the rookery: the more rugged the coast, the more
rocky it is and the more abundant the reefs on it, the larger the sites
suitable for rookeries. Here the seals are more numerous but the number
of animals in each rookery is small.

Hunters distinguish three main types of rookeries (Tikhomirov,
1966a). One is the rookery located in a highly rugged locality .where
sections of large pebbles are interspersed with rocky hummocks and
large isolated rocks. Such, for example, are the rookeries on the Sivuch’i
Islands. The second type is found on exposed sandy or pebbly laidas
(low coastal plains dissected by tortuous rills), islets, spits, gently
descending into the water (rookeries on Cape Polonsk, in Tauisk Bay in
Melkovodnaya Gulf, and in western Kamchatka in the Moroshchechnaya
River). The third, and most common type of rookery, is represented by
coastal reefs and rocks, for example on Capes Khairyuzovo, Yuzhnyi,
Utkholoksk, etc. (western Kamchatka).

At several places the larga inhabits land together with or in the
neighborhood of pagophobic island seals. Such instances are particularly
numerous in the northern and southern parts of the Kuril Islands; in the
southern parts of the range, largas are even permanent settlers (Belkin,
1964). The biotopes of both forms of seals there are almost identical. A
somewhat similar situation is noticed on the Commander Islands where
the larga is, however, far less numerous, constituting a rare find in the
rookeries of island seals.

Food. The common seal in general is a distinct fish-eating animal
but not specialized for a limited number of any particular species and
feeds on quite diverse, mainly large fishes available on the coasts at a
given time of the year. At the same time, various invertebrates constitute
a significant proportion of its food ration.

For nearly two-thirds of the year, from spring to autumn, the larga
is assured of abundant fish food. In eastern Sakhalin, in spring, it takes

256

339

advantage of the approaching herring; in summer it feeds partly оп
herring and also on humpbacked salmon [pink salmon, Oncorhynchus
gorbuscha] and Siberian salmon [chum, O. keta]. When the summer
run (migration) of salmon slackens, the larga takes to consuming river
fish—“kundzha,” [Sakhalin char, Salvelinus leacomaenis] rudd etc. Some
seals while chasing these fishes reach far up the rivers. The larga takes to
capelin during its en masse availability in the first half of July (Ambroz,
1931).

In western Kamchatka, in late spring and early summer, the larga
feeds on char and sea trout which descend from the rivers into the fore-
estuarine regions of the sea, and later on smelt and capelin which form
spawning schools on the coasts. Later, from the middle and second half
of June, the larga takes to feeding on herring which comes close to the
coasts in large numbers. At the end of the herring season, usually from
early July, it takes to chasing salmons—chum, red salmon, pink salmon,
etc. (Lun’, 1936). On the opposite side of the Sea of Okhotsk, on the
Shantarsk Islands, it takes to navaga (Pacific navaga [saffron cod, Eleginus
gracilis) in spring as supported by the coincident en masse approach
of this fish with the increasing population of the seal (Lindberg and
Dul’keit, 1929). In Schast’ya Bay (western part of the Sea of Okhotsk)
a similar relation has been reported between the gathering of the larga
and the availability of smelt in shoals (S. Naumov, 1941). In Tatar Strait,
at the end of May, numerous largas were noticed; they were evidently
drawn there by the availability of herring.

In the snowy period, when the adult seals are confined to the regions
of drifting ice floes far removed from the coasts, the food regime of
the seal is somewhat different. Thus in the southern part of the Sea of
Okhotsk, not far from the Hokkaido coasts (Wilke, 1954), the larga feeds
in April mainly on pollock (Theragra chalcogramma) followed by herring,
while cephalopod mollusks, especially the octopus (Octopus daeffeni),
account for a much smaller quantity of food intake. The ratios between
these food groups are hardly constant. This seal readily feeds on whatever
species are available at the time of foraging for food.

The larga concentrations in the southeastern part of the Sea of
Okhotsk, on the Kuril Islands, feed in the spring-summer period quite
intensively, consuming roughly equal proportions of fish and cephalopod
mollusks. The following fishes were detected in the stomach of largas:
Atka mackerel, rockfish (Sebastodes sp.), walleye pollock, and saffron
cod; octopuses from among cephalopods; and squids of the genus Gona-
tus to a much lesser extent (Panina, 1966).

At the end of the reproductive cycle, quite often even after incom-
plete molting, the larga migrates into the coastal zone, concentrating

257

340

mainly in the estuarine sections and the lower courses of rivers where
salmon migrate in large numbers for spawning. The seals reach there
either at the time of high tide or at a much later period or are even
confined to the fore-estuary or bar sections. On the western coast of
Kamchatka, in the region of Tigil’ River, even brooklets to which the
pink salmon go for spawning are quite often literally blocked with the
seals, which are caught there in tens or even more. Not being able to
penetrate the brooklets, the seals at times even crawl in high tide to the
estuary, thereby blocking the entry of fish until their hunger is satiated.
Tens of larga transgress into very deep and broad rivers and in rivers
such as the Tigil’ even 200-300 or more may be seen (Lun’, 1936). The
same pattern of transgression is seen at other places also. While chasing
the fish, the larga can dive under water without respite for up to 400 m,
gaining a speed of over 4 m/sec, or it can dart several meters into the air
(Chugunkov, 1969).

For 1.5-2 months (August and even September), the seals are not
satisfied with the fore-estuary sections but transgress even into the lower
courses Of salmon-bearing rivers and brooks. This transgression ceases
abruptly only in October. Only rare seals, not yet fully fed, hunt for food
at this time even in the largest rivers.

The affinity of the larga for coastal sections and rivers abundant
in salmon, consuming much and damaging the commercial fish (often
snatching only the daintier and meatier portions of the back), is respon-
sible for its reputation as a carnivore inimical to the fishing economy.
However, because of incomplete data, it is yet difficult to establish the
exact extent of damage inflicted by the larga.

In addition to fish and cephalopod mollusks, crustaceans occupy a
definite place in the diet of the larga. Among these crustaceans, the main
ones are shrimps and partly amphipods. In Tatar Strait common shrimps
of the genus Sclerocrangon and amphipods of the genus Gammarus have
been found time and again in large quantities in the stomach of seals
caught at the end of April; in some cases the stomachs were literally
crammed with shrimps (Freiman, 1936b). How important shrimps are as
food items for the larga in this region is deomonstrated by the following
data. Of 36 seals caught on May 12-14 in Nevel’sk Bay, the stomach
of 21 animals was filled with shrimps; of the remaining 15 caught on
May 15-19 in the environs of Cape Lak, the stomach of nine seals also
contained only shrimps while the remaining six seals contained shrimps
with fish remnants (S. Naumov, 1941).

In the spring-summer period the adult and semiadult Bering largas
feed mainly in the morning (up to 9:00 a.m.) and in the evening (after

341

4:00 p.m.) and consume a large amount of mixed food although fish con-
stitutes the main item. at this time. Slightly less than one-half of the seals
feed on a single type of food at this time of year: only fish 29%, only
crustaceans 11%, and only cephalopods 7% (Gol’tsev, 1969). In Tatar
Strait (Gol’tsev, 1971) the larga in spring (in March - April) feeds mainly
on pollock, octopuses, navaga, squids, and sand eels. The food of the
larga in Peter the Great Gulf has not been adequately studied. Evidently
in Spring (in March), navaga constitutes the main food followed by floun-
der (Liopseta sp.) and thirdly by perches (of the family Scorpaenidae); in
addition to these, the larga there also consumes shrimps (Pandalus sp.)
(Goltsev, 1971).

In the gulfs of Anadyr and Karagin, where studies were carried out
for three years (Gol’tsev, 1971), the larga feeds in spring on three groups
of animals: (1) different species of fish, mainly the polar cod, pollock,
sand eel, goby, and to a lesser extent navaga, armed bullheads (Ago-
nidae), stichaeid blennies, etc.; (2) octopuses; and (3) crustaceans (mainly
the shrimp Pandalus goniurus and more rarely the following species:
Spirontocaris macarovi, Eualus gaimardi, and Е. fabricii). Groups of crus-
taceans, such as amphipods, crabs, and hermit crabs are very rarely found
in the stomach (usually in not more than 2%).

The following aspects could be set down with respect to the food of
the larga: (1) evidently there are no total or prolonged seasonal absti-
nences in the feeding of the larga even at the time of molt; (2) the “food
spectrum” of the larga is quite broad. Evidently it is this phenomenon
that helps them to survive under diverse biotopical conditions which, in
the ultimate analysis, explains the unusually extensive distribution of this
species, extending from the Chukchi to the Yellow seas; and (3) eurytro-
phy in turn is ensured by the fact that the large can get at food not only
in the shallow and surface sections of the sea, but also at considerable
depths, of the order of 300 m or more.

Some other feeding patterns have also been pointed out (Gol’tsev,
1971), especially the daily feeding rhythm: the animals of this species feed
mainly in the morning and evening hours. Some age-related changes of
food intake were also noticed; the juveniles after lactation feed initially
on amphipods, shrimps, and schools of small fishes while the proportion
of pelagic fishes (navaga, polar cod, and sand eel) as also cephalopod
mollusks increases later. Evidently bits of algae and stones and sand
accidentally enter the stomach along with the food and are totally unin-
tentional.

Amphipods disappear from the food of adult largas and the spe-
cific proportion of benthic organisms, fishes, and crustaceans (decapods)

258

342

correspondingly increases. Among the benthic and demersal fishes con-
sumed are the flounder, halibut, goby, stichaeid blennies, and some
poachers; among decapods the snow crab and others. The quantum of
cephalopod mollusks consumed also increases considerably.

Although there are no clear ideas so far on the seasonal changes
in the feeding intensity of the larga, it is not entirely uniform. During
reproduction and lactation feeding is most intense and weakens sharply
during molt although food intake does not cease altogether, as shown
by the data given above. However, after the end of the snowy period,
feeding is again intense. As the seals become well fed, the intake again
decreases. In any case, the Okhotsk larga at the end of summer is so
well fed that its food hunting impulse weakens sharply. It is at August
end and early September that the larga loses interest in food in spite of
the continuing arrival of salmon at places and the availability of other
food. The larga then prefers to rest on the beach rocks or shoals in the
interval between high tides.

Evidently there do exist some minor interruptions in feeding caused
by seasonal massive availability of one or the other food items, i.e., short
interruptions in their en masse availability, when the numbers of some
species decrease and the migration of others has not yet peaked.

The Far Eastern pagophobic (island) seal differs vitally from the
larga in its food regime. Being a more settled animal, it relies on
various food items available in the region of its habitation which, on
the Commander Islands, is restricted to just a mile-long coastal strip
(Marakov, 1968). Such a relative stability of habitation of the local
populations of island seals is evidently due to the extremely abundant
benthic biomass, up to 20-30 kg/m on the coasts of the Commander
Islands (Marakov, 1969). Invertebrates play a relatively important role
in the food of these Commander Island seals although the seals feed
there on fish too, especially the smooth lumpsucker Cyclopterichthys
ventricosus, and, when it is no longer available, on sculpins (genus Сойи5)
and greenlings (Hexagrammidae); from among the invertebrates, they
consume predominantly cephalopod mollusks, crabs, even amphipods,
gephyreans, etc. (Barabash-Nikiforov, 1936). Instances were also noticed
of the intake of rockfish (Sebastodes sp.) and mysid. In the winter-spring
season the Commander Island population survives almost exclusively on
invertebrates; these food components, it should be assumed, occupy an
important position in different seasons of the year. There are other
indications too that smaller fish are consumed by local seals. For
example, these seals are not seen in Saranna Bay at the confluence of
the Saranna River, which has the most abundant salmon reserves in the
Commander Islands, although hundreds of these seals regularly inhabit

259

343

the coasts 10 km away from its estuary, оп Саре Tonk (Marakov, 1967).
Neither were seal concentrations observed around the estuaries of other
rivers in spite of the availability of salmon in them.

The corresponding Kuril populations feed on crustaceans (shrimps),
cephalopod mollusks (squids) Pacific ruff, bullheads, and other marine
animals of the coastal zone (Velizhanin, 1967). Another list of food
items has also been given for these same regions: greenling, rockfish,
saffron cod (navaga), goby, walleye pollock, flounder, cephalopod mol-
lusks (especially octopuses, more rarely squids) and also diverse types
of shrimp (Crangon dalli, Sclerocrangon sp., and Lebeus polaris) (Panina,
1966Ъ).

The nearest kin in the American territorial waters of the Bering Sea
and in the more southern territorial waters of the USA and Canada,
Richard’s seal, also feeds on extremely diverse foods. On the Pribilov
Islands, as far as can be judged from the limited data (Lukas, 1899*), it
feeds in summer mainly on octopuses (Octopus punctatus) and to a lesser
extent on crabs. On the Aleutian Islands (on Amchitka Island), its food
mainly consists of greenling and octopuses (Kenyon, 1965). The more
southern seal populations, living beyond the Bering Sea limits, consume
mainly fish, including cod, herring, salmon, flounder, rockfish (Sebas-
todes sp.), goby, and even skate; they also consume lamprey, cephalopod
mollusks, and crustaceans (Scheffer, 1928*; Scheffer and Sperry, 1931;
Imler and Sarber, 1947; Fisher, 1952). In the season of spawning migra-
tions of river smelt the seals feed almost exclusively on this fish but,
when available, herring becomes their main food item; likewise salmon,
when available, is consumed in significant quantities.

Nothing is known about the food of the common seal on the Murman
coasts. On the British and Dutch coasts, and generally on the coasts of
the North Sea, the food of these seals includes herring, flounder, cod,
goby, eel, and occasionally salmon. Shrimps are consumed but less often.

The quantum of the daily food intake, according to the data of
zoological gardens, averages 5.5-7.5 kg of fish depending on the age of
the animals and the nutritional value of the food; the annual intake thus
amounts to 1,800 kg (Moore, 1955*). Taking into consideration the fairly
prolonged intervals when there is no feeding in some other seasons, this
norm cannot be extrapolated to the food ration under natural conditions.
At the same time, the experience of feeding seal pups weighing 17.5 kg at
the time of capture on the western coast of Kamchatka, and in captivity
at Utkin, confirms the phenomenal appetite of these animals. They do
not feed regularly or even daily, but in captivity consumed 1.8-2.6 kg
per day of different types of salmon, 1.е., roughly 15- 19.5% of the pup’s
own body weight. The young seals voraciously consumed various types

260

344

of local salmon: Chinook, masu, pink, chum, silver, arctic, and “Kam-
chatka steelhead” (with preference for small live fish). On catching a
large fish, the larga kills it first by crushing the head between its teeth
while small fish are swallowed whole. Larger fish are torn to bits using
the teeth aided by the fore flippers. When satiated, the seal desists from
fish pursuit (Chugunkov, 1967).

Home range. The distinctly settled population, characteristic of the |
Commander and Kuril (island) seals, as already mentioned (p. 335), occu-
pies relatively small sections exposed to very little seasonal variation.
The Commander Islands are inhabited by individual herds which form
extremely close-knit rookeéries at various places and remain almost sta-
tionary year round. Their precise boundaries have not been determined

and establishing them is not easy since the animals in search of food or

impelled by other causes can be dispersed quite widely and encountered
almost everywhere on the coast (Marakov, 1966*).

The herding tendency of the seals varies sharply in the different
seasons while the area covered by their habitation in a given season
undergoes wide variation. A herd of 50-100 seals inhabiting an area of
a few hundred square meters, entering the water at high tide, is confined
within some 3-5 km. But in the winter-spring season the population is
“dispersed” in an extensive expanse of drifting ice floes far away from
the coasts. This process of animal scattering peaks by spring when the
mating pairs are often seen at a distance of several kilometers or many
hundreds of meters from each other. The density of largas at the time
of whelping in some regions of the Sea of Okhotsk averages 1.2 animals
per km? (G.A. Fedoseev). At the end of lactation and mating, they again
form groups. Initially these are small and are not as coherent as the much
later beach rookeries in which the seals are often disposed shoulder to
shoulder. The herd begins to grow only in summer when the animals
arrive from every direction and form the final rookeries. The density of
such colonies is dependent on the total local population concentration
of food items and the nature of the coastline.

Daily activity and behavior. Alternation of periods of wakefulness
(activity) and sleep (rest) is mainly guided by the need for feeding and
at places, especially in the formation period of the beach rookeries, by
tidal conditions. The periodicity of daily activity under the influence of
the latter factor is very sharply manifest among the larga of the Sea of
Okhotsk in the beach rookeries on islets and reefs that are inundated.
Right at the commencement of low tide, the animals gather around an
exposed shoal in anticipation of rapid food availability. As soon as the
first patch of their favorite land opens up, they rush to occupy the site
right at the water’s edge. As the water recedes, new animals crawl to

345

the newly exposed sections of the shoal ог reef, forming а large herd т
a comparatively short time. Land occupancy and the first residency of
animals on exposed land close to one another is accompanied by loud
discordant noises, restless movements, and sometimes even scuffles. Ulti-
mately the animals become quiet and begin to rest. They lie somnolently
even under unfavorable conditions—frost, rain, or snow—being driven
from their place only by a high tide (Lun’, 1936).

The animals in the beach rookeries sleep very lightly. There are no
special guards in the rookery. From time to time, one or the other seal
awakens, raises its head, peers about, and on sighting no danger, drops
its head and sleeps again (Tikhomirov, 1966).

A reverse process occurs with the commencement of high tide as the
land becomes flooded. The water steadily drives the animals from the
peripheral sections and they quietly move sideways in order to gather
again at the former site when the tide recedes.

When sleeping or quietly resting seals are disturbed by danger, say
the intrusion of man, the entire rookery is instantly enlivened and the
animals rush to the water pell-mell like an avalanche. As soon as the
danger has passed, they gradually return to their former places on the
coast.

Opinions on which of the sense organs are better developed in the
larga are somewhat contradictory and the matter still not wholly resolved.
According to some, sight and smell (Tikhomirov, 1966) and, according
to others, sight and hearing (Marakov, 1966*) are the best developed.
Whichever, the larga is one of the most cautious seals.

It is difficult to discuss the daily regime of the larga in winter because
of inadequate data. Before the period of reproduction commences, the
animals are evidently quite scattered, far away from the coasts among
drifting ice floes, and take advantage of open water pools for respiration.
The larga usually does not make air holes in the ice. With the onset of
the mating period, the activity of the males naturally intensifies. Since
ultimately only one male remains for long alongside the female, and the
males and females are numerically equal, animal pairing can be assumed
as relatively peaceful.

The activity of whelped females is subordinate to the lactation
rhythm; in the intervals between suckling her pup the female has
sufficient time for rest although she has to ensure the safety of her pup
and also feed herself. At the end of the feeding period the adult animals
split into groups of some tens each, sleep long periods on a large ice floe
or simply rest in a somnolent state. In spring immature animals remain
apart from the adults but no information is available on their behavior
at this time.

261

346

At the conclusion of molt the larga again becomes highly active,
partly because of the preceding period of relative abstinence from food.
The herds break up and abandon the ice floe. Further, in the period of
intensive feeding, the animals attracted by schools of fish again gather
into groups. Now, once again, their activity alternates with hours of
rest.

The pagophobic (island) seals on the Kuril and Commander Islands
are quiet and peace-loving animals with an even more distinct herding
tendency than the larga. Their lairs consist mostly of 20-50 animals
though some contain even up to a hundred. Quite often the same lair
also holds one or more single seals without animosity. Similarly, They
exhibit no enmity toward other warm-blooded animals. Scuffles are pos-
sible only between the males in the mating season. Affinity for a given
site is more pronounced in them than among the larga. While the lat-
ter usually inhabit the lower coasts and advance toward the coastline
with the receding waterfront during low tide, the island seals can be
seen 30-60 m away from the waterline (Velizhanin, 1967). In spite of all
their quiescence, however, the screams of the Commander Island seals,
similar to those of bitterns, can be heard during the period of lactation;
the voice of a female calling her pup in case of danger sounds the same
(Marakov, 1966*).

Seasonal migrations and transgressions. These aspects have not been
adequately studied. Unlike the harp and some other species of seals,
the common seal is usualiy regarded as a settled animal, albeit this is
not entirely correct. Only the Atlantic common seal and the Pacific
island seal can be regarded as fairly settled animals. Contrarily, the larga,
except for the not so numerous Kuril population, should be regarded as
a migratory animal. At least the Okhotsk and Bering populations are
such.

Although pagophilic, the larga nevertheless avoids compact ice floes.
It therefore abandons coastal regions covered with stable ice floes in
winter and returns to them only after the floes break up. The period of
its appearance on the coasts depends on the time they are freed of ice
floes. In the Sea of Okhotsk, in the Yamsk-Siglansk and Tauisk regions,
it approaches the coast for the first time usually at the end of May
(Fig. 157). In the Tigil’sk region (Kamchatka) the first, but still very
rare, predominantly old, animals are sighted in April. On the contrary,
in the Shantar Sea, where the ice floes prevail for a much longer duration,
the larga approaches the coast only in July and at some places only in
August in some years. Until recently, it was not clear where the largas,
especially, the Okhotsk form, spent the winter and early spring months,
having abandoned the coast. Some authors assumed that they probably

262

261

347

winter close to the coasts among the drifting ice floes (Freiman, 1936).
Others held that some part of the Okhotsk population, perhaps even a
significant percentage, winters in the seas of Japan and Kamchatka (S.
Naumov, 1941). The second view is supported by the total disappearance
of the larga in winter from the Shantar Sea, from Sakhalin Bay, and later
even from Tatar Strait.

Wintering of the larga in the Sea of Okhotsk in the 1950s and 1960s
was demonstrated by direct observations. It was assumed that having
left the coastal sections for wintering, the larga localized “in certain
regions confined to the sites of their coastal rookeries” (Tikhomirov,
1961). Mapping of the main concentrations of the larga showed that by
early spring they are at least distributed in “patches” along the periph-
ery of the Sea of Okhotsk, but far away from the coast on the edges
of drifting ice floes in the central sections of the basin (Tikhomirov,
1966).

In fact, in the Sea of Okhotsk the seals of most local populations
spend the winter-spring period quite far from the land but nevertheless in
a “traverse” opposite the coastal sections where the beach rookeries are
formed in the middle or end of summer. The animals forced to abandon
their coastal sections are attracted toward them in winter and, moving
away from the shore ice and in general the stationary coastal ice floes in
the zone of drifting ice floes, they continue to remain as long as the ice

Wy
о
Гм
[п
и
2

Fig. 157. Figure showing the distribution of the larga in the Sea of Okhotsk
during whelping and the main migratory directions to the molting sites in 1969
(by G.A. Fedoseev).

348

conditions suit them in such places as are closer to the coast abandoned
by them. Whether or not the food factor influences the winter migrations
of these animals is not known for certain.

Thus the largas scattered in winter and early spring among drift-
ing ice floes in the Kamchatka belt of the Sea of Okhotsk evidently
represent the Kamchatka population. The most active among them are
sighted in spring along the coasts in the very first favorable conditions.
The larga population remaining on the ice in the northeastern fringe of
the Sea of Okhotsk is evidently confined to the coastal rookeries of. the
Yamsk-Tauisk region. The distances separating the closest beach rook-
eries, according to winter finds of their populations, are at least 150 km
in a straight line, which the animals traverse twice a year.

Perhaps even more distant migrations also occur. There is a view
that the larga population forming innumerable summer-autumn rook-
eries on the Shantarsk Islands and around them depart for wintering
and for reproduction far southward and winter in the southwestern-
most corner of the Sea of Okhotsk, toward the northeastern coast of
Hokkaido between La Perouse Strait and the southern part of Sakhalin,
on the one hand, and the southern islands of the Kuril range on the
other (Tikhomirov, 1961). The autumn-winter courses of the Shantarsk
populations to the sites for wintering and reproduction are not known.
The spring migrations, however, have been confirmed as running along
the eastern coast of Sakhalin (Tikhomirov, 1961, 1966). Thus the large
congregations of “whelped larga” noticed in April in the above region in
the extreme south of the sea, begin to migrate northward in May along
the eastern coast of Sakhalin. In June the largas are concentrated in the
region of the northern extremity of Sakhalin and remain there as long
as the ice floes prevail. The thawing of ice floes and the disappearance
of the seals along with them correspond in time with the arrival of the
larga in the Shantarsk region (Tikhomirov, 1961).

Thus, in order to reach its summer-autumn sites, the Shantarsk pop-
ulation (assuming that the pattern of its migrations is accurate) has to
traverse a distance of nearly a thousand kilometers. It has to negotiate
the same distance back with the approach of winter. The graphic variants
of the larga migrations in the Sea of Okhotsk (Fig. 157) agree to some
extent with these accounts.

The seasonal migrations of the other populations of the Okhotsk
larga are considerably shorter. They are the shortest in the case of some
small local groups concentrated in summer оп the northern and southern
islands of the Kuril range. These groups of larga lead an almost settled
way of life, remaining for more than three-fourths of the year near their

263

349

beach rookeries. They abandon them only at the time of reproduction,
which takes place in the neighboring ice floes (Belkin, 1964).

Most of the larga population of the Bering Sea undertake migra-
tions in the same manner as the Okhotsk larga and are dependent on
the very same factors—formation of compact stationary ice floes in the
winter-spring months on the coasts and migration to the reproduction
sites away from the coasts. These sites represent massive drifting ice floes
with a fairly large number of open water pools and shore ice, protected
simultaneously from storm waves. Most of the Bering population is dis-
posed in the winter-spring months in the eastern part of the sea; the
larga from Soviet waters also reaches there.

Seasonal migrations on one scale or another exist among all the pop-
ulations of the basin of the Sea of Japan. The larga begins to migrate
southward from Tatar Strait in November and disappears from there
finally in December. It reappears there again in spring only after the
thinning of ice floes (Dorofeev, 1936; S. Naumov, 1941). In Peter the
Great Gulf the number of larga increases noticeably in autumn and
decreases in spring while it abandons these sites almost altogether in
summer (Ognev, 1935) and is evidently dispersed widely in the coastal
waters to the north and south; some remain in their original sites.

The Far Eastern pagophobic populations of the Kuril and Comman-
der Islands, eastern Kamchatka, and some other sections of the range
not exposed to prolonged winter glaciation behave altogether differ-
ently. These seals lead a fairly well-settled way of life and undertake
no significant migrations whatsoever from their coastal sections. Such
behavioral characteristics are evidently typical of the local groups of
pagophobic seals inhabiting the Aleutian and Pribilov Islands, coastal
sections of Alaska, British Columbia, and the USA. The Atlantic seal
belongs to the same type as its migrations occur as a rule within the
confines of a limited expanse inhabited by various populations. These
seals migrate under the influence of weather and human persecution,
food availability, need for more isolated sites for reproduction, and other
factors, as exemplified by the behavior of the Dutch populations (Bem-
mel, 1956).

Reproduction. The pagophilic (larga) and the pagophobic (island)
seals mate at different times. Among the former, this period sets in ear-
lier. In the Sea of Okhotsk and the Bering Sea the larga mates soon
after whelping. The presence of an adult male hanging around almost
each suckling female is the first sign of mating activity. These observa-
tions have been supplemented by a study of the genital organs (includ-
ing the presence of sperm in the vagina). The total period of mating

350

in the northern part of the Sea of Okhotsk and in the Bering Sea con-
tinues for about one month, from April 20 to mid-May; the bulk of
the females, however, mate in the period from May 1 to 10. Mating
pairs start to form roughly one week or 10 days before whelping and
break up by the commencement of the period of molt (Tikhomirov,
1964, 1966).83

In the more southern regions of the Soviet Far East, in the southern
parts of the Sea of Okhotsk, and in the Sea of Japan, mating occurs early,
corresponding to the much earlier periods of whelping, i.e., in March and
April (Tikhomirov, 1966a).

The island seals reproducing on the coasts, in the Kuril and Com-
mander Islands and on the coasts of eastern Kamchatka, have not been
adequately studied in this regard. Mating of the Commander Island pop-
ulations occurs at the end of May (Barabash-Nikiforov, 1936) or more
probably in June-July (Marakov, 1964*, 1966*); it occurs obviously at
May end and in June on the Kuril Islands (Inukai, 1942; Belkin, 1964).
Information about the mating season of the seals on the Murman coasts
is not available. It is also not available for the population entering the
Soviet Baltic waters. In general, however, the European, especially the
Scandinavian, populations of the common seal have long been thought to
mate in the autumn months: in September (Holmgren, 1865; Lilljeborg,
1874) or even early October (Collett, 1911-1912). The mating of seals on
the southern coasts of the North Sea has been placed in August (Havinga,
1933; Moore, 1952*, 1955*). Despite the earlier views followed by sev-
eral Soviet authors (Ognev, 1935; Bobrinskii, 1944*; Vinogradov, 1949;
and others), the European common seals mate at the end of July and in
August (Harrison, 1960). Among the American (eastern Canadian) pop-
ulations of the Atlantic common seal, mating occurs not in the autumn,
as was thought earlier, but immediately after lactation ceases, i.e., June
end or early July (Fisher, 1954).

As in the case of other seals, there is no distinct polygamy among
common seals albeit the bulls in one way or another claim the right to
mate with a particular female. Furious and prolonged scuffles evidently
do not occur among them although marks of seizure in the form of

8 In the light of these data it is difficult to agree with the earlier viewpoint (Sleptsov,
1943) that the Okhotsk largas mate late, from June through the first half of August (peak
at July end), when most of these seals have already completed molt. S.P. Naumov (1941)
places the mating of the Okhotsk larga as late as August-September, restricting it thus to
the coastal rookeries in which it is assumed to occur.

E.S. Chuzhakina (1955), using this very information, contradicts herself a few lines
later by correctly pointing out that mating precedes molt, which occurs from April end to
early June.

265

351

scratches and scars are often seen on the skin of the Atlantic as well as
the Pacific pagophobic seals (Allen, 1880; Havinga, 1933; Averin, 1948).
Among the larga, fights for the possession of a female are evidently few.
The literature adduces neither direct evidence on scuffles among males,
nor indirect evidence. The skin of the males reveals no injuries inflicted
by competing suitors. In general, the period preceding the formation
of mating pairs, noticed from the commencement of lactation, proceeds
imperceptibly among largas.

Direct observations of the act of mating of the larga have also not
been reported in the literature. On the coasts of eastern Kamchatka,
it was observed in water on July 7, 1942 (Averin, 1948). The region,
as also the date of observations, indicate that the mating seals in this
case were probably pagophobic island seals. On the coasts of Scotland,
eastern England, and Holland, mating of the common seal was invari-
ably observed in water (Havinga, 1933; Venables and Venables, 1957;
Harrison, 1960).

Mating pairs of island seals (the male pursuing the female in water)
can be seen on the Commander Island coasts in calm weather in June
and July. Both animals loudly whip the water incessantly with the hind
flippers, often dive, and on surfacing sniff each other’s snout (Marakov,
1966* ).

A study of the genitals and vaginal smears convincingly demonstrated
that ovulation of the Bering larga commences even around the 20th of
April, with peak mating in the first 10 days of May (at this time, only
virgin females and those that have not whelped in the current year mate),
while the total duration of the mating season extends from April 20
through May 15; by this last date almost all the females are inseminated
(Tikhomirov, 1964, 1966, 1970). These periods apply to a great extent to
the Okhotsk larga also.

No unanimous opinion was available until recently about the dura-
tion of gestation. The assumption of late (autumn) mating and dif-
ferences of opinion regarding the existence and duration of a latent
period gave rise to controversies in determining the duration of gestation
also. It was assumed as seven months (S. Naumov, 1941), nine months
(Sleptsov, 1943; Chuzhakina, 1955; Venables and Venables, 1955), close
to ten months (Havinga, 1933; Scheffer and Smith, 1944*), or 11 months
(Moore, 1952*, and others). At present, a fairly long (two to four months)
lag in the implantation of the blastocyst is recognized among the pago-
phobic seals (Ph. v. richardi) and largas (Fisher, 1954; Harrison, 1960).
Thus, of the nearly 10.5 to 11 months, on average, from the time the
female mates up to her parturition, active embryogeny lasts only seven
to nine months.

352

Twins have not been reported among largas while they are not alto-
gether rare among the European common seals. Thus, of the 12 births
recorded from 1933 through 1940 in the Bremerhaufen zoological gar-
den, two were twins (Moore, 1955*). Of the 70-80 births occurring annu-
ally in nature in these same years on the coasts between Vezer and the
Elba, about 10 were twins (Juncker, 1940). In the Far East the larga
becomes capable of reproduction at three years of age at the earliest
but the majority about one year later. Some animals even after attain-
ing maturity lag behind until the fifth and even the sixth year of age
(Fedoseev and Shustov, 1964*; Tikhomirov, 1966). Immature females
at four years of age constitute 7% and at five years 6%. All males are
mature from the fourth year of age (Tikhomirov, 1968). The island seals
have not been adequately studied in this respect; it has been assumed
that maturation of the Commander seals sets in at the age of two to
three years (Marakov, 1966*). The European common seals are capa-
ble of reproducing in the third or fourth year of age (Havinga, 1933;
Heinroth, 1958).

By the time maturity is achieved, the testes of the larga weigh at
least 100 g (without appendages), rising to 185 g by the mating season
(Tikhomirov, 19664*).

No data are available on the time of whelping of the common seal
inhabiting the Murman coasts. In the Norwegian waters, however (in the
southern as well as northern parts), the seals whelp roughly at nearly the
same time, from the first half of June to July 20, predominantly at the
end of June (Collett, 1911-1912; Eines, 1964*). Along the southern and
western extremities of the North Sea and in the southwestern Baltic Sea,
pups begin to be seen in most cases in the second half of June to mid-
July though much earlier as well as much later dates of birth are known
(Havinga, 1933; Wagner, 1936; Venables and Venables, 1957; Harrison,
1960; Moore, 1955*, 1965*; and others).

In the Pacific Ocean part of the range, the pagophobic (island) seals
of the more southern populations forming rookeries on the Kuril Islands
whelp from mid-May through the second half of July (Belkin, 1964,
1966; Velizhanin, 1967); those on the Commander Islands mainly do
so in June (Marakov, 1964). The much earlier dates pointed out for
the latter region, 1.е., from April end through early May (Barabash-
Nikiforov, 1936), are evidently not wholly accurate or pertain to the
pagophilic larga.

Largas whelp in a much earlier winter-spring season. Those on the
southern boundaries of the range in Peter the Great Gulf whelp right in
February (Ognev, 1935; Nikulin, 1937) and even at the end of January
(Pikharev, 1948). According to the latest data (Kosygin and Tikhomirov,

266

955

1969), the period of whelping in this region extends “roughly from the
middle 10 days of February and includes the first 10 days of March”. In
Tatar Strait this period shifts to a much later period. A large number of
females give birth there in March and the remainder in April; the total
period of whelping, however, extends from mid-February to the second
half of April. Instances are known of pups being caught there even on
May 19 (Dorofeev, 1936; Nikulin, 1937; S. Naumov, 1941; Pikharev, 1948;
Tikhomirov, 1966; O.A. Salmin).

In the Sea of Okhotsk, the pups appear earlier in the south, mainly
in mid-March, than in the north where whelping in most cases occurs
in mid-April (Tikhomirov, 1966). In general, however, this period even
in the southern part of the sea is also evidently extended since, in the
region of the southern Kuril Islands, newborns were sighted on the ice
floes at March end to the first half of April (Belkin, 1964).

Newborns were noticed at the earliest in the Bering Sea in 1962 on
April 3 (Tikhomirov, 1964) and in 1963 on March 26 (Kosygin, 1966).
Until the beginning of the middle 10 days of April, however, pups were
very rare and most of the adult females caught at this time were gestating.
En masse whelping occurs there from the second week of April and
extends roughly to the end of that month, peaking in the middle of the
month. Newborns were seen in 1963 up to mid-May (Kosygin, 1966). The
solicitude of the mother for her pup in the period of lactation is great;
she does not tolerate even the proximity of people or ships, considering
them a mortal threat to her offspring.

The larga usually whelps every year; in any case, barren females
hardly exceed 10%.8* In the period of reproduction the maternal
population and the bulls are concentrated mainly in the pelagic strip
of drifting ice floes 20 km or less wide (Tikhomirov, 1966b) though in
the Bering Sea they whelp even deep among sparse ice masses (Kosygin,
1966a). The larga selects for whelping ice floes that are not particularly
large and preferably not too hummocky but firm and clean with open
water pools among them. They do not whelp on compact stationary ice
floes (in the coastal fast ice) and within intensively broken massive ice.
Parturition occurs not far from the edge of the ice floe.

These seals do not form concentrated. nurseries and whelp far from
each other; but in Tatar Strait scattered groups of females, sometimes
with up to 20 to 30 pups in the range of vision, have been observed. In
the Sea of Okhotsk and in the Bering Sea they remain more scattered,

81 According to Е.А. Tikhomirov (1966), it is roughly 5% of the eligible females; Gol’tsev
and Fedoseev (1970) assume it to be 15% for the Okhotsk population and 8% for the Bering
population.

267

354

usually at a distance of at least 0.5 km from each other but sometimes
even farther apart. Thus no more than two or three suckling females with
pups are seen in 1 km? (usually with an adult male alongside). On the
whole, the area of the ice floes thinly populated with animals extends for
150-200 km? (Tikhomirov, 19655*). Even the term “rookery” cannot be
applied to such thinly scattered seals.

The pagophobic Commander Island seals undergo parturition some-
times on the coastal rocks or in sand spits which are sometimes even
flooded during high tide so that the pups in such cases are “introduced”
to the water immediately after birth (Marakov, 1967). The newborn is
quite prepared for this: it is born with a short hair coat, having shed the
preceding embryonic fur in the mother’s womb. The island seals, also
on the Kuril range and outside the USSR waters on the Aleutian and
other islands, undergo parturition under fairly similar conditions. The
seals inhabiting the more southern regions of the Pacific Ocean coasts
of North America whelp on sandy-pebbly shoals (especially in the estuar-
ine sections and the lower courses of rivers) and also on small rocky islets
and reefs along the continental coast. The Atlantic seals along the-coasts
of the North Sea, in the southwestern Baltic, and in all the other parts of
the range, including Soviet western Murman, undergo parturition under
nearly similar conditions.

Growth, development, and molt. Newborn largas weigh 7.5-8 kg,
somewhat more, possibly up to 10 kg in some cases. The body length
in a straight line (Lcv) varies from 65 to 80 cm (length measured up to
tip of tail along the body curvature varies from 75 to 90 cm and pups
already somewhat grown up to 110 cm). They have the typical white coat
of dense, long, silky hairs of almost pure white color with creamy tones
and resemble the white pups of other pagophilic seals, especially of the
harp and ribbon seals.®° Their teeth still sit deep in the alveoli and are
covered by the gums. Their milk teeth, however, are extremely reduced
and may be imperceptible. The subcutaneous adipose tissue has almost
not yet developed.

The pups of the Atlantic common seal born on the beach almost do
not differ in size and weight from the pups of the larga,®° are devoid of
the embryonic coat, and are born almost covered with the short, smooth,
coarse hair coat that is characteristic of older animals. However, during

85 The hair coat of the newborn in the region of Peter the Great Gulf is a smoky-gray
(Kosygin and Tikhomirov, 1969).

8 The maximum of 15 kg (Havinga, 1933) should be regarded as an exaggeration; evi-
dently this pup had already completed suckling.

355

the embryonic growth of this seal and other ecologically similar pagopho-
bic seals, i.e., Kuril (island) and Richard’s seals, the embryonic white coat
is seen, as in the larga. Just before birth, the hair coat loses strength and
most often is shed immediately before parturition or right at the moment
of birth, more rarely in the first few hours after birth. Only in extremely
rare cases do the pups sport this coat for a few days after birth. Yet many
such extraordinary cases have been recorded in the last 20 years (Moore,
1965*; Stutz, 1966).

Since the newborn pagophobic seals have no embryonic coat, they
can enter water within a few hours of birth. As noted above, such pups
may even be born right in water; in such cases the pup nevertheless
scurries toward land with the mother’s help. The pup of the common
seal has a color similar to that of the adult but with a more distinct
monochromatic middorsal strip.

Suckling of larga pups continues (as far as can be judged from the
average birth periods and from the time at which the female abandons
her pup) for about one month. For much of this time the mother remains
on the ice floe, never leaving her pup and displaying extraordinary solic-
itude. In the Bering Sea lactation continues from April 10 to May 10-15
(Tikhomirov, 1964). In this interval the pup adds 12-20 cm and mea-
sures 80-90 cm in a Straight line or 95-107 cm along the body curva-
ture (Kosygin, 1966) ог 101.5 -111 cm (Chapskii, 1967). The pup grows
rapidly during lactation and its weight increases largely as a result of
fat deposition. At the end of suckling the initial weight of the newborn
quadruples to 30 kg or more.

As this period draws to a close, the embryonic coat weakens, is later
gradually shed, and soon replaced by a new smooth, short, and very firm
coat with sparse hair. Opinion on the duration of the primary embryonic
coat is not unanimous. According to some, it remains strong for roughly
two weeks or slightly less (Tikhomirov, 1964) and in the Bering Sea
up to April 15; according to others (Kosygin, 1966), only 5-7 days, the
pups being fully molted at 15 days of age (Fig. 158). Casting of the
embryonic coat proceeds in the very same sequence as in pups of other
species of seals; after a brief latent period during which the embryonic
coat loses strength and the new coat still covered with short and smooth
hairs grows, molting commences and the old embryonic fur is shed in
clumps on large sections of the head, flippers, and tail. In this state, given
the sharp contrast between the dark-colored molted sites and the still
preserved light-colored embryonic fur, the pups appear to sport a white
fur vest (Kosygin, 1966). Later, the hairs on the back and on the ventral
side of the trunk are shed. Most pups of the larga are completely free of
their embryonic fur by mid-May (Tikhomirov, 1964).

268

268

356

Fig. 158. Intensely molting white large pup. Bering Sea (photograph Бу С.М. Kosygin).

Only on completion of molt does the pup begin to enter water and
adjust to the new environment, remaining in it almost up to autumn.
The color of the new hair coat after the embryonic coat has been shed
is very Similar to that of the adult but is also variable with near total
absence of light-colored streaks along the spine.

The total duration of lactation of the Kuril (island) seal is consid-
erably more extended: 3.5 months from mid-May through August end
(Velizhanin, 1967). This is evidently due to the prolonged whelping sea-
son. The individual duration of suckling, though undoubtedly short, is
evidently much longer than the corresponding duration in the larga.
Many newborns of the island seal originally sport a very deep dark,
almost black, coat with diffuse light-colored annular spots. Their body
length along the dorsal curvature (Lc) varies from 94 - 104.5 cm and they
weigh 19-19.4 kg (Belkin, 1964). There is no information on the increase
in these values after lactation.

The duration of lactation among the European common seal (British
population on the east coast) is thought to be three weeks (Harrison,
1960). For seals on the coasts of Holland, the Federal Republic of Ger-
many, and the German Democratic Republic, double this duration has

269

357

been indicated. There is basis for preferring the data of direct obser-
vations on the duration of suckling of the animals born in zoological
gardens, which indicate a figure of six weeks (Heinroth, 1958).

The rapid growth and weight increase of pups of all seals, includ-
ing those of the species under consideration, over a comparatively short
duration of lactation is explained by the very high nutritional value of
the milk. The milk of the common seal has 45% fat and 9% protein
(Harrison, 1960). Later, the growth and development of the youngster
feeding on its own are characterized by slow tempos. It has been indi-
cated (Havinga, 1933) that, after the first year, the Dutch seals grow
to 95.7-113 cm (length in a straight line, Lcv); this is only 5-10 cm
more than the length achieved by the pup of the larga in less than a
month of lactation. Further growth of the larga is illustrated in Table 14
(Tikhomirov, 1968).

The molting of immature (commencing from yearlings, Fig. 159) and
adult largas is mainly confined to the regions of reproduction and only
partly to the beach rookeries where probably some adult animals may
complete molt, if they have not completed it on the ice floes. In the Sea
of Japan, only one molting region is known for certain in the northern
part of Tatar Strait. The molting sites of the larga reproducing in Peter
the Great Gulf are not yet completely known. It may be assumed that
the animals spend a part of their molting period there itself on the ice
floes and complete the molting process on the coastal reefs.

In the Sea of Okhotsk molting colonies have been detected in the
southwestern region of Shelikhov Gulf (in Yamsk Bay), in the northern
part of the Sea of Okhotsk itself, west of Shelikhov Gulf to Tauisk Bay,
and on the eastern coast of Sakhalin. The local groups perhaps molt
also in the proximity of the western coast of Kamchatka. Opinions vary
concerning the westernmost part of the sea, 1.е., Gulf of Sakhalin and
the Shantarsk archipelago. According to some (S. Naumov, 1941), there
are no genuine molting colonies of the larga in the Gulf of Sakhalin;

Table 14. Body length of the larga along the dorsal curvature (Lc), (cm)

Female Male
Age in
years
Range Average Range Average

1 105 - 136 123 113-136 125
2 120-143 134 132-150 140
3 137-150 143 142-160 151
4 141-155 149 159-161 160
5

140-152 149 148-165 162

358

270 Е. 159. Under-yearling larga. Bering Island, May, 1969 (photograph by I.P. Tomatov).

at the very end of the icy season “only insignificant remnants of molted
colonies formed in the other regions” are seen there. In fact, throughout
June the larga is mainly confined to northern Sakhalin, or even farther
away, toward the Gulf of Sakhalin and more so toward—Shantar Sea
[Island], which is blocked by heavy ice floes. Only in the second half of
this month does the larga localize in a comparatively restricted space
between Cape Litke and Men’shikov Island. This region represents the
main residence of the larga in spring, on drifting ice floes all along the
western part of the Sea of Okhotsk. The seal penetrates west of this
region only from the first half of July, by which time most of the animals
have completed molting (Pikharev, 1941). Other authors unreservedly
include, in addition to the above regions, not only Sakhalin Bay, but
also the Shantar Sea [Islands] among the molting regions of the Okhotsk
larga (Nikulin, 1937; Tikhomirov, 1966).

In the Bering Sea the main concentrations of molting largas are seen
in the same two main regions in which whelping occurs: the southeastern
fringe of the ice masses (mainly to the north and northeast of the Pribilov
Islands), the Gulf of Anadyr and partly the eastern coast of Kamchatka.
Molting largas do not congregate in the Chukchi Sea (Tikhomirov, 1966);
there are no accurate data whatsoever on the molting of this seal there.

270

359

The information available in the literature on the period of molting
varies widely. In Tatar Strait molting occurs at April end and in May
(Dorofeev, 1936) or at the end of May (Yu.A. Salmin). In the Sea of
Okhotsk, on the eastern coast of Sakhalin, molting larga juveniles were
encountered (in twos and threes) on small broken ice floes at the end
of May (Nikulin, 1937) and somewhat farther away from Sakhalin but in
the same southwestern part of the sea molting adults were encountered
at the beginning, middle, and end of June (Pikharev, 1941); further, in
1939, an adult larga caught on June 17 was still in the initial stage of
molt while others caught later (up to July 1) were at the peak of molt.
In the northern part of the Sea of Okhotsk the first of the molting largas
were found among the molting bearded and ringed seals from May 20
(Freiman, 1936). On the whole, the molting period in the Sea of Okhotsk
continues, according to some (Tikhomirov, 1961), from the end of April
and, according to others (Fedoseev and Shustov, 1964), from around May
10 to mid-July, i.e., roughly оуега period of 2-2.5 months. |

In the Bering Sea most of the adult largas generally molt during the
same period: from the middle or end of the second ten-day period of
May to the middle or even the end of June. It has been pointed out
that males and females (which for some reason have not undergone par-
turition) begin to molt 10 to 15 days earlier (it is quite possible that
some males molt even during the mating season) compared to females
that have whelped (Tikhomirov, 1964). Young animals, however, molt
somewhat earlier than the adults, from the last 10 days of April (Kosygin,
1966), although the reference to a much earlier period, 1.е., from the first
few days of March (Tikhomirov, 1964), is due to an incorrect understand-
ing (Or pertains to the southernmost populations). More recent authors
(Gol’tsev and Fedoseev, 1970) have indicated that en masse molting of
the larga occurs in June.

Information on the molting of land-loving forms is extremely scant.
Only general references are available on the European-Atlantic seals
which undergo molt in summer (Millais, 1904; Collett, 1911-1912);
young seals molt earlier, in July, and older ones in August and even
early September (Havinga, 1933). The first signs of molting among the
pagophobic island seals inhabiting the Kuril Islands are noticed early in
July. At the end of this month and in early August, intensely molting
animals were noticed on Makaarushi Island; molted animals were caught
in mid-August (Belkin, 1964).

On the Pacific coast of Canada and the USA (especially on the coasts
of British Columbia and Washington state), molting continues from the
first half of August to September end (Fisher, 1952).

271

360

Enemies, diseases, parasites, mortality, competitors, and population
dynamics. The common seal in the Baltic section of the range is threat-
ened by no natural enemies. One of the most important factors responsi-
ble for natural mortality there is stormy weather, which can take a toll of
newborns. Some pups may perish due to unfavorable birth conditions, as
sometimes happens in zoological gardens. Nothing is known about the
natural mortality of the Murman populations of the common seal.

Information on the larga in this respect is more specific. Although
itself quite aggressive, it is attacked at places by carnivores. Bears attack
the larga in many regions on the coasts of the Sea of Okhotsk during the
formation of beach rookeries. Sometimes bears even stack their quarry in
some ravine and cover it with soil. Largas torn apart by bears have been
found time and again along the banks of the Moroshchechnaya River in
western Kamchatka and in some bear dens along the northern coast of
the Sea of Okhotsk, including Amakhton Bay (S. Naumov, 1933, 1941;
Tikhomirov, 1966). There is evidence of wolves attacking the larga. A
body with the internal organs and brain eaten out was found on the same
Moroshchechnaya River with many wolf tracks around it (Tikhomirov,
1966).

Even large predaceous birds can peck a newborn larga to death. An
indirect proof of this is the hovering of the golden eagle and the bald
eagle around seal rookeries. The Kamchatka residents on the coasts of
the Sea of Okhotsk speak of seal fights with eagles. On the ice floes close
to Sakhalin, pups of the larga pecked to death by eagles were also seen
(Inukai, 1942; Wilke, 1954). A similar instance was noticed in the Bering
Sea too on the coast of Alaska (Wilke, 1954). Nevertheless, instances of
predaceous birds attacking pups, less so fully grown largas, are not very
frequent since the breeding sites of the latter are usually quite far from
the coasts; further, the adult animals protect their offspring.

The shark could be regarded as a natural enemy of the larga though
no concrete data are available for recent years. This factor has been cited
for the Atlantic common seal (Sutton and Hamilton, 1932*). At places,
the larga is victimized by the killer whale (Orcinus orca). On the Japanese
coastal whaling base near Abashiri, seals of this species were the most
frequent food item found in the stomach of killer whales caught from
August through October, 1948 (Wilke, 1954).

Abiotic factors are equally responsible for the mortality of the young.
Pups on sparse drifting ice floes among extensive open water pools and
more 50 оп being transported to the fringe are exposed to dual fatal
factors. In highly windy weather the embryonic fur coat sprayed or even
drenched by an icy wave simply cannot protect the pup from cold. A
powerful wave can sweep a pup off an ice floe or even topple a floe with

272

361

pups into the water. Ice floes quickly break up under wave action and are
transported to more southern regions where they soon disintegrate and
thaw. Contrarily, during hummocking of ice floes, pups face the danger
of being crushed. Every year, from April end to the first 10 days of May,
carcasses of larga pups in white fur coat are scattered on the coast of
eastern Kamchatka. Particularly large numbers of them were thrown up
from May 1 through 5, 1942. On the coast of Kronotsk sanctuary, in
Olga Bay, 17 such dead pups were collected (Averin, 1948). These have
been regarded as important food supplements to bears awakening from
winter slumber.

Insofar as the Pacific pagophobic island seals are concerned, espe-
cially those inhabiting the Commander Islands, they have practically no
natural enemies except the blue fox, which attacks newborns only as a
very rare exception (Marakov, 1966*). There are apparently more ene-
mies on the Kuril Islands but practically speaking, neither the bear nor
the wild dog, nor even the fox can inflict harm since the rookeries are
generally located on small islands or on reefs where the seals are inac-
cessible to the carnivores.

Our literature contains no definite data on the diseases of this seal.
Among the external parasites recorded, the louse Echinophthirius hor-
ridus localizes mainly on the back, upper side of the tail, and at the
base of the hind flippers, often in large numbers (Freund, 1933; Moore,
1955*). This parasite is a carrier of microfilaria in the blood.

Information on the helminth fauna of the Atlantic common seal
appeared for the first time in the early nineteenth century (Rudolphi,
1819). Later, the helminth fauna of this seal was studied quite thor-
oughly (Monticelli, 1889; Reie, 1899*; Stiles and Hassall, 1899; Linstow,
1905; Ransom, 1920; Baylis and Daubney, 1925; Lyubimov, 1927; Delya-
mure, 1955; and others) but the helminth fauna of the Far Eastern larga
had almost not been studied until quite recently. Only nine species of
helminths of this seal were known by the early 1960s (Belopol’skaya,
1960). Much progress has been made in recent years in identifying the
helminth fauna of the larga. In 1966-1967, 152 animals were autopsied
for helminthological studies: 116 from the Bering Sea and 36 from the
Sea of Okhotsk. As of date (M.V. Yurakhno), 29 species of helminths
and 4 of their larval forms have been identified in the Atlantic common
seal and larga. From among the trematodes these were found: Orthos-
planchnus arcticus (infects the liver, gall bladder, and pancreas; from
1 up to 69 specimens were detected in a single animal), Cryptocotyle
lingua, Echinostoma acanthoides, Rossicotrema venustus, and Phocitrema
fusiforme (parasites of the intestine), and Pseudamphistomum truncatum

273

362

(liver). From among the cestodes these were found: Diphyllobothrium cor-
datum, D. hianus, D. schistochilus, D. latum, Diphyllobothrium sp., Pyram-
icocephalus phocarum, Diplogonoporus tetrapterus, D. mutabilis, Trigono-
cotyle skrjabini, and Trigonocotyle sp. (all these species of cestodes infect
the intestine). The nematodes infecting the gastrointestinal tract are:
Anisakis sp., Contracaecum osculatum, Phocascaris phocae, Terranova
decipiens, T. azarasi, Terranova sp., and species of genera of Anisaki-
dae; Skrjabinaria spirocauda infects the heart, blood vessels, and lungs;
Parafilaroides gymnurus, P. krascheninnikovi, and Otostrongylus circum-
litus infect only the lungs; the nematode Ph. phocae infects the larga
more often than other seals. The acanthocephalans infecting the intes-
tine are: Corynosoma strumosum, C. semerme, C. validum, C. hadveni, C.
ventronudum, and C. osmeri.

The helminth fauna of the Atlantic common seal and the larga vary
significantly. There are large differences in the species composition of
trematodes and cestodes; there is none common among the 14 species
and, further, these groups of helminths are more abundantly represented
in the larga. The latter is highly afflicted by helminths, especially in the
Bering Sea, where all the animals commencing from yearlings were found
infected by hundreds and even thousands of C. strumosum (there were
more than 10,000 of them in some individuals) and also by a large num-
ber of other helminths, among which the cestode Т. skrjabini was the
most numerous (from a few tens to some hundreds) (M.V. Yurakhno).
The more commonly infected organs were: small and large intestine
(80% of the animals), stomach (72.2%), duodenum (66.9%), and гес-
tum (60%). The following organs were more rarely infected: liver and
gall bladder (7.8%), lungs (6.9%), heart and large blood vessels (5.3%),
and pancreas (0.9%). The most pathogenic helminths of the common
seal include S. spirocauda, parasitizing the heart, large blood vessels,
and lungs.®’

The population dynamics of seals in our territorial waters and in the
seas of the Far East has practically not been studied in recent decades.
The degradation (which was not very intense) of some beach rookeries of
the larga in the 1930s in the region of Tatar Strait and at some points on
the coasts of the Sea of Okhotsk as a result of hunting was minimal and
intermittent and hence led to no particularly serious consequences (see
pp. 364-366). Hunting from ships in recent years in the Sea of Okhotsk

87 Information on the helminths of the species under study published here was specially
prepared for this publication by Prof. S.L. Delyamure and A.S. Skryabin, scientists at the
Helminthological Laboratory, Crimean State University.

363

and the Bering Sea (see р. 365) has likewise exerted no perceptible influ-
ence on the population of the larga. Only some local populations of
pagophobic seals, especially on the Commander Islands, have not so far
regained their earlier population level due to uncontrolled hunting. The
Kuril seal too possibly shares the same fate. The extremely small pop-
ulation of Baltic seals is explained as a result of hunting and the high
degree of economic exploitation of the coasts, particularly the marginal
sections of the range. The small Murman populations at present have
clearly gained the maximum levels.

While calculating the reproductive capacities of larga populations,
the following basic data should be taken into consideration: (1) the quan-
titative proportion of the males and females remaining equal, the mature
animals in the populations of the Sea of Okhotsk and the Bering Sea
average 55.7%; (2) barrenness among the females is not identical in
both populations but is generally not high (8% in the Bering Sea and
15% in the Sea of Okhotsk populations). Thus the annual increment is
about 25% of the total population; (3) the mortality of underyearlings
is high at an average of 42.5% pups and thus the increment in herd is
roughly 14.5%; and (4) a comparison of these indices with the number
of pups points to an increment of not less than 10% of the popula-
tion. However, by the time the newborns become yearlings, the ratio
between the additions and losses is equalized (Gol’tsev and Fedoseev,
1970).

Field characteristics. The adult common seal on Murman can be dis-
tinguished in external appearance from the ringed seal by its much larger
size; slightly longer snout; small, almost black speckles on the skin; and
the absence of a fairly broad monochromatically dark longitudinal band
along the spine. It differs from the harp seal (gray-spotted animals) in the
presence of light-colored ring-shaped patches on the back and the body
flanks and the highly vivid dark color of the flanks in general; moreover,
the harp seal does not come onto the coast. It is easily distinguished
from the gray seal by its shorter snout, absence of a convex profile in the
interorbital zone, and presence of light-colored ring-shaped patches on
the dark-colored background of the skin. Unlike the bearded seal, the
common seal is small and, moreover, spottiness is more sharply manifest
and streak-like gaps visible.

The Far Eastern pagaphobic seals are recognized by their habi-
tat (encountered in the coastal regions which do not freeze as a rule)
and their ability to inhabit the elevated sections of the coast at a dis-
tance of up to 25-30 m from the waterline (the larga is usually found
right in the water), by their size (usually larger than the larga), by the
broad somewhat puffed up snout, and by bright spotty coloration; quite

274

364

often, contrasting light-colored, predominantly distinct ringlets without
blackish-brown fine uneven speckles are scattered on the monochromat-
ically dark-colored background. The adult larga during residence on the
ice floes (in spring) is confined in groups consisting of a female with pup
and an adult male. Later, in the molting period, the seals form hards of
dozens of animals. They differ in coloration from the ribbon seal and the
bearded seal in a variegated, spotted pattern on the skin, from the ringed
seal in fine speckles scattered randomly on a gray background on which
the ringed pattern does not form a continuous grid or lattice, formed by
the fusion of ringlets, as in the ringed seal. Moreover, the ringed seal
is much smaller than the larga and has a relatively shorter trunk and
snout.

In the beach rookeries largas are readily recognized by their close
disposition, their movements (scuffles are frequent in the rookeries), and
mainly from the racket they raise, especially at the moment of seizure,
and their screams, which sound like a cacophany of barking dogs and
bellowing cows (S. Naumov, 1941). The racket can be heard for distances
of 2 km or more in quiet weather.

Unlike the larga, the island seal is quiet and almost does not raise
its voice in the rookery. Only in the case of danger do the females sig-
nal the pups with guttural sounds resembling the “boom” of a bittern
(Marakov, 1966*). It has been pointed out that the island seal lying on
an isolated boulder jutting out not very high above the water usually
lies with the head as well as the hind flippers held high, its silhouette
resembling a boat with a high bow and stern (Marakov, 1966*). Fur-
ther, the larga can also be seen on ice floes with raised hind flippers.
(K.Ch.)

Economic Importance

The European common seal, being extremely few within out waters, is
of negligible commercial importance. This seal is generally not caught at
all on Murman. The pagophobic populations of the Kuril or island seals
have no commercial prospects since their number is negligible, although
the local people use a few of them here and there in Kamchatka and at
some other places.

The larga commands considerable commercial interest. In fact, it is
not the only one among our Far Eastern seals whose reserves, in most
regions, are being exploited much below the population abundance. The
larga has been practically of no interest to the state shipping industry in
the Sea of Okhotsk from the early 1930s. It occupies the last and negligi-
bly small position: in the Sea of Okhotsk in 1937 to 1939 its proportion

275

365

varied from 4.8 to 8.0% (average 6.5%) of the total catch; the bulk of the
catch in these years (as also in the 1960s and 1970s) comprised the ringed
seal, Pacific bearded seal, and the ribbon seal. Over two-thirds of the
catch of larga came in the summer-autumn hunting season in the beach
rookeries of Gulf of Sakhalin, Shantarsk Islands, and partly of Tatar
Strait where 500 to 1,000 animals were caught annually. State hunting in
the beach rookeries commenced in 1934, was quite irregular, and con-
tinued only up to the end of the 1930s in spite of the larga being readily
accessible. Thus, in 1934, some 1,500 animals were caught in 22 days
(Tikhomirov, 1966). The catch from 1937 to ‘1939 was high although
aparently uneven. Hunting was not restored in subsequent years.

In spring, the main season of hunting using ships among drifting ice
floes in the western regions of the Sea of Okhotsk, the larga accounted
for 1.3 to 2.3% (average 1.8%) of the total catch from 1937-1939
(Pikharev, 1941). The reason for this low representation of the larga lay
in the characteristic dislocations of its major concentrations on the ice
floes in the Sea of Okhotsk as also in its behavioral characteristics. In the
early spring the whelped and lactating mothers with pups as also the adult
males are concentrated in the Sea of Okhotsk among compact frozen ice
floes that are difficult for ships to penetrate. Later, however, when the ice
floes become thin and the reproduction period has concluded, the largas
gather in the molting rookeries, which are unfavorable for hunting for
the sole reason that it is difficult to approach the cautious animals: at
the very first rifle shot all the animals disappear into the water where
they are inaccessible to the hunters.

In the early postwar years hunting of larga in the Sea of Okhotsk
intensified perceptibly as a result of enlarging the fleet and the region
of its activity. Thus from 1954-1958, the annual catch went up to 3,100
animals, or 4.3% of the total during intense seal hunting. In the next
spurt of activity the specific proportion of the larga rose further: from
1959 - 1963, an average of up to 6,000 animals was caught (Fedoseev,
1966) or 6.8% of all the seals killed in these years.

From the very beginning of the 1960s, hunting by means of ships
occurred in the Bering Sea too, although the larga was caught there in
the smallest numbers compared with other seal species: in the early years
its proportion in the total catch was only 2.1% in spite of the fact that
its population was sufficiently large to support a higher level of kill and
to reduce the pressure on the other seal species.

In the early half of the 1960s, the total catch of the larga in the
entire Far Eastern basin, including hunting by the local people, went up
10 10,000- 15,000 animals. Its proportion in the seals killed by hunting

276

366

using ships did not, however, exceed 9.4% in the best years (Tikhomirov,
1966a).

Some intensification of larga hunting with reduced killing every-
where of the Pacific bearded seal as well as the ribbon seal and partly
the ringed seal is one of the possible methods of rational utilization of
the seal resources using ships in the Far East. Intense hunting of the larga
would evidently reduce the damage caused by these seals to the salmon
reserves. As already pointed out (p. 339), the larga consumes and dam-
ages quite a large quantity of Siberian salmon, humpbacked salmon, and
other even more valuable types of salmon, which serve as food for the
larga in summer. The largas gather in large herds in the fore-estuarine
sections, directly in the estuaries, and right in the lower courses of rivers
during the period of arrival of salmon for spawning and do not so much
consume them as damage them (especially when the fish are abundant),
by selectively nipping small bits of flesh along the spine. This adverse
role of the larga cannot be ignored, at least at places where fish catching,
preparation, and processing have been organized. The Canadian gov-
ernment offers a handsome reward for killing even one seal (Phoca v.
richardi). In the most important salmon rivers of British Columbia, over
12,000 seals were destroyed over a 10-year period, from 1939 through
1948. From 1941/42 through 1946 alone, some $32,000 were spent for the
destruction of 10,000 seals (Fisher, 1952). The loss inflicted by the seal
to the fishing industry in Alaska, especially in the Mednaya River region,
is nearly 2-3% of the total salmon catch here (Imler and Sarber, 1947).
In the spawning rivers of British Columbia during the arrival of salmon,
these fish constitute about 30% of the food of seals (Spalding, 1964).

The technique of hunting using ships is generally as follows. Motor
schooners capable of negotiating among ice floes set out with several
motorboats on board, which are dropped in the water in the region of
hunting as soon as a sufficiently large number of animals is sighted. The
ship heaves to and each boat (with a crew of three in white masks) is
assigned a particular direction. The ship maintains radio contact with
the motorboats. The teams must not only not lose track of the sighted
animals, but also attempt to approach them within rifle range while
maneuvering among the ice floes. Depending on the situation, the boats
move far away from the ship for several hours or even the whole day. The
skin with blubber is recovered from every killed animal while the rest of
the carcass (skeleton with musculature) is only carried if space permits.

During lactation close approach to the larga depends exclusively on
ice conditions. If the ice is dense and the motorboats cannot negoti-
ate, the ship approaches the animals. The larga can be shot directly

367

from the ship or experienced hunters climbing overboard and jump-
ing from one floe to another can approach the animals singly. Instinct
holds the suckling mother close to her pup in spite of fright. The adult
male in the proximity of the female with a pup is usually the prized
trophy of the hunters. Killing of isolated young (immature) larga dis-
persed on the ice floe poses no problem. The situation changes, how-
ever, at the end of the lactation period. Then hunting becomes quite
difficult.

Until comparatively recently, the larga was killed even in the beach
rookeries. For this purpose, 15 to 20 hunters in two boats dropped from
a ship in the vicinity of a rookery would set out for it taking care not
to frighten the animals. Landing away from the rookery and armed with
clubs, they would approach the animals as closely as possible by crawling
under cover, ready to attack at a signal from the leader. This method was
not always successful but often yieled over a hundred animals. In a suc-
cessful hunt, from the viewpoint of the hunters, 5 - 10% of the animals in
the rookery could be killed thusly, or 500- 1,000 animals throughout the
autumn hunting season (Pikharev, 1941). But not all autumn hunts for
the larga in rookeries were successful (sometimes the frightened animals
escaped before the hunters could reach them); moreover, the killed ani-
mals had to be dressed before high tide as otherwise the waves carried
it away and recovery was difficult.

On the open coasts, where it is practically impossible to approach
the rookery, nets are more commonly used to catch the animals. With
the onset of darkness, one group of hunters in a boat carrying a sweepnet
comes within 50-75 m of the rookery and carefully spreads the net over
this area. An auxiliary group of hunters, waiting in another boat at some
distance from the rookery, then approaches the net-covered site and both
parties draw the ends of the net toward the coast. Between 200-250 seals
were caught in one such net in the estuary of the Moroshchechnaya River
(western Kamchatka) (Tikhomirov, 1966a).

Typical methods of killing the larga in the autumn rookeries are
practiced in Tauisk Bay by the Orokhets. On the open rocky coasts the
hunters make a preliminary hideout near the rookery, using for this pur-
pose a boat priorly cast off near the site. As soon as the seals have
become habituated to the hideout, the hunters, taking advantage of high
tide, when the animals abandon the rookery, gather in the hideout and
wait for the animals to return. Allowing some time for the animals to
settle down, the hunters then pounce with clubs and quickly kill as many
seals running panic-stricken into the water as possible (Tikhomirov,
1966).

277

368

The Sakhalin Gilyaks have long adopted a very unique but highly
ineffective method, using a boat and a long (4.0-4.5 m) flexible pole to
which a large ski-like float with a harpoon is attached. Dropping this
equipment in the water, the hunter attempts to take it to the closest
diving seal and, when successful, pierces the animal with a powerful
thrust of the harpoon (Nikol’skii, 1889).

In various sections of the vast Far Eastern coast, other hunting meth-
ods are known but none are specific for killing the larga. Two methods
are very common and extensively used. One is practiced from the end
of summer and in autumn. The hunter crawls up to the seal from the
coast, shoots it, and rapidly drops a light boat to catch the wounded ani-
mal; quite often, however, the seal drowns. In spring the hunters from
the coastal villages (mostly from the Bering Sea area) come out in large
motorboats and seek the animal among thin coastal ice in quite the
same way as done by the hunting parties from a ship in the more pelagic
regions.

The skin of the newborn larga (white pup) up to five days of age
does not command the same high price as that of a harp seal (greenish-
white pup) or some other seal born on the ice. The skin of the larga can
be used in collars, caps, and other fur goods in a natural state or dyed.
However, the number of such skins produced in the early spring hunting
season is negligible. Given the prevailing hunting conditions, especially
in the Bering Sea, it is difficult to say whether the hunters in one ship
can catch over 200 undamaged skins of white pups of this species during
the season.

The skins of molted pups (killed somewhat later) and juveniles (one
to two-three years of age) with a more beautiful, brightly spotted color
and relatively thin skin are of much greater value. Because of these
qualities and also because of the relative rarity of such skins, the larga
has long received particular attention from the coastal people, especially
the Chukchis, Koryaks, and Eskimos. They use these skins for making
dresses and generally for more delicate work, in particular for fashioning
women’s apparel.

Until quite recently (1960s), the prevailing fashion for seal skin arti-
cles such as caps, jackets, cloaks, shoes, etc. put the seal skins, especially
of young ones, in high demand in the fur market. The fashion for short,
rigid furs of natural color necessitated catching not the white pups, but
the molted pups and the older juveniles. This is all the more rational as
such skins are more accessible and hunting is not restricted to the brief
spell during which the pups sport the neonatal embryonic fur. Even the
skins of the adult larga are perhaps suitable for this purpose through

278

369

their hair coat is slightly sparser and more rigid, the color less attractive,
and the hide considerably thicker and heavier.

The quantity and thickness of the subcutaneous adipose tissue varies
according to the season and the age of the animal. Newborns are almost
devoid of it but accumulate a 3-4 cm thick fat layer (together with the
skin) in the short lactation period and weigh roughly 15 kg (also with
the skin). The skin with the subcutaneous fat (blubber) of animals in
a transitional age weighs 15-25 kg. The thickness of the fat layer in
adult animals in spring differs little from that of the immature animals
but exhibits sharp seasonal changes. The weight of the blubber of the
larga in spring (especially after lactation and mating) averages 20 kg; it
doubles to 40 kg by autumn (Fedoseev and Shustov, 1964*).

The meat of the larga is used as required by the trade. The demand
for it has been increasing year by year with the expanding animal farms,
particularly in the local coastal collective farms of the Far East. The
meat with bones of a young larga weigh 15-20 kg and of semi-adult and
adult animals 35-40 kg.

Rules governing the utilization of the Far Eastern marine animal
resources have not been properly drawn up to date. This applies to the
larga also, although there is no need for special controls on its hunting.
However, the need to reorganize the hunting activity is quite consider-
able. Any seal providing fur (and hence the larga too) deserves the same
attention as the fur seals.

There is also need for special supervision over the utilization and
study of the less abundant island (pagophobic) seals on the Kuril Islands,
Commander Islands, and along the eastern coast of Kamchatka, all the
more since these seals, as far as is known, do not affect the fishing
industry. (K.Ch.)

Subgenus of Harp or Greenland Seals
Subgenus Pagophilus Gray, 1844

HARP OR GREENLAND SEAL
Phoca (Pagophilus) groenlandica Erxleben, 1777

1777. Phoca groenlandica. Erxleben. Syst. Reg. Anim., p. 588. Greenland.

1778. Phoca oceanica. Lepechin. Acta Academ. Petropol., I, p. 259,
Tables 6 and 7. White Sea.

1785. Phoca semilunaris. Boddaert. Elen. Anim., р. 170. Greenland, !ce-
land.

1811. Phoca dorsata. Pallas. Zoogr. Rosso-Asiatica, I, р. 112. White Sea.

(V.H.)

279

370
Diagnosis

These are relatively large seals. The body length of the adult reaches
2 m (even more in some cases) up to tip of tail along the dorsal sur-
face (Lc). The adult skull is 185-240 mm long. The color of the hair
coat varies (depending on the age and to some extent on the sex) from
spotted gray (with haphazard but not very densely scattered dark, mostly
angular patches in a gray background) to a bright contrasting “wing pat-
tern” with two very large highly elongated dark-colored, almost black
patches, sharply prominent in the light-colored background. This “wing
pattern” is disposed symmetrically on both sides and the anterior tips
converge on the back. In the final dress of the animals the color of
the anterior portion of the head is the same as that of the “wing pat-
tern”.

The skull is quite massive, with thick bones. The upper and lower
processes of the posterior edge of the zygomatic bones are nearly equal in
length; the bony nasal septum in the choanae reaches the posterior edge
of the bony palate. The palate has no significant notches but a central
prominence, turned backward, is usually seen. The bony lobe of the
external auditory meatus is genuflexed forward. Molars and premolars
(except the first) have two roots while the crowns, especially of the lower
jaw, bear well-developed accessory cusps. (K.Ch.)

Fig. 160. A—male harp seal, Pagophilus groenlandica, with the final color of

the “wing pattern”; B—adult female harp seal in the “semiwinged” phase of

color (incomplete pattern of the “wings” covered in sparse spots) (figure by
N.N. Kondakov).

279

371

Description

The external appearance is typical of seals of the genus Phoca but the
adult animals are perceptibly larger than their counterparts in other
species of this genus. The body is almost perfectly streamlined and its
hydrodynamic characteristics improve with growth (Alekseev, 1966). The
claws on the fore flippers are quite massive, blackish-gray, with a distinct
transverse rib formation among adults; on the hind flippers these age-
related bands or segments are less prominent or distinct. The length of
the digits on the fore flippers decreases successively from the first and
the second (which are nearly equal in length) to the fifth.

The whiskers are dark gray, flattened, with wavy edges. The labial
whiskers are usually disposed in 7 rows; each side of the lowermost row
most often has 7 (6 to 9) whiskers, the second to the fourth 9-10 (8 to
11), the fifth 4-9, the sixth an average of 4, and the last an average of
2. Sometimes a lone whisker is seen above the seventh row. The labial
whiskers total 46-47 on each side. The supraorbital whiskers number
most often 3 each but vary generally from 1 to 4. As a rule, there are
two whiskers around the nostrils on each side (Yablokov and Klevezal’,
1964).

Two types of hair coat are common among pagophilous seals. These
are successively the neonatal (juvenile, white) and the definitive form
acquired on shedding it. The first consists of tender wavy fur hairs set
very densely in tufts to form luxuriant and concomitantly dense and long
creamy-white fur. The definitive hair coat of the harp seal in structure,
strength, and notable slant (pile) is the same as in the other seals. It con-
sists of uniformly distributed tufts, usually comprising three categories
of hairs: guard hair (12.5 mm long), intermediary 1-2 (6.2 mm long),
and 4-5 fine and wavy fur hairs (5.5-7.7 mm long). These are disposed
in a definite sequence; intermediary hairs anterior to the fur hairs and
the guard hairs shifted forward even more, covering the rest of the tuft
constituents. A 1 cm? area has 1,700 such tufts with a total number of
about 12,000 hairs (Bel’kovich, 1964).

The color of the hair coat varies greatly with age (Fig. 161) and the
final wing pattern is highly typical and contrasting: one large, long, dis-
tinctly contoured, vivid brownish-black patch each on the right and left
sides of the trunk on a very light, white, almost pure white, or slightly
silvery background. The anterior tips of the patches fuse roughly in the
zone of the scapula on the dorsum and with a slight divergence extend
backward and downward along the flanks; the patches enlarge percep-
tibly in this process with a slight crescent-like form in the midportion.
Later, they gradually narrow and disappear in the sacral zone near the

281

372

Fig. 161. Main changes in age-related skin patterns among harp seals. Top to bot-

tom: gray form (1-3 years old); transitional phase of female (average 3-5 years

old); and final coloration, 1.е., the winged form (5 years and above) (photograph
by R.Sh. Khuzin).

280

‚ 373

base of the tail. On a flat skin they resemble a typical horseshoe ог
are somewhat lyre-shaped. The head almost up to the neck, ear open-
ings, and up to the anterior front is the same color as the “wing pat-
tern”.

Such a coloration is acquired by successive transformation which
proceeds slightly differently among males and females. The color of
the first definitive hair coat (juvenile) acquired on shedding the white
coat (neonatal) consists of a light ash-gray main background (quite
often, slightly darker on the upper side) and usually a few dark gray or
brownish-black spots. These spots vary widely in number, are randomly
scattered, often in small groups with small and very large ones intermixed,
sometimes with fairly angular outlines. This is the color of the young seal
(one to three years).

The white coat at birth (neonatal) in males and females is altogether
identical (see above). The infantile coloration succeeding it on shedding
of the white coat is also typical with no specific differences between
males and females.

The transitional spotted gray color is sported, almost without change,
for several years in spite of annual molt. It shortest life occurs in males:
less than half the generation has it for five years and not more than a
quarter for six years. As far as is known, older males with a spotted gray
coloration do not exist. Females, howeger, are seen much longer in this
infantile coloration, 1.е., up to 10 or more years of age; even at eight
years, their proportion can be 50% (Khuzin, 1964). Stray spotted gray
seals are encountered from time to time even among 14- or even 17- or
20-year-olds (Potelov and Mikhnevich, 1967).

The next transitional phase of coloration is the formation of an
incomplete “wing pattern”. It reveals a mixture of the original features,
1.е., spotted gray (or grayish) and the final “wing рацегп”.8 The life of
this stage too varies in males and females. Only the very beginning of
this phase is generally similar in both sexes.

Very indistinct, highly diffuse darkening is seen initially at the site of
the future “wing patterns”. Their coloration is most vivid on the upper
(internal, or dorsal) edge which is very distinct and sharply demarcated
from the gray background. The “wing pattern” rudiments turn increas-
ingly pale toward the lower lateral edges as though faded and gradually
merge with the light gray main background of the body flanks and under-
side. This shadow of the wing pattern is covered at places by stray, rare,

88 Our special literature used to refer to such 20-year-old females as the “second phase,
with gray wings” (М. Smirnov, 1927) but this phrase was later replaced by the term “‘semi-
winged,” first applied to males in which the pattern was not fully formed (Chapskii, 1952).

281

374

small spots that are new and bright as well as the remnants of infantile
coloration. After the next molt, the “wing pattern” appears more dis-
tinctly in some while the features and the color vividness vary little in
others and, as before, remain more distinct only along the upper edge
and are covered with spots.

At this stage, sex-related differences appear in the very nature as also
the tempo of color variations. In the semiwinged phase with an incom-
pletely formed “wing pattern,” the color transformation process is usually
quite delayed among females. This age-color group commencing from
the first five or six years (although relatively few in these generations®’)
constitutes in succeeding years a perceptible proportion of almost up
to two-thirds of the generation. Even among 17-year-old females, the
number of semiwinged animals is obviously quite large. It is quite pos-
sible that a significant number of females never attain the final “wing
pattern” phase but remain in the “semiwinged” coat to the end of their
life (Khuzin, 1964; Potelov and Mikhnevich, 1967). Most females never-
theless reach the final color earliest at 6-7 years of age (roughly 10%
each of the animals in these age groups). At 10 to 17 years of age the
number of females with a “wing pattern” increases perceptibly to around
two-thirds of the generation.

Detailed descriptions are not available of the successive color gra-
dations on transition from the semiwinged to the winged pattern. It can
only be assumed that this transition is quite simple and sharp since the
first of the youngest females with a “wing pattern” are just a year older
than the first of the animals with a semiwinged pattern.

It therefore appears that some definite quantitative proportions of
females of all coloration phases exist simultaneously in the six- to 14-
year-old generations although the representation of the spotted gray
variety is the least, semiwinged more (one-fourth to one-third), and the
winged variety about two-thirds.”

The transitional coloration among males is characterized by some
specific features. This transition covers only three groups of ages from
five to seven years while seals with the full “wing pattern” are also seen
in these groups. It may therefore be assumed that the transition from

89 According to some authors (Khuzin, 1964), the females acquire the semiwinged pattern
coloration at six years of age; according to others (Potelov and Mikhnevich, 1967), however,
this type of coloration is seen among 10% of even three-year-olds.

% Some differences are noticed in these proportions among various populations. The
White Sea females preserve the infantile (spotted gray) coloration only up to 10 years of age
and the semiwinged pattern up to 16 years; the corresponding ages among the Jan Mayen
females are 15 and 22 years; a similar lag of color transformation is also characteristic of
the Newfoundland females (R.Sh. Khuzin).

282

575

the spotted gray to the final phase is extremely short among males and
often the intermediate semiwinged phase is bypassed. Nevertheiess, the
latter phase in turn can be subdivided into a few stages (Chapskii, 1967).

The first stage is the very dark “wingless” type: the dark spots are
so densely scattered that they cover the main background almost wholly
so that the “wing pattern” is not visible; such animals look like a black,
silhouette from a distance.?! Animals with a faintly identifiable “wing
pattern,” lusterless like a dim shadow, can be seen in this group. The ages
of the males sporting both color variations are wholly identical: roughly
40-43% each of four- and five-year-olds and 15-20% of six-year-olds.

The second stage, forms with a “semiwinged pattern,” is distin-
guished by fully developed contours of the lower edge of the “wing
pattern” and considerably lighter but densely spotted main background
(except for a dorsal, very dark clearance between the “wing patterns”)
and even more vividly colored snout (Fig. 161). The age of such males is
predominantly six years but even five-year-olds (possibly even four-year-
olds) are encountered in this stage.

The third stage of coloration, spotted “wing pattern,” is the final
phase with a sharply distinct posterior boundary in the dark coloration
of the head but still with quite a large number of spots or dabs which are
generally dull and scattered in the bright main background. Such males
are close to the forms with a “semiwinged pattern” in age: more than
one-half of them are six-year-olds, about one-third seven-year-olds, while
the five-year-olds constitute roughly one-seventh.

Thus the transition from infantile to final adult coloration among
males occurs over three years (from the fourth through the sixth inclu-
sive). All the males do not necessarily undergo the intermediate stages of
coloration. The majority enter the “winged” phase, either totally bypass-
ing the intermediate phases or passing through only one or two of the
later stages. Thus the fourth year is the last year of the infantile period
in which all the males still sport the spotted gray coloration (or gray
animals), and the fifth year is the first year in which one-fifth or one-
third of them acquire the final wing coloration. All the males develop
the wing pattern by the eighth year (Potelov and Mikhnevich, 1964, 1967;
R.Sh. Khuzin; Chapskii, 1967).

The maximum range of individual color variation is noticed in the
period of growth and formation of final coloration. It is high among ani-
mals preserving the infantile type of coloration as also among mature
females of a much younger age and partly also among males in the
transitional “semiwinged” phase. Among the younger animals in the

91 White Sea hunters call them “ogar,” “salovar,” etc.

376

population bearing a spotted gray skin, it is difficult to find animals
in which the disposition, form, density, and even the color of the spots,
and partly the main background are totally identical. There are very light-
colored animals, sometimes with dense variegated spots and sometimes
with extremely few spots; there are several variations between these types.
Among the females with a prominent “wing pattern,” their contours,
superimposition with stray spots, and pattern of spots vary markedly.
Among the males in the transitional phase to the final form of “winged”
type of coloration, all possible variants of dark aberrations and spot
patterns are encountered.

The skull (Fig. 162), seen from above, is somewhat similar to that of
the large larga (Phoca vitulina largha) but the zygomatic arches do not
protrude markedly into the sides: the width at the zygoma in adults is only
slightly more than that of the cranium measured” between the mastoid
processes. It constitutes 90-110%* (x = 102%) of the mastoid width

We

ye
uy

lg
( Ky". er:

My, Wess :

Fig. 162. Skull of the Greenland (harp) seal, Phoca (Pagophilus) groenlandica
(figure by N.N. Kondakov).

% The craniometric data are from the author’s materials with some corrections by
R.Sh. Khuzin (1967a*) marked with and asterisk.

283

ЭЙ

or about 55.5% of the condylobasal length. The interorbital constriction
varies from 9.5- 13% of the mastoid width. The width of the rostrum
(at the level of the upper canines) varies from 23-36% (x = 29%)*.
The length of the facial portion (up to the uncinate processes of the
pterygoid bone) is roughly one-half (x not over 52%) of the condylobasal
length.

The basic skull features of the species mainly occur in the basal
side of the axis of the skull and also in the structure of the lower jaw
and the zygomatic bones. The midportions of the posterior edge of the
bony palate are shifted relatively far back and are fairly at level with
their lateral sections without forming a distinct palatal notch; a reverse
situation is also seen quite often when, at the end of the median suture, a
perceptible inverse projection is formed. This formation is promoted by
the longitudinal bony septum in the choanae reaching the posterior edge
of the bony palate and quite often extending slightly beyond. In such cases
the rear section of the palate is in the form of two highly flattened fused
arches. The zygomatic process of the temporal bone is turned forward
and not perceptibly enlarged. The choanae are low (height roughly one-
half their width).

The tympanic bullae are roomy, with quite complex contours, and
rounded-triangular in the horizontal plane. The angularity intensifies
somewhat with age. Their width (together with the lobe of the audi-
tory meatus) is greater than their length and averages slightly less than
20% of the condylobasal length. The bony lobe of the external audi-
tory meatus is massive and genuflexed forward. The jugular processes
in most cases project above the skull surface and are bent backward.
The uncinate processes are not bent outward. The nasal bones, consti-
tuting about one-fifth of the total length of the skull, are wedge-shaped
with a highly enlarged anterior tridentate margin. The nasal processes
of the maxillary bones extend along the nasal bones usually for not
less than one-fifth the length of the latter. The sagittal crest is almost
absent while the occipital crest is not strongly pronounced. The sub-
condylar process is well developed in the lower jaw and is perceptibly
incurved.

The molars and three posterior premolars and the molar on the
lower jaw bear well-developed accessory cusps spread fanlike on both
sides of the main cusp (one each in front and, in most individuals, two
each at the back). In the corresponding teeth of the upper jaw, the
anterior accessory cusp is poorly developed (altogether absent in most
premolars but quite often preserved in the molar).

Skull differences between males and females are not striking but
become distinct on statistical processing of the data. The upper contour

284

378

of the skull profile in the frontal portion among males is in the form
of a perceptibly curved line; this curvature is far less noticeable among
females.” А paired fold on the lateral fronto-sincipital surface from
where the temporal muscles originates is more intensely manifest among
the males. The bony nasal septum in the choanae is more developed
among males while it is often incomplete (with a notch on the posterior
margin) among females. The anterior margin of the nasal bones in males
has very long lateral teeth while the median teeth are more often the
longest among females. The width of the rostrum at the level of the
canines in males is 26-33.5% (M 30%) of the width at the mastoid and
in the females 23-32% (M 27%). The width at the zygomatic arches
quite often slightly exceeds that at the mastoids in males but not in
females. Statistical processing of the data revealed that these and several
Other craniological values are somewhat higher in males than among
females. In general, the skull of the male is perceptibly more massive
and heavier.

Age-related variability of the skull is very high. Along with an
increase in its dimensions and weight, an extremely distinct age-related
feature is the development of its crests and the general surface relief of
the cranium. Among newborns, its dorsal surface has an almost wholly
smooth fronto-sincipital obtuse area angularly turned toward the nose
bridge. The sides of this “angle” from where the paired temporal muscles
emerge are even less prominent. With age, they become increasingly
contoured and are displaced toward the midline. As a result, the anterior
width of the area steadily decreases and the anterior angle becomes
increasingly acute while the marginal folds formed are transformed
almost into a crest and come increasingly closer. Concomitantly, the
very faint rugosity observed along the lambdoid suture only among the
newborn, develops into a sharp crest with an overhung pointed dentate
cornice in the adult.

The age-related changes of other craniological characteristics are
less striking but are readily discernible in statistical processing of the
data. The most significant are the following:** (1) the posterior margin
of the bony palate in the newborn usually has slightly more concave
contours, sometimes with a mark on the notch, the latter filling up dur-
ing growth and even transforming into a reverse fold in the midportion;
(2) the base alone of the bony nasal septum in the choanae in most
young animals reaches (not invariably though) the posterior margin of

33 This is not a characteristic of the race (М. Smirnov, 1929) but is of sex-related sec-
ondary importance (Plekhanov, 1932*; Chapskii, 1952; and others).
% Data pertain to the White Sea population.

285

379

the dony palate; hence, there is an angular notch along its height. In
adult and older animals, however, it is not only filled up but its upper
portion even emerges beyond the margin of the bony palate; (3) the
subcondylar process of the lower jaw is fully developed (i.e., reaches the
vertical of the condylar process), also only in the adult; (4) the tympanic
bullae in young animals are relatively more swollen, with a smoother,
round surface, and more simply contoured than in adults. Their posi-
tion is more firmly fixed with age; in the young they may be disposed
anterior to the rear crest of the articular fossa while they are invariably
posterior to the latter in the adult; (5) the flexure of the upper pro-
file in the zone of the nose bridge becomes apparent with age but is
absent in the newborn; (6) the premolars and molars which have already
emerged from the alveoli are disposed in young animals almost with-
Out gaps, their edges often touching each other and sometimes even
slightly extending beyond the crowns one behind the other. With age,
gaps arise between them as a result of the elongation of the jaws and
become widest by the time total maturity sets in. Tooth wearing becomes
perceptible only in very old animals; in males and females with the
“wing pattern,” the accessory cusps of the crowns become worn to a
variable extent; in much older animals, even the main cusps are highly
worn down but the crowns are not usually completely worn down. The
cusp and partly the side of the canines are worn but fully worn down
canines are almost never found. The incisors in adults are also quite
often highly worn down; moreover, with increasing age, some straight-
ening of their upper row is noticed (the incisors are set anterior to the
margin in an even file while they are arranged in a slightly semicir-
cular fashion in the young); and (7) the apex of the nasal bones (the
depth of their wedging into the frontals) changes with advancing age in
the following manner, as a percentage of the total length of the nasal
bones:

Mean
Wi patozone months ni) yaa beh emia tee tae ee ae 41.5 (n = 28)
Fromvone tosthree уваги eo at oor 43 (n = 20)
Males with “semiwinged pattern” (five to six years old) 44 (n = 12)
Males with “winged pattern” (adults) ................ 44.5 (n = 13)

The skull proportions too vary with age, especially the ratio between
total skull length and width of the cerebral portion (the skull becomes
increasingly longitudinal).

The width of the skull in the region of the zygomatic arches increases
Steadily (toward the time of attaining maturity, it is comparable to and
even Slightly exceeds the mastoid width). The rostral width (level of the

380

canines) expressed as an average percentage of the mastoid width reveals
the following course of changes:

%
Age up to one month ......... 227
One-year-olds ................ 24.5
Two- and three-year-olds ..... 26.2
Mature animals .............. 28.6

In young animals the rostral portion is not only noticeably narrower, but
also considerably shorter and lower than in adults. The cranium reveals
a reverse tendency (in the young it is relatively more swollen; the length
of the tympanic bullae as a percentage of condylobasal length decreases
from the first year to old age by 4%), etc.

The range of individual variation in craniological features is quite
significant even among such animals as reveal systematic changes sexwise.
Thus the index of rostral width which points to a harmonious increase
from one age level to another in animals of nearly the same age (for
example, in males with the “winged pattern’) varies from 26 to 34 while
the index of the interorbital constriction varies from 10.5 to 16.5, index
of the width at the zygoma from 98 to 109.5, etc. Skull elements such as
length of the nasal bones, shape of their anterior notch, contours of the
posterior margin of the bony palate, etc. also vary.

The body length of the adult, fully grown. White Sea males
measured between tip of nose and tip of tail in a straight line
(Lev) is 155-188 cm. However, the length measured along the dorsal
surface (Lc) is 169-205 cm. In the White Sea adult females, Гсу
varies from 153-188 cm and Lc from 167-202 ст? (Khuzin, 1963;
M.Ya. Yakovenko).

The length of the os penis in the adults averages 160 mm.

The total weight of the White Sea well-fed adult can reach a maxi-
mum of 164 kg and that of the skin with blubber removed by the com-
mercial method up to 69 kg (Yu.I. Nazarenko).

Weight of heart 600-960 g; liver 1,500-2,400 g; total length of the
intestine at 1,900-2,870 cm exceeds the body length of the adult 13.6
times (Yablokov, 1963).

The condylobasal length of the skull of adult males (37 animals
from the White Sea) was 199.8 - 235.8 mm (х = 217.0); zygomatic width
107.0- 142.0 mm; mastoid width 113.0- 133.3 mm (x = 122.4); width of

9° 229 cm in an extraordinary case.

286

381

the snout above the canines 30.0 -43.5 mm (х = 36.2); and the smallest
interorbital width 8.8-21.0 mm ( = 15.0).

The condylobasal length of the skull of adult females (27 animals)
from the White Sea was 193.0-223.2 mm (x = 210.6); zygomatic width
105.3 - 130.3 mm; mastoid width 11.4-138.0 mm (х = 119.5); the ros-
tral width at the canines 28.6-40.0 mm (х = 32.8); and the smallest
interorbital width 9.1- 16.0 mm ( = 12.7) (Khuzin, 1963, 1967).

The reliability of the craniological differences between males and
females is also confirmed by statistical variance analysis. (¢ more than 3
or 2). The differences are reliable in all the populations: condylobasal
length, width at level of canines, and smallest interorbital width (Khuzin,
1963; Yablokov and Sergeant, 1963; Khuzin, 1967). The difference in
the index of rostral width (as percentage of the condylobasal length) is
reliably or significantly higher in males than in females (Khuzin, 1967).

Within the territorial waters of the USSR, the overall dimensions
and craniometric values of the harp seal reveal no geographic variation
since Our waters are host to animals of one single population that repro-
duces in the White Sea. The breeding center of the other, i.e., the Jan
Mayen population, nearest to the White Sea population, is separated by
over 2,000 km from the breeding site of the latter and the transgressions
of Jan Mayen animals into our waters are very rare (see р. 407). The
differences between the Jan Mayen population, which is isolated from
the White Sea population spatially and in breeding sites, are neverthe-
less very minor (see pp. 387-390) although the Jan Mayen population
is Classified as a special group. (K.Ch.)

Taxonomy

Pagophilus Gray may be regarded as a special subgenus within the genus
Phoca $5. 1., Ph. groenlandica. The seals of Phoca groenlandica differ more
from those of the subgenera Pusa and Phoca s. str. than from species
of the subgenus Histriophoca. The harp seal has several characteristics
proximating it with Histriophoca, the most important being the common
final coloration of the hair coat, i.e., alternation of large sections of very
dark and bright coloration. Some craniological features too are similar:
(1) relatively poorly developed and shortened lower posterior process
of the zygomatic bones, the length of which often does not exceed or
only slightly exceeds the length of the upper process; (2) forward projec-
tion of the temporal bone without enlargement on the anterior margin
of the terminal section of the zygomatic process; and (3) some simi-
larity in the shape of the posterior margin of the bony palate and in
the development of the compact longitudinal septum in the choanae;
etc. There are elements of similarity in ecology also: the seals of both

287

382

these species are confined to drifting ice floes without emerging, as a
rule, onto the coasts, lead a pelagic mode of life, and perform fairly
significant migrations.

Based on these elements of similarity, an attempt was made (how-
ever, not supported by later authors) to combine the harp seal with
the ribbon seal into one genus, Histriophoca (N. Smirnov, 1929, 1935).
Much later, it was proposed that these species be combined (at the level
of monotypical genera) into a subtribe, Histriophocina (Chapskii, 1948,
1955). These species figure in different genera in several contemporary.
works (Scheffer, 1958; king, 1963*; Chapskii, 1963).

The similarity between these genera is also seen in the number of
cartilaginous rings of the trachea (average 43), relative length of the
intestine (13.6 to 14.4 times the body length), variation coefficient of the
weight of some internal organs, and relative similarity of the number of
labial whiskers (Shustov and Yablokov, 1967).

A greater generic proximity between the species compared, than
with any others, is also detected in the response to precipitation
(V.I. Borisov). The evolution of the harp seal from a common ancestor
with the ribbon seal is hardly debatable, but the divergence of these
species should be placed not in the Quaternary period (Davies, 1958),
but in a much earlier period, probably the Pliocene.

There is a view (Winge, 1924, 1941) that the harp seal is more
advanced in some respects than the other species, especially subgenera
Phoca s. str. and Pusa. The more advanced evolutionary features of this
branch are seen in the elongation of the bony palate (also due to the
equalizing of its posterior margin) and in the growth of the longitudinal
bony septum in the same direction (up to the posterior margin of the
bony palate).”

Geographic Distribution

The subarctic and arctic expanses of the Atlantic Ocean and parts of
the northern Arctic Ocean adjoining the eastern fringes of the Atlantic
Ocean.

Geographic Range in the USSR
Constitutes the easternmost part of the general range (Fig. 163). The

range of this species within our territorial regions and the adjoining
international waters covers the entire coastal belt of the Barents Sea

% Such an interpretation of the structural features the skull is somewhat debatable in
the light of some new facts (Chapskii, in litt.).

383

along Murman from the boundary with Norway to the White Sea inlet,
including all the bays and straits, even those penetrating deeply inland,
such as the Kola and Motovsk. Farther east, the range encompasses the
entire Kanin-Kolguev shallow-water zone, Cheshsk and Indigsk bays, the
mainland portion of the sea to the north and northeast of Cape Timansk
of St. Nos, extending along the Timansk coast toward the Pechora Sea
from its northwestern, northern, and northeastern regions right up to
Vaigach itself. However, the southern and southeastern continental sec-
tions of the mainland sections of the Pechora Sea bound roughly by a
line traversing from the Russkii Zavorot to Yugorsk Shar, including the
latter, and also the coastal waters of Dolgii Island, and all other expanses
south of the above line fall outside the limits of the range.

Farther north, the region of the regular habitation of the White Sea
harp seal covers the Kara Strait zone, coastal waters of the northern one-
third and probably also one-half of the western coast of Vaigach, south-
ern extremity and entire western coast of Novaya Zemlya, and almost
all the rest of the wide expanses of the Barents Sea except evidently its
extreme southwestern pelagic portion falling under the warming influ-
ence of the Nordkapp branch of the Gulf Stream and bound by the actual
position of the edges of drifting ice floes in the period of their maximum
distribution. It is difficult to draw any precise boundary here.

287

Fig. 163. Range of the harp seal, Phoca (Pagophilus) groenlandica in the USSR
(K.K. Chapskii).

384

The northern limits of the range in the Barents Sea cover the Franz
Josef Land archipelago and roughly the same latitudes between it and ~
Spitsbergen.

The Kara Sea does not wholly fall within the range: the extensive
mainland expanses which, with some approximation, can be described by
a wide arc from Vaigach roughly to Minin skerry fall outside the range.
The harp seal is evidently not found southeast of this line. Thus it is
wholly absent in Baidaratsk Bay, on the western coast of Yamal, in all
the bays and straits fed by the waters of the Ob’, Taz, Yenisey, Pyasina,
and other rivers (Chapskii, 1938). The harp seal is extremely rare in the
eastern mainland regions of this sea but is more common in the Novaya
Zemlya strip from the Kara inlet right up to Cape Zhelaniya.

In the northeastern regions of the Kara Sea at 77 to 80° lat., the
range extends in the form of a fairly long tongue from the line separat-
ing this sea from the Barents Sea, i.e., through a corridor between the
northern extremity of Novaya Zemlya and Franz Josef Land. Much of
the population entering here is evidently scattered in the western sec-
tions while a small proportion reaches Severnaya Zemlya. In very rare
cases, under a favorable icy environment, extremely small stray groups
are evidently capable of entering even into Vil’kitsk Strait.

The range of the harp seal in the White Sea is nowhere restricted
and covers its entire area, including the bays. In the last century, seals
often penetrated deep into Dvina Bay and were encountered even on
Mud’yug Island (Danilevskii, 1862), 1.е., almost up to the estuary of the
Northern Dvina. They have been seen around those places and also on
the Letnii coast, e.g., in Unsk Bay, even in this century. They transgress
into Kandalakshsk Bay although they usually do not traverse far into
its deep northern portion; however, under unfavorable conditions of the
ice drifts stray young animals may be seen even in its extreme cul-de-sac
sections. In Onezhsk Bay adults are not usually seen except for very rare
transgressions into its northernmost section not far from the Solovetsk
Islands; but young ones are seen there at times (rather rarely) (in the
years of broken drifting “young” ice).”” In Mezensk Bay this seal is quite
common except perhaps for the extreme coastal waters along the south-
eastern fringe.

A feature of the range of the harp seal is its sharply manifest dynamic
character caused by the migratory nature of the animal. These seals are
not encountered simultaneously throughout the entire expanse of the
range. They cover parts of the range in a certain sequence of regular

97 The “invasion” of molted juveniles in 1966 following drifting “young” ice pools in a
reverse direction against the general flow, was an unusual event (see page 425).

289

385

seasonal migration of the entire population or its individual age and
sex groups. From late autumn throughout winter and in early spring all
the harp seals are concentrated at the inlet to the White Sea and in
the adjoining sections of the Barents Sea, from western Murman to the
Pechora Sea. The range undergoes maximum constriction in the winter-
spring period when almost all the White Sea population is concentrated
in the White Sea and in the very near border sections of the Barents
Sea. In this period the range evidently forms a ring of broad ice floe
fringes whose rough position can be schematically depicted in the form
of an uneven arc, with one end resting on the eastern Murman coast
somewhere in the region of Cape St. Nos (and from there extends in a
narrow fringe along the Murman coast even farther west and, in some
years, up to the Norwegian coastal waters). The other end of the arc
runs northeast beyond Kanin Nos toward the northern coasts of Kolguev
Island and farther in the direction of Novaya Zemlya somewhat taking
advantage of the edges of the ice floe belt. In a prolonged autumn and
late winter, this expanse is more enlarged and extends farther northeast
toward the Kara inlet. Such was the situation particularly in the first half
of the winter of 1966 -67, when the seals were confined in isolated herds
throughout such a long broken edge (Beloborodov, 1969).

From autumn and very early in winter, the seals are often seen close
to the coasts but there is not much information about their encounters
due to the darkness and cessation of navigation. The animals remain
exclusively in the water and tracing out their distribution even in the
White Sea is quite difficult. Only from the beginning of February, when
the period of reproduction is quite close and ice rookeries begin to form,
is it possible to establish the location of the breeding section of the
population. However, even right at the peak of whelping and lactation,
when the female population with offspring is localized in certain parts of
the White Sea, the location is not clearly known of the immature portion
of the population and of those adults which, for some reason, have not
participated in reproduction in a given season.

In April and up to early May, almost all the White Sea population is
even more localized, mainly at the inlet and in the adjoining regions of
Mezensk Bay, but quite often in the neck and even in the central basin of
the White Sea, and partly right in the nearest regions of the Barents Sea,
in the zones of the ice fringe. From mid-May the seals usually desert the
White Sea, move rightward on the fringes in the most proximate region
of the southern part of the Barents Sea, into the region of Kanin and
Kolguev, and farther toward Gusin Land (Novaya Zemlya); later they
even scatter on the new ice fringes of the Barents Sea right up to the
Medvezhii and Spitsbergen islands, moving from the fringe increasingly

290

386

north and northeastward. At the end of July and in August, they reach
the northern limits of the Barents Sea, transgress into the Franz Josef
Land archipelago, in the northwestern regions of the Kara Sea, totally
disappearing at this time not only from the White Sea, but also from the
immense expanse of the Barents Sea.

In the high latitudes, along the edges of the arctic packs to the east
of Spitsbergen and among the sparse drifting ice north and east of the
northern extremity of Novaya Zemlya, the White Sea harp seal is quite
scattered in small groups (sometimes adult males even singly”*) right
up to early September, and later extends southward into the wintering
ground. It is again concentrated at the time of reproduction in the White
Sea region and the southeastern parts of the Barents Sea and small
numbers are even seen in Cheshsk Bay.

This, then, is a general outline of the seasonal dynamics of the range
of the easternmost White Sea populations of the harp seal.

Geographic Range outside the USSR (Fig. 164)

Extends from the central and peripheral regions of the Barents Sea in a
broad strip through the entire northern half of the Norwegian Sea and
Spitsbergen Strait, northeastern and southwestern regions of the Green-
land Sea (to the west and northwest of the Martovsk ice fringe) into
Denmark Strait. The range encompasses the coastal waters of northern
Iceland and encircles the eastern and western coast of southern Green-
land in a relatively narrow strip. To the west of the meridian passing
through Cape Farewell, the range includes Davis Strait and Baffin Bay
(including the whole coastal periphery on the Greenland side as also
on the western Canadian fringes). The seals are encountered in small
numbers in Kane Basin and are more abundant in Jones Sound and
traverse along Lancaster Sound up to the Wellington, Barrow, and the
northern part of Peele straits inclusive. Depending on ice conditions,
advance populations enter the Gulf of Boothia through Prince Regent
Strait. The seals are common along the entire eastern coast of Baffin
Island, the Labrador coast, in Hudson Strait, and in the northern part of
Hudson Bay, descending along it almost up to the Belcher Islands. The
southernmost part of the range encompasses all the sides of Newfound-
land including the Gulf of St. Lawrence, the shelf along Nova Scotia,
and probably the region of the Great Newfoundland coast.

The branch of the range diverging from the White Sea basin into the
northwest along Murman extends rather irregularly around the Varanger

98 Observations in the Kara Sea, southeast of the Blagopoluchiya Strait (К.К. Chapskii).

290

291

и
FOR
ока

Fig. 164. Species range of the harp seal, Phoca (Pagophilus) groenlandica, and the
site of its concentration during whelping and molt (dotted spots) (K.K. Chapskii).

Peninsula and the more western highly rugged Finmark coasts, descend-
ing to the Vesteralen Islands and even to the southern Lofoten Islands
sporadically and only in cold winters and in spring. In extremely severe
winters, аз in the very early years of this century (especially the win-
ter of 1902-1903), the herd of “invading” seals spread out even farther.
Somewhat similar but much smaller “invasions” were observed on the
Norwegian coasts even later.

The presence of stray specimens far beyond the limits of the normal
range in the seas of Western Europe is extremely rare, episodic, at times
even totally improbable. Such was the case of a gestating animal found
floating in the Elba River 500 km upward from the estuary in 1896. Stray
animals were sighted time and again on the coasts of Great Britain, from
Scotland to the Thames. Stray animals were found on the northern coast
of France. There is no reliable information about the sighting of these
seals on the coasts of Sweden. In Norway the transgression of a stray
juvenile into the extreme south was reported at Oslo Fjord in 1936.”
(K.Ch.)

Geographic Variation

The intraspecific structure of the harp seal was discussed even forty years
ago. It has long been known that the entire population of this species is
divided into at least three geographically isolated populations (or herds),

% Data on the episodic transgressions of seals were taken from Collett (1911-1912) and
Moore (1952*).

388

each of which has its own features, extremely restricted areas of repro- `
duction, and different regions of wintering and molt. Based on this, it was
quite natural to decipher some morphological features characteristic of
each of these populations. Even the earliest attempts to identify specific
features of craniological indices (N. Smirnov, 1924, 1927, 1929) recorded
some differences in average values between the different populations.
These differences were subsequently refined.

Only one subspecies is known within the USSR.

White Sea harp seal, or “Lysun,” Phoca (P.) groenlandica oceanica
Lepechin (1778).

This is the largest form of the species.

The body length measured along the dorsal surface (Lc), according
to the accurate data of Khuzin (1963), for males (100) is 169-204 cm
(x = 185), for females (300) 163-229 cm (х = 183). The skull dimen-
sions (according to the same source) are: condylobasal length in males
(37) 200-234 mm (х = 217), in females (300) 200.5-223 mm ( =
209.5); mastoid width in males (37) 113-127 mm ( = 121), in females
(27) 111-124.5 mm ( = 117); rostral width (at level of canines) ш.
males (38) 32-46 mm (x = 36.2), in females (27) 27.5-37.0 mm ( =
31.5).

White Sea, waters of the USSR in the Barents Sea, and Kara Sea.
The White Sea represents the zone of breeding.

Outside the USSR, this species is found in the waters of northern
Norway and Spitsbergen and western and northern parts of the Barents
Sea.

* OK *

Outside the USSR, usually only one subspecies is recognized, 1.е.,
the Newfoundland harp seal, Phoca (P.) g. groenlandica Erxl., 1777; dur-
ing reproduction and molt, these are concentrated in two sections of the
range: slightly northeast of Newfoundland Island and in the Gulf of St.
Lawrence.

According to the accurate data of Khuzin (1963), the body length
of males (83) is 152-195 cm (5 = 176.5), of females (127) 156-201 cm
(х = 175.5); condylobasal length in males (39) 200.0-219.5 mm ( =
208.5), in females (41) 191.0-219.0 mm (х = 204.5); mastoid width in
males (40) 109.0- 124.0 mm (х = 117.5), in females (41) 109.0- 123.5 mm
(< = 116.0 mm); rostral width in males (40) 30.0- 40.5 mm (х = 33.5),
in females (41) 27.0-36.0 mm (x = 31.5).

Statistically, the most reliable differences between the White Sea and
Newfoundland populations are seen in the overall body length (¢ = 8.52)
and in the length of the palate (¢ = 4.61) as well as in the condylobasal
length and mastoid width of the skull, etc.

292

389

The range includes the northwestern, predominantly subarctic and
arctic Atlantic including Davis Strait, Baffin and Hudson bays, and
regions of the southwestern and perhaps southeastern coastal waters of
Greenland.

The real subspecific differences between the Newfoundland and east-
ern Atlantic populations are also confirmed by the presence of a char-
acteristic genotype in each of these groups (Naevdal, 1965).

Insofar as the Jan Mayen (Greenland) population, which is spatially
well isolated, is concerned, it reproduces in the region of Jan Mayen
Island and is generally distributed in Denmark Strait and the Greenland
Sea; it reveals almost no statistically reliable craniological differences
from the White Sea subspecies (Khuzin, 1963; Yablokov and Sergeant,
1963) and does not differ from the latter biochemically in the protein
polymorphism (Meller, Naevdal and Valen, 1966*).100

Thus this population evidently cannot yet be regarded as an inde-
pendent subspecies; it represents a variety, tribe, or herd closely related
to the White Sea harp seal; in fact, the two were regarded in the past as
essentially similar (N. Smirnov, 1929). (K.Ch.)

Biology

Population. Although the total world population of the harp seal has
decreased very significantly over the long period of its hunting, it was
still regarded as considerable in the mid-1950s. It was approximately, but
evidently overestimated, put at 5.5 million (Dorofeev, 1956) or 4.5 -7 mil-
lion (Scheffer, 1958). Out of the former assessment, 3 to 4.5 million were
regarded as the Canadian-Newfoundland herd. This was followed by the
White Sea population at 1-1.5 million. The Jan Mayen population at
0.5-1 million came third. A total figure of 3-3.5 million is evidently
closer to the real position (Chapskii, 1966).

The harp seal is one of the few species for which the population
has been determined by different methods, including aerial photographic
survey. The latter procedure was used for the first time in the world
in 1927 to count our White Sea herd. Part of the concentrated molting
rookery was photographed, its area determined, and by extrapolation and
assumption the population put at one million (Dorofeev and Freiman,
1928). This operation was repeated in 1928 and the total population of
the White Sea herd was placed at 3-3.5 million (Dorofeev, 1939).

100 At the same time, some other morphological features of the rank of the population
were also identified; differences in body length, in distance between teats, and in number
of asternal ribs and whiskers (Yablokov, 1963).

390

In the second half of the 1930s, new population counts were
made based on different methodological procedures, i.e., by counting
the mother seals (or the pups). The main stock (М. Smirnov,
1928)101 was initially put at 600,000-700,000 (Р.А. Rudakov and
N.V. Provorov). Somewhat later, double this figure or 1,300,000 was
cited (N.V. Provorov).

The first postwar experiment in studying the White Sea population
enabled a rough evaluation of this herd in 1947 at 1,200,000 (K.K. Chap-
skii).!°? A quarter of a century after the above surveys and calculations,
the same aerial photographic survey was again employed, which gave a
figure of 1.2-1.5 million (Surkov, 1957). These figures incontrovertibly
confirmed the sharp (by more than one-half) reduction in the popu-
lation compared with the situation prevailing some 30 years ago. On
the basis of this figure, however, no conclusion was drawn “about the

- adverse effect on the status of the reserves” by excessive hunting (Surkov

293

and Khuzin, 1959)103 and the herd continued to degrade for some time
thereafter. In the 1960s, extremely disturbing warnings about its distinctly
unfavorable position appeared in the press (Nazarenko and Yablokov,
1962; Yablokov, 1962; Yakovenko and Nazarenko, 1962; Yakovenko,
Nazarenko, and Timoshenko, 1963; Yakovenko, 1963, 1967, etc.). The
total population of the herd in the early mid-1960s was put at differ-
ent levels: 400,000 -700,000 (Nazarenko and Yablokov, 1962), 750,000
(Yablokov, 1962), a minimum of 400,000 (Chapskii, 1966), and a min-
imum of 225,000 (Yakovenko, 1963). The last figure is regarded as an
underestimation (Khuzin, 1972*) although the mother population based
on the data of a 1963 aerial photographic survey of rookeries was put
at only 65,000. The total population in all the photographed molting
rookeries in 1962 and 1963 did not amount even to 200,000. The total
of the White Sea herd calculated on this basis did not exceed 300,000 by
1963 (Yakovenko, 1967).

The reduction in total population of the White Sea harp seal in the
1950s is also strikingly demonstrated by the dynamics of the shrinking
areas of its nurseries and molting rookeries (Table 15).

The age-related composition of the productive population (Doro-
feev, 1939) also served as an index of the adverse status of the pro-
ductive proportion of the herd in which animals older than 10 years
became very few in the early 1960s (Khuzin and Potelov, 1963; Khuzin,

101 That is, without pups.
102 Data for the 1930s-1940s have been taken from Khuzin (1972*).
103 Assuming that all the 100,000 animals killed were pups.

293

293

294

Sil

Table 15. Variation in the area of concentration (rookeries) of harp seals in the White
Sea, km? (Yakovenko, 1967)

Year of observation
Type of rookery А ео ава oe crt 28 беби в
1953 1954 1955 1956 1957 1958 1959 1960

Rookeries in the lactation period (nurseries) 294 290 200 167 143 126 130 120
Rookeries in the molting period (molting) 851 27564 58) 58 48) 46 48

Table 16. Total average annual kill of the harp seal in the White Sea and in adjoining
areas of the Barents Sea at five-year intervals from 1947 - 1964 (in thousands)

1947-1951 1952-1956 1957-1961 1962-1964
190.0 125.2 117.6 89.7

1964). Hunting statistics too provided an equally striking account of the
population reduction (Table 16).

AS a result, an agreement was reaced with Norway in 1965 to sharply
reduce and modify the trend of marine hunting in this region. The mea-
sures adopted have already begun to show positive results.

A similar process of population reduction of the harp seal was also
reported in the western fringe of its range where the Canadian population
concentrates in the period of reproduction and molt. According to the
calculations based on aerial photographic surveys in 1950 and 1951 of the
nurseries in the Gulf of St. Lawrence and in the open Atlantic northwest
of Newfoundland Island, the total reserves of these herds were put at
roughly 3.3 million at the beginning of the second half of this century; of
this, some 650,000 were regarded as pups (Fisher, 1952, 1955; Sergeant,
1959). Survey-based calculations gave somewhat lower figures for 1959
and 1960: the total population of pups in both the Newfoundland regions
(in the gulf and in the “front” region) was put at 365,000. The basic stock
(1.е., without pups) of both these herds was, however, put at only 1.2
million (Sergeant, 1963, 1965). The 1964 census,confirmed these figures
(350,000 pups) (Sergeant, 1965).

No census has been done of the Jan Mayen population but it has
been roughly put at a maximum of a million (Fisher, 1954; Scheffer,
1958; Dorofeev, 1965; and others).

With such a dynamic nature of the range of the harp seal, its local
population year round is also highly variable. By changing location from
season to season, the population gathers sometimes almost wholly in
relatively small sections of the range and at other times is scattered over
a wide expanse (see above).

392

Habitat. The harp seal is a distinct inhabitant of cold waters but
avoids the arctic pack ice, preferring the marginal zones and regions of
stable drifting ice floes prevailing year round (as near Jan Mayen Island)
or during low and high tides (as in the neck or inlet zones of the White
Sea). It is this type of biotopic conditions that is selected by this seal in
the periods of breeding and molt. In the course of these biological cycles,
its herds appear deep in the ice massifs quite far from the marginal zones.
In the breeding period it selects large stable ice floes and ice fields even
with hummocks as a solid substratum. It is less choosy in the molting
period and is seen on the fringes of large ice fields as also on groups of
small ice floes if they provide a good link with the water body. This seal
does not use the stationary shore ice (fast ice), as far as is known, under
any circumstances.

Like the other pagophilic seals, the harp seal also crawls onto the
ice floe but resorts to this mainly when the open water pools between
the drifting ice floes on which it whelps are frozen.

At the end of the winter-spring period of reproduction and molt,
the seals move to the fringes of drifting ice floes (at the beginning of
summer) or are confined generally among the fairly sparse ice floes (in
summer). At the same time, it is not quite correct to stress that the “harp
seal spends its life among the ice floes round the year” (Freiman, 1939).
At the end of summer, in autumn, and even in early winter, when these
seals begin and continue to migrate southward and even appear in the
region of breeding, the environment is no longer icebound (Chapskii,
1961).

Pelagic life is highly characteristic of the White Sea harp seal but
nevertheless during the autumn migrations and on Murman even in
spring, it is quite often seen around the coasts, transgresses into the
bays, even those penetrating deep inland, such as the Kola, and may
traverse through narrow Straits, at times extremely narrow, such as the
Zheleznye Inlet (on Loginov Island south of Novaya Zemlya). In the
ice-free period, however, the seals distinctly exhibit a preference for the
coastal strip and do not stray far from the coasts.

Food. The food of the harp seal, including our herd, in spite of
the earlier optimistic assessment (Ognev, 1935) has not been adequately
studied to date; the list of food objects is far from complete for much
of the annual cycle, especially from June through January and February,
when its feeding is very intense.

Among the invertebrates found in its food are the crustaceans,
mainly euphausids (genus Thysanoessa) and amphipods (genera Anonyx,
Gammarus, Themisto, Gammarocanthys, and Others), and also shrimps
(Crangon sp., Sclerocrangon boreas, Pandalus sp., etc.); among the.

295

393

plankton, mollusks of the genera Clio and Limacina. There is no accurate
list of the species of caphalopods consumed, but these are mainly squids
as also cuttlefish. Among the fishes consumed are capelin, polar cod,
navaga, cod, coalfish [pollack], herring, plaice (Hippoglossus), sea bass,
and even goby. At the end of the last and the beginning of the present
century, harp seals fed along the Murman coasts quite regularly in spring
as well as in autumn. This was the period of considerable increase in ice
coverage of the Barents Sea and hence of a sharply reduced ice-free
expanse, which constituted a seasonal contraction of effective area for
the White Sea population of this seal. Fish was evidently the mainstay
here, as confirmed by actual observations.

Earlier investigators may have erred in assuming that the harp
seal consumed a sizable quantity of cod and even chased this fish
away from the Murman coast (Knipovich, 1895; Knipovich, Yagodovskii,
and Zhikharev, 1902; partly N. Smirnov, 1903; Breitfus, 1903) and also
evidently in other regions (Allen, 1880; Wolleback, 1907; and others).
Direct observations and dissection of the animals contradict these views.
In the spring of 1900, from early March through the 20s of April,
along the Murman coast, especially in the region of Kil’din Island, seals
fed intensively on capelin (Mallotus villosus) and the stomach of 34 of
58 dissected animals contained fairly significant quantities of this fish
(Breitfus, 1903). From the end of November, throughout December,
1902, and also in January and February, 1903, herds of seals wandered
in the coastal waters of the western Murman and “their stomachs were
stuffed with herring” (Breitfus, 1906). From the last 10 days of February
through April end, 1905, the arrival of groups of harp seals on the
Murman coasts was associated with the abundance of small coalfish
(Prigorovskii and Breitfus, 1912).

Commencing from the end of lactation and molt, the juvaniles seek
food independently, feeding initially on large components of zooplank-
ton present right among the ice floes. At the end of March the molted
pups almost cease to look for small crustaceans and prefer to rest on the
ice floes, resuming an active search for food in April. In this month their
stomach or intestines mainly contained remnants of euphausids (Thysa-
noessa inermis and Th. газспи) and amphipods (Anonyx nugax, Parath-
emisto sp., etc.) (N. Smirnov, 1903, 1927; Dorofeev, 1936; Sivertsen, 1941;
Chapskii, 1961, 1964; and others). It is possible that the list includes the
pteropod mollusk (sea butterfly) in the more northern regions.

Data are inadequate on the subsequent diet changes of pups on
emerging (or drifting) into the fringes of the Barents Sea and also on
the food of the adults there. At the very beginning of the 1920s, when
the ice conditions along the Murman and northern Norwegian coasts

296

394

were unusual, the pattern of gradual intensification of feeding of the
underyearlings and the enlarged species range of food items used by them
were as follows. Initially feeding on minute plankton, the pups “later take
to fish, at first polar cod ... available abundantly among the ice floes ...
and later take to the food generally characteristic of this species, leaving
no fish that is readily available, be it capelin or cod, surfacing from the
bottom after it, and herring, and others” (N.A. Smirnov, 1903). There is
some exaggeration here too about the cod. Smirnov also stated that “if
herring were plentiful in the Murman, the bulk of ‘skin’ (1.е., harp seal,
K.Ch.), mainly the young ones, would remain there” (N. Smirnov, 1903).
Later, however, the views on this subject were more reserved and mainly
“small pelagic fish or those confined close to the ice fringes” were added
to the list of fishes consumed by the White Sea harp seal (N. Smirnov,
1924, 1927, 1935).104

According to some investigators (М. Smirnov, 1903; Ognev, 1935),
there are no differences in the nature of feeding between underyearlings
and yearlings; according to others, however (Sivertsen, 1941), some dif-
ferences do exist. Fish and deep-water and benthic invertebrates occupy
a prominent position in the food of yearlings in addition to the pelagic
crustaceans On which 1.5-month-old pups feed. Thus, the stomach and
intestines of some yearlings caught in the White Sea in the last 10
days of April, 1934, sometimes contained in addition to the euphausid,
much larger crustaceans (Crangon crangon and Spirontocaris turgida) and
amphipods, capelin being found more often (Sivertsen, 1941). Regard-
less of the foregoing situation, there is evidently no significant difference
in the food of yearlings and adults.

The seals do not feed regularly during reproduction and molt, at
least not daily; many generally remain hungry. Their stomach is often
empty and highly shrunk. Of the hundreds of adult animals dissected at
hunting sites in the 1920s and 1930s (Sivertsen, 1961), food was found in
the stomach of two: March 15, 1929—male (the stomach was filled with
euphausids) and April 3, 1931—female (the stomach contained shrimps).
A similar picture was observed in special investigations conducted at
the beginning of the 1960s in the White, Barents and also Greenland
seas. Not even one of the hundreds of animals examined contained food
remains in the stomach. In most cases the stomach was markedly shrunk
but its wall was very thick. In this period of such physical starvation the
gastric juices were neutral (Shepeleva, 1963). According to other reports,
the lactating females satiate themselves from time to time “if some food

104 The “invasion” in Murman waters of this seal and the disappearance of cod there is
a consequence of sharp atmospheric cooling (Linko, 1912; and others).

395

is available in the nursing region”. The food comprises mainly planktonic
or nectonic crustaceans (Surkov, 1960).

The seals not engaged in lactation or reproduction and living in
this season in the more northern regions of the White Sea, along the
fringes in the adjoining sections of the Barents Sea, and on the open
Murman coasts, evidently feed regularly. Feeding is interrupted only in
the period of molt when the animals mainly rest on the ice floes. In the
years of deep penetration of these seals westward, into the Norwegian
Sea, some instances are known of their being trapped in fishing nets set
at a depth of about 200 m. Two animals were trapped even in the Arctic
Circle, one of them in the region of Vardo even at a depth of 280 m
(Collett, 1911-1912). With inadequate information on the behavior of
such deep submergence, Nansen (1924, 1939) himself pointed out that,
apart from the common fish food, the White Sea harp seal was capable
of surviving on sea bass and flounder as well as on cuttlefish. Thus the
“food spectrum” of the White Sea harp seal is evidently quite broad and
has not been adequately studied.

In the 1920s, when the nurseries were located more northward, on
the threshold of the Barents Sea, it was affirmed that the females at the
end of the breeding season migrated for sometime, especially for feeding,
to the Murman coast (Skvortsov, 1927; Dorofeev and Freiman, 1928). It
is possible that such local migrations did in fact occur since the “young”
ice floes at the time of the cessation of lactation often drifted away toward
the Barents Sea. At the same time, such an interpretation of the move-
ments of mother animals can be the result of inadequate reliable data.
Regardless, such massive migrations have not been recorded in the past
40 years. In a much earlier period, as already pointed out, not only the
adult females, but also other groups of the White Sea population were
generally encountered feeding in considerable numbers on the Murman
coast commencing from late autumn through the end of spring. Harp
seals in particular were noticed here right from early March through the
last 10 days of April, i.e., in the period covered by the formation of nurs-
eries and molting rookeries. Evidently, the severe weather conditions in
winter, as for example in 1901-1902 and some other years, exerted an
unusual influence; in such years the edges of ice floes extended almost
up to Kil’din Island.

The period of intense feeding now commences evidently from May,
immediately after the cycle of reproduction, after which the seals desert
the White Sea to molt. Breaking up into small groups along the entire
edge and among thin ice floes, they feed mainly on macroplankton (the
more massive species of crustaceans) and the polar cod scattered over
immense expanses of the arctic seas. The summer-autumn range of the

297

396

harp seal is mainly dependent оп the regions of distribution of this fish,
which has great food value for the seal. Feeding in the high latitudes
continues all through summer. With the onset of autumn, when the
polar cod begins to form large schools and moves toward the coastal
regions for spawning, harp seals pursue it. This fish is now almost the
mainstay of these seals on the coasts of Novaya Zemlya, Kolguev, and
Kanin, and especially in the northern regions of the White Sea. In the
latter region, apart from polar cod, herring and partly perhaps navaga,
represent important food items. Feeding in the autumn-winter period is
no less intensive than in summer: everywhere in the coastal waters of the
Soviet arctic, concentrations of seals and en masse arrivals of polar cod
are noticed simultaneously (Klyuche,* 1936; Manteifel’, 1943; Chapskii,
1938, 1961; and others).

Home range. The concept of home range is difficult to apply to the
ecological features of this species whose populations not only migrate
widely, but vary sharply in concentration in different seasons. The herd
instinct in the harp seal is quite sharply manifest; it is evident even in
summer when the population is highly dispersed, in small groups of usu-
ally not less than some 10 animals. One such “congregation” in which
the animals are confined sometimes more closely and at other times less
so (although usually not closer than 5-10 m) is separated from another
by varying distances that are not conducive to numerical averaging. Evi-
dently in regions of highly dispersed food objects (for example, the polar
cod), the seals can be found in some cases even singly and far removed
from each other.

During the autumn migrations, the animals gather into much closer
and larger “congregations” but usually do not remain for long at one
place. The age-sex structure of the feeding and migrating groups is not
yet clearly understood; however, there is considerable justification for
assuming that the migrants segregate into young and adults; the latter,
even in high latitudes, then separate into sex groups, which is particularly
noticeable among animals on the threshold of the winter-spring grounds.
Here the concentration coefficient is also highly increased. The maximum
density of animals in the White Sea occurs at the time of formation
of the rookeries (Fig. 165). In 1963, mother seals were concentrated
with their offspring at three sites over a total area of 108.3 km? where
60,000 whelped females were photographed (Yakovenko, 1967). In fact,
the females there must have been somewhat more since not all of them
were basking on the ice floes at the time of the aerial photographic
survey. Assuming that some 10% of them had remained in water and
making a correction of at least 5,000 accordingly, the average density of
disposition of lactating females in the nurseries works out to roughly 600

298

397

Fig. 165. Part of the harp seal rookery on ice floes in the White Sea. Pho-
tographed from an airplane (material of PINRO) [Polar Scientific Research
Institute of Sea Fisheries and Oceanography, named after N.M. Knipovich].

animals per km? or one female in an area of roughly 1,660 m2, 1.е., ава
distance of roughly 40 m from each other.

Actually, however, the mother seals are disposed far more densely
and are not evenly dispersed, remaining in groups concentrated mainly
along the fringes of large or finely broken ice floes that refreeze due
to the formation of young ice floes at the site of open water pools or
crevices. On these new ice floes extending in weird, twisted, and inter-
laced strips, tongues and much broader sections, the females and their
pups gather in far denser groups than in the central parts of the ice

298

299

398

floes. Moreover, drifting ice floes with animals on them generally alter-
nate with “vacant” ice floes (see p. 397). Thus the lactating females in
fact live within some 5-10 m of, or even closer to each other.

In the molting rookeries the density is much higher: 165,000 animals
of mixed composition (adult males, immature animals of either sex in
the age of one year to five or six years, and adult females) covered a total
area of 76 km? in 1963 (Yakovenko, 1967), 1.е., average of 2,200 animals
рег km?. In other words, there was one animal in an area of 455 п?
(average distance of 20 m from each other). In fact, however, in this
case too the molting animals are usually much closer, especially in the
initial, exclusive male preserves where they literally rub shoulders, quite
often in very dense rows almost fringing the edge of the ice floes (see
Fig. 165). The sequence in the selection of biotopes is determined not
only by the requirements under the definitive conditions of reproduction
and migration, but also by the food factor, in particular the nature of the
seasonal distribution of the polar cod, which is almost the mainstay of the
White Sea harp seal and hence a factor responsible for the disposition
of these seals.

Hideouts and shelters. A hard substratum (ice) is used by harp seals
almost exclusively in the period of reproduction (for whelping, lacta-
tion, and partly mating) and molt, i.e., in the early spring (winter-spring)
period. Very rarely, animals are encountered on ice floes even in the
midsummer season in high latitudes (evidently the sick or well-satisfied
animals induced by the excellent weather). Thus these seals reside in
water for an incomparably long part of the year.

This seal never attempts to build hideouts in the snow on top of an
ice floe, as done by the ringed or the Baikal seal. The only thing the
White Sea harp seal is capable of doing is making air holes in the ice
floe, a characteristic of most other species of pagophilic seals.

The mechanism of formation of the air holes evidently has no dis-
tinctive features. The seals resort to making them only in those cases
when the open water pools between ice floes with pups on them begin
to be covered by thick ice. The animals surfacing for respiration pierce
the ice with their head. As long as they do not move to another site, they
use the openings already formed, which are kept unfrozen by the animals
through frequent surfacing and crawling out onto the ice. In the course
of time these air holes assume the form of a low crater initially fringed by
broken bits of ice and later by the water splashed during surfacing. Many
animals can use the same air hole and hence there are usually fewer air
holes in a nursery section compared to the number of lactating females.
Since the ice floes in the breeding region are on the move constantly
with open water pools usually occurring between them, the air holes do

399

not represent the lone means of contact for the animals between water
and air.

The seals hardly use air holes in the much later spring molting rook-
eries, being accommodated by the natural openings in the ice floes in the
form of open water pools, crevices, “gapes,”!° and “partitions,” which
never close for long because of high and low tides, drifting ice floes,
and floe movements caused by winds. In fact, there have been instances
when the open water pools closed in high tide and froze when filled
with ice bits. The seals surprised by hunters found themselves separated
from the water. If, however, the “gape” is not densely or firmly frozen,
the animals can put to advantage their weight and pierce the ice cover
to hide below it. Similarly, the heavy animals can break up even thin
young ice extending in quiet frosty weather over an open water pool
or in cracks and crevices in the ice floes, especially by their combined
might.

These seals resort to no other shelters. Only the newborn in the very
first days of life on the ice floes seek some shelter from foul weather,
creeping time and again into niches or crevices formed before they were
born during the hummocking of the ice floes, or hiding under an ice floe
overhang. Sometimes the pups hide so deeply under the ledges in piled
up ice floes that they readily escape detection.

Daily activity and behaviour. The activity of the harp seal is not stereo-
typical from day to day; it differs in different seasons of the year depend-
ing on the characteristics of the successively changing biological cycles
to which not all the age and sex groups of the population respond to the
same extent. Thus there are periods of high and low activity. Animals
participating in reproduction exhibit hyperactivity, which is somewhat
more prolonged among females than males. For much of this period the
males are even passive (up to the moment when their sexuality peaks,
which is of relatively short duration). Females with pups, however, are
active for at least 2-2.5 weeks of very intense lactation, though with
intervals for whole days, since the pups are suckled not only during
the day but at night, and go into the water many times between suck-
lings.

An extremely high motor activity, increasingly manifest in daylight, is
noticed in the short mating period, accompanied by chasing competitors
and seizures (see p. 400).

105 Discontinuities formed between the frozen masses of ice floes, sometimes compressed
(during compression) and sometimes diverging or enlarging (in low tide) are called “рарез”
by the coastal hunters.

300

400

The last period, 1.е., molt, is characterized on the contrary by maxi-
mum passivity, especially among males who rest in small groups on the
ice for days on end without going into the water. The females, however,
and also the young (commencing from yearlings) are still in the water
at this time and join the males later. At the end of molt, the animals
again enter a period of high motor activity and migrate into the zone of
summer residence and intense feeding. Their “activity,” directed toward |
procuring food in the zone of ice fringes in this period, can be assumed
to be manifested predominantly in daylight hours which rapidly increase.
Nevertheless, their activity does not cease at night.

The summer behavior has not yet been clearly understood. Evidently,
having expended considerable effort on migration into higher latitudes
and feeding well on the way, the animals are now widely scattered in the
northern limits of the range and do not exhibit much activity. The latter
increases somewhat later with the commencement of reverse migrations,
formation of herds, and transition to intensive feeding on fish in the
autumn. Moving from place to place in autumn and early winter, on their
way to the White Sea and in it, the seals avidly hunt for polar cod and
other fishes during the day and also at night. Further, they are capable
of submerging quite deeply, as demonstrated earlier by the example of
some stray animals being trapped in fish nets set at considerable depths
(see below).

The herd instinct is highly typical of the harp seal and is mani-
fest not only in the formation of strictly localized rookeries in which
almost the entire population of a region gathers. It is also reflected in
all the behavioral features of these animals. The seals remain in herds
everywhere, on the migratory courses, in the wintering sites, and in the
feeding grounds. Only in the summer period is some deviation sometimes
seen.

No aggressive tendency whatsoever is exhibited toward man under
normal conditions; on the contrary, with the approach of man, the herd
tends to leave the ice floe. However, the maternal instinct is so intense
among the females that it suppresses the instinct for self-preservation
and they bravely guard their pups without regard for themselves. There
are no reports of any scuffle whatsoever among the animals except during
the mating season.

Adults and immature animals are not heard but a hungry pup calls
its mother with a loud wail, quite similar to the cry of a child.

It is difficult to establish with certainty which of the sense organs
are better developed. Reports that these seals are “frightened” by ship
smoke have no serious basis as the olfactory faculty is the least devel-
oped in all the pinnipeds. Their vision is quite good in water as well as

301

401

on land. Near, rather than distant, sight is probably better on land. The
auditory faculty is evidently well developed. The ability to echolocate is
beyond doubt among harp seals; otherwise, it is difficult to explain how
this animal can orientate itself in water and catch quarry at depth even
under conditions of the polar night. This faculty has been demonstrated
experimentally by recording underwater sounds in a hydrophone (Mohl,

1968; L. Popov and Pleshakov, 1970).

Seasonal migrations and transgressions. In the nature of its migra-
tion, the harp seal has almost no peer in the family of true seals (Pho-
cidae). Only the hooded seal and partly probably the Caspian seal are
comparable, but the latter falls behind sharply in the magnitude of this
phenomenon. Although this aspect of the ecology of the harp seal was
known from the earliest published works, i.e., at least from the last quar-
ter of the eighteenth century, it has not been thoroughly investigated so
far and much is yet unknown.

In particular, all the details of the migrations of a well-fed juvenile
which leaves the breeding site for the first time, are not known for cer-
tain. In fact, the initial migrations of the under-yearlings bear a passive
character depending on the general drift of the ice floes on which the
pup continues to rest for sometimes (up to March end to early April)
after the final molt (i.e., transformed into a gray pup).

Under normal conditions of the White Sea basin with sharp drifts
(i.e., when the ice floes drift northward through the neck into the inlet),
the gray pups present partly on the ice floes and partly in the open water
pools between them, drift in the same direction as the ice floes. Under
the influence of this drifting, the pups ultimately find themselves in the
northernmost regions of the White Sea or even beyond, on the fringes
of ice floes in the southern sections of the Barents Sea. Here the gray
pups feed from time to time and remain for sometime, possibly until they
are overtaken by the molted adults, 1.е., firstly the adult males and later |
the immature animals of both sexes, as well as gestating and wandering
females. All this represents only a working model as accurately as the
actual picture can be deciphered.

One cannot entirely agree that the gray pups abandon “young” ice
floes for arctic latitudes solely guided by some subconscious desire to
reach the north (Danilevskii, 1862). The situation that prevailed in 1966
demonstrated that such is not so. The exceptional complexity of the icy
environment in that year, caused by steady northeastern stormy winds,
disturbed the normal pattern of the ice drifts. The ice floes on which
whelping occurred were not transported into the White Sea inlet nor
into the Barents Sea, but were pressed to the western and southwestern
coasts of the White Sea where they thawed with the onset of warmth. The

402

molted juveniles on these ice floes and in the water around them thus
found themselves not in the northern sections of the White Sea, but on
its western and southern fringes where they strayed in early summer into
totally unexpected places, up to Arkhangel’sk and Kandalaksh inclusive.

By the end of April to early May, when the White Sea harp seal is
quite prepared to emerge from the region of breeding but not yet wholly
joined by the migrating stream, almost all the White Sea population is
concentrated in the northern parts of the inlet zone, on the threshold
of the Barents Sea, and in its southermost sections. Here the seal pop-
ulation is partly confined to large rookeries and partly highly scattered
in the form of small herds and groups on the ice floes as well as in the
water. Their actual disposition, no doubt influenced in the recent past
by hunting, depends mainly on the position of the ice fringes, on the
characteristics of the ice regime, the nature of distribution of the ice
floes, and on other as yet unidentified factors.

As soon as the majority of the seals complete molt and the molting
rookeries in the White Sea are disbanded, the movement of the animals
along the fringes of the ice floes of the Barents Sea becomes evidently
somewhat more active. The herds spread increasingly along the fringes
and, as they recede, move farther northward and ultimately reach the
summer range.

The directions of spring migrations (Fig. 166) do not remain strictly
constant but depend on the contour of the ice fringes. However, these
migrations can be generalized schematically as follows. During May and
June the movement of the seals from the southern parts of the Barents
Sea proceeds initially along an arc approaching quite closely to Kanin
Nos, Kolguev Island, and Gusin Land. Roughly on the latitude of the
latter, in spite of the disposition of dense massive ice floes, the animals
sometimes move north, sometimes to the northwest along the ice fringe
receding gradually northward, and scatter almost up to Medvezhii Island
and to the ice floes surrounding Spitsbergen. When, however, a rather
large batch of the population thus moves west of the meridian of the
Kola Peninsula, the ice floes even to the east of the Barents Sea rise
markedly northward; they open up in July and by August provide access
for the seals into the northeastern regions of the sea.

Following the receding ice floes and holding on mainly to their
fringes, large numbers of the White Sea harp seal reach high latitudes in
August, presumably spreading increasingly predominantly in the eastern
regions of the summer range. However, on their northward journey in
the spring-summer period, they often cannot reach the coasts of Novaya
Zemlya directly which are still blocked by ice floes, although the seals are

303 evidently numerous along the outer edges of this icy belt. In July-August,

403

"ош pue чоцопролАэл jo $1018э1—ф ‘зиоцел8ци ишизие—$ pue ‘8и19$—2 ‘э8иел ay] JO зэперипоа—1
:(нуз4ецо yy) вофрив]изо48 (snjiydo3vg) vo0yg Чеэз diey еэб эйцм ey) JO зиопел8ни ay) BuIMoYsS эла8 ‘99т ‘ЗЫ ZOE

71 IWN Е-7
ef") 1b]

404

а considerable number of them are seen in the broad corridor between
Novaya Zemlya and Franz Josef Land. Some herds penetrate the straits
of this archipelago while a much larger number of them transgress into
the northwestern part of the Kara Sea. Small groups go east, sometimes
very far, reaching the western coasts of Severnaya Zemlya.

Migration in the reverse direction commences in September. Now,
abandoning the zone of the ice fringes, which in the autumn holds no
special interest in the context of food availability, the seals go south and
southwest (and in the western boundaries of the range possibly even
southeast), and probably through the open sea. Unlike the spring migra-
tions, on their reverse course, which has not been thoroughly studied,
the seals determinedly adhere to the Novaya Zemlya coasts. One of the
most important stimuli for their approach to land is the commencement
of arrival of spawning polar cod along the coasts on which all the arctic
seals feed in this period.

In the region of Cape Zhelaniya, harp séals are seen even in Septem-
ber and later at almost all the other points on the west coast right up to
the southern extremity of this vast twin island. More often, they move
from one cape to another and more rarely are seen in the deep bays and
straits. The White Sea harp seals migrate southward in small numbers
along the Kara Sea, 1.е., the eastern side of Novaya Zemlya, emerging
into the Barents Sea through the Kara Strait. Further advance to the
White Sea has not been well traced but nevertheless it may be assumed
that the migratory routes in the Pechora Sea run for the most part away
from the mainland coast, which the herds of seals rarely approach and,
that too, only on the western side of the Russkii bend. They probably
do not transgress deep into the Cheshsk Bay; they are, however, more
common on the northeastern coasts of the Kanin and partly Kolguev
islands.

The seals arrive in the White Sea usually slightly earlier and begin
to form their first nurseries there. In one or the other sections of it,
predominantly on the right bank, these seals are encountered even from
November end to early December, their appearance being also asso-
ciated with the arrival of the polar cod. The autumn migrations, like
the summer-spring ones, are mainly based on the food factor, especially
the schools of spawning polar cod. It is therefore not wholly correct to
interpret the autumn course of the harp seal to the White Sea as exclu-
sively “migration for reproduction” (Freiman, 1939). This phenomenon
is evidently more complex (Chapskii, 1961) since it begins to manifest
at least 5.5 months before the forthcoming actual breeding season and
the phenomenon covers even immature animals. This, if one may say
so, complex stage of stereotyped behavior (countless generations have

304

405

well-worked out the course) in which the feeding seasons and the return
of the animals to their original sites, guaranteeing the performance of
the concluding stages of the annual cycle, has evolved with an accuracy
that is perhaps maximal for biological systems.

In the past, in very cold years, when drifting ice floes in the Barents
Sea came close to the western Murman and the White Sea harp seal
rookeries were disposed in the inlet of the White Sea and the entire
seal population in the spring period gathered in the inlet and in the
comparatively small space extremely close to the Barents Sea bound by
the fringes of drifting ice floes, the seals undertook extensive migrations
along the Murman coast to the west. In particularly icy, cold springs,
they moved not only up to Finmarken, but even to the coasts of northern
Norway; turning round Nordkapp, they approached Lofoten and moved
even more southward. Particularly memorable are the invasions of seals
in 1901-1902 and 1902-1903 when the winters were unusually severe
and ice floes were seen even in May close to the western Murman coasts
and the southeastern fringes of the compact floating ice floes “lay” on
the Murman coast considerably more westward of the White Sea inlet,
almost around Kil’din Island. The large concentration of seals pressed
by the ice floes at this time toward the Murman coasts caused the distant
movement of some herds into Norwegian waters.

In the relatively less snowy years, however, when the waters on the
coasts of the western Murman were not so intensely cooled and the
winter-spring fringes of the ice floes remained far away in the north and
east, transgressions of the seals along the Murman coasts were not so
massive. Until the beginning of the 1930s, harp seals were quite common
in the winter-spring months all along the Murman and were trapped
there at many places in nets. Invariably, the herds initially came from
the east, from the side of the White Sea and, after wandering, invariably
returned again eastward. Their encounters continued usually throughout
the winter-spring period, mainly in the spring, up to May, and sometimes
even in June.

Now, in the 1960s and the 1970s, the nurseries are usually disposed
deep in the White Sea while the spring boundary of the drifting ice floes
in the Barents Sea runs far northward of the Murman. For this reason
and also because of the sharp population reduction on the Murman
coasts, harp seals are encountered comparatively rarely.

The Newfoundland group of seals begins to riove northward from
the regions of reproduction and molt even from the end of April. They
move along the fringes of the ice floes blocking the Labrador coast
toward Davis Strait and their courses branch somewhere at the latitude
of 60° and along 60° W long. A relatively small portion of the population

305

406

continues, as far as the snowy environment permits, to move northward
up to the southeastern region of Baffin Island. Another, much larger
portion, as far as possible, moves through Hudson Strait into the north-
western and eastern regions of Hudson Bay. A third group, representing
the greater bulk of the migrants, deviates into the northeast and reaches
Greenland roughly at the latitude of 65° or slightly more southward;
later, the majority of them proceed northward and, depending on the
situation of the ice floes, diverge radially along the straits of the Cana-
dian archipelago, along the eastern coast of Baffin Island, and north into
Kane Basin.

A small group of the population turns southward up to Cape
Farewell and, probably running round it, reaches the eastern side of
southern Greenland (Sergeant, 1963; Mansfield, 1963).

The reverse migrations, traced slightly better, proceed from Baffin
Bay along two main routes: one runs along the coasts of Greenland up to
its southern tip and from there traverses through the open sea toward the
southern part of Labrador. Another group, gathering from the Canadian
Straits, runs along Baffin Island and taking the branch from Hudson
Strait continues along the Labrador coast. Evidently, at the southern
corner of the Labrador Peninsula, which the seals reach by November,
both the routes converge and deviate again along the Strait of Belle
Isle. Along one of them following the eastern banks of Newfoundland,
the animals turn farther southward. It was formerly assumed (Robinson,
1897; Chafe, 1923; Nansen, 1927*, 1939) that the seals rushed toward the
great fish banks in the southwest of Newfoundland. New information
(Sergeant, 1963) does not, however, confirm this view. Another path
proceeds through the Strait of Belle Isle into the Gulf of St. Lawrence.
It is not clear where the seals of this herd remain from the middle of
January to February end and which courses they take to gather in the
nurseries.

Exact information is not available on the migrations of the Jan
Mayen seals. At the end of the lactation period, the pups drift with
the ice floes and later emerge onto the fringes; the rest of the animals
wander in small herds to the northeast and southwest of the region of the
rookeries. It is difficult to say whether the seals diverge simultaneously
in Opposite directions or these directions alternate in some manner.

Judging from tagged pups, in some years (for example, 1953), the
animals are mainly carried away in one direction toward Denmark Strait;
in other years (especially in 1955), mainly in the same direction, but
also in the opposite direction (Rasmussen and Oritsland, 1964). The
zone of summer-autumn dispersal of the Jan Mayen herd extends along
the fringes of the ice floes blocking the eastern coasts of. Greenland,

407

from Denmark Strait (and probably even from the southernmost part of
Greenland) to Spitsbergen. In the latter region these seals probably meet
with their White Sea kin and sometimes are drawn by them onto the icy
rookeries in the White Sea. Transgressions of stray tagged animals from
one region to another are known.

Reproduction. Although the harp seal forms no harems whatsoever
like the other seals reproducing on ice floes, it should not evidently be
regarded as strictly monogamous. This concept is in general not appli-
cable to the White Sea harp seal. Even in the days preceding mating,
no single male picks up a female for mating as happens in the case of
the larga. In the same manner, and immediately after mating, the male
does not stay with the female nor does he accompany her in the follow-
ing days. The males and females converge only for coitus and the entire
mating behavior of any given pair, including “heat” and contentions over
a female among the suitors, lasts barely for a day.

Not long before the onset of the mating period, which becomes
evident from the growing activity of the males, the latter gather near
the whelped females in small but fairly close rookeries (“monasteries”)
as though in anticipation of mating. When the time comes, they are
aroused from their torpor and these heavy and awkward animals exhibit
surprising activity on the ice floe. They crawl along it, dive into the water
one after the other, surface from the open water pools, noisily chase each
other, as if in play exhibit surprising dexterity, and again dive into the
water. In earlier years it was not always possible to perceive from afar in
this melee, even with binoculars, who was chasing whom—whether the
males were chasing the females or the suitors were chasing each other.
As a result of organized scientific floating stations on ice floes, it has
become possible to study the intimate life of the White Sea harp seals
quite closely and quite completely.

The period of mating is accompanied by powerful excitation of the
productive males and rather serious scuffles occur among competitors
which can even draw blood (R.Sh. Khuzin). Sometimes the competitors
are severely injured with bleeding wounds. The victor remains alone with
the chosen female far from the rest of the animals. Coitus proceeds in
water (evidently more often) as well as ice floes.!°° In the latter case, it
extends for 20-25 min (Yakovenko and Nazarenko, 1971*). It is highly
possible that the scuffles taking place between productive males is a
consequence of the acute competition among them arising from the

106 There are some other views, although hardly substantiated, that the animals do not
mate in water but only on ice floes (L. Popov, 1966).

306

306

408

quantitative disproportion between males and females due to long-time
preferential killing of the latter.

The mating period among White Sea seals sets in at the beginning
of the first week of March and terminates evidently around the 20th of
the same month. Thus the total duration of the mating period is not
long—about 1.5-2 weeks for most of the animals. However, there are
various views on this subject: not more than 2 weeks (Surkov, 1957),
10-12 days (Rasmussen, 1957), about 2 or 2-3 weeks (Dorofeev, 1960;
L. Popov, 1960), and 10-20 days (Freiman, 1939). The views of some
about a late mating season (as against the average). from March end
to mid-April (G. Nikol’skii, 1933) and more so that mating occurred
after the cessation of molt (Sleptsev, 1949) are erroneous. Even in 1902,
the first of the mating pairs on ice floes were noticed on March 10
(N. Smirnov, 1903).

Most of the whelped females mate even before suckling of the pups
has ceased—roughly in the second half of the period of lactation, closer to
its end. The earliest recorded date of mating is March 4 (Sivertsen, 1941).
Rarely, the last of the winged males was observed hanging around suckling
females in 1948 on March 20. Twenty years later, these dates were rendered
more accurate by researchers on the floating research stationsin the White
Sea (Yu.I. Nazarenko, L.A. Popov, M.Ya. Yakovenko). The first pair in
coitus was observed on March 10 and the last on March 24 (Yakovenko
and Nazarenko, 1961). At the beginning of April the testes of the White
Sea harp seals revealed no mature sperm (Surkov, 1957).

Fig. 167. Mother suckling her pup. White Sea, March, 1967 (photograph by
M. Ya. Yakovenko).

409

The conclusion of the mating period is marked by the quiescence of
the males, cessation of scuffles among them, a distinct coolness toward
the females, and their departure with the young ice floes. By this time
(in the second half of March), the males begin to form different herds
(rookeries) initially consisting exclusively of adult males. These very inti-
mate and massive groups, extending for kilometers along the edges of
the ice floes, along the open water pools and partitions, mark the com-
mencement of a new cycle of life, i.e., the molting period.

The period between mating and whelping is about 11.5 months. In
fact, however, embryogeny occurs with a fairly prolonged lag in the stage
of the blastocyst. The growth of the embryo commences roughly 2.5 -3
months later when implantation sets in. Excluding this duration, the
actual development of the fetus extends for slightly over 8.5-9 months.
Until June, no distinct signs of gestation are seen in the womb (Nansen,
1924, 1939; G. Nikol’skii, 1933; Sivertsen, 1941; Fisher, 1955).

Having attained maturity, the females usually undergo parturition
every year although there are no accurate data on barrenness to date.
The view was expressed before (N. Smirnov, 1903, 1927, 1935; Kulagin,
1929) that the females of the White Sea herd do not give birth to young
ones every year.. However, this view was based on incomplete data and
has not been adequately proved. A thorough analysis of the generative
organs leads more to a contrary conclusion (С. Nikol’skii, 1983; Chapskii,
1963): most of the whelped females mate again in the same year. In any
case, there is no large-scale barrenness among the White Sea females; it
does not exceed 10-15% of the mother population in this herd.!°”

The proportion of barren females among the Canadian population too
is not high though it has been put at 10 to 16% (Sergeant, 1966) and even
20% (Fisher, 1952, 1955) for the population of the Gulf of St. Lawrence.

According to the morphological indices of age used at present, males
as well as females retain the reproductive capacity for over 15 years after
attaining sexual maturity. The oldest of the investigated females that
had whelped in a season were aged 20 years while the productive males
included even 25-year-olds (Fisher, 1952; Rasmussen, 1957; Sergeant,
1966; Yakovenko, 1967). Thus it may be assumed that males as well as
females retain their reproductive capacity up to 25-30 years although
under conditions of extremely intense hunting it is indeed a rare animal
that attains this age (fraction of a percentage).

Unlike the other arctic seals, harp seals bring forth their offspring
in strictly localized sections of the winter-spring range, which for the

107 Tn calculating the population dynamics, the barren animals among the females were
frequently not taken into consideration (Yakovenko, 1967).

308

410

species as а whole number four: (1) Gulf of St. Lawrence; (2) region to
the north-northeast of Newfoundland Island close to the Strait of Belle
Isles (seals of the Canadian population); (3) region of Jan Mayen Island;
and (4) the White Sea. The ice regime characteristics in a given year or
even a very prolonged period of this regime exert an influence on the
actual location of the nurseries, which therefore vary from year to year
within certain limits. These locations also change in the course of a given
season under the influence of drifting ice floes.

Seals of the White Sea herd form nurseries depending on the condi- -
tions of the ice formation either in the neck or in the central basin (more
often on the threshold of the neck and sometimes even in Dvina Bay)
or in the inlet of the White Sea. The possibility in some rare years of
a small proportion of mothers giving birth in Cheshsk Bay (Danilevskii,
1862; Zhitkov, 1904; Suvorov, 1913*; Vinogradov, 1949; Dorofeev, 1956)
has not been confirmed and should be regarded as erroneous.

Over the last 30 years the formation centers of White Sea nurseries
have undergone significant changes. In the 1920s, rookeries were seen
almost exclusively in the inlet comparatively close to the fringes of the
ice floes in the Barents Sea. From the mid-1930s (1935 - 1936), however,
they began being formed considerably farther away, toward the south-
west, i.e., in the neck and the central basin. The desire to provide for the
juvenile adequate conditions in which it can grow well until it becomes
self-supporting has compelled the females to select regions much before
parturition with sufficiently large and stable ice floes capable of with-
standing compression and hence able to serve as a reliable substratum for
the pup’s residence for many days. Another invariable condition for the
formation of the nursery is the presence of a fairly dense network of open
water pools, fissures, and gapes, which permits the female to approach
the chosen ice floes while maintaining contact with the water (Fig. 165).

The sum total of these conditions together with the herding ten-
dency (which to a large extent is probably caused by these factors) and
also the sharply manifest seasonal migrations due to concentrations of
the population in a restricted section of the range, necessitate that ges-
tating females gather periodically in large numbers in the regions of
reproduction. Immediately before whelping, they crawl onto the ice to
form nurseries. A nursery may run into several tens of kilometers and the
area several hundreds of square kilometers. Initially, there is only one
nursery (rarely two or more) but over time, under the influence of drifts
and shuffling of the ice floe, the single massive nursery is fragmented.
As a result, at the end of lactation, when the suckling females leave the
young ice floes, several individual groups of juveniles are seen in what

411

was once a large single nursery. Usually, by this time the juveniles are
transported by ice drifts for long distances from the site of whelping.

Under the influence of the prevailing surface flow from the White
Sea into the Barents, nurseries formed initially in the central basin or in
the neck are gradually transported into the inlet where they cease to exist.
There are, of course, exceptions to this rule. An example is the situation
that prevailed in 1937 when ice floes with pups in the final stages of
molt were transported far westward into Kandalakshsk Bay. The very
same phenomenon, far more acutely, was repeated in 1966. This clarifies
the view expressed by Acad. I. Lepekhin (1805) about some retreats in
the spring migrations of the White Sea harp seal juveniles when they
attempted the inner regions of the White Sea.

The animals are disposed unevenly in the nurseries. The gestating
females usually select large ice floes and are disposed initially along
their peripheries. In time, they often crawl toward the center of the
ice fields and thus their distribution becomes more uniform. On the
whole, however, within the nursery the animals occupy not all the ice
floes but only some of them, usually those along the open water pools
and fissures. Therefore, the rookery does not appear compact but has
alternating occupied and vacant sections on the ice floes. Ice floes with
seals densely or sparsely disposed on them alternating with “vacant” ice
floes, cover an extensive area away from the coastal regions of the sea.
The lesser the density of the rookery, the wider its area, and vice versa.
Further, much depends here on the population of the animals, which is
affected by the magnitude of hunting.

Shortened periods of whelping are a characteristic feature. Among
the majority of White Sea mothers, parturition extends for not more
than a week, from the last few days of February to the first few days
of March. In 1968, by March 4, almost all the gestating females had
undergone parturition. П is the rare female that gives birth earlier than
this period (but nevertheless not before the 20th of February) or slightly
later. A survey of the animals in the month of March revealed that of the
total number of white pups (916), the newborn (aged less than a day) on
March 5 were only 12, or 1.4%; on March 7, 0.8%; March 9 about 0.4%;
and on March 12 about 0.3% (Khuzin, 1970). In fact, some very late dates
of the appearance of newborns, for example March 18 and 31 and even
mid-April (Sivertsen, 1941) have been recorded but such instances are
totally sporadic.1°* Evidently, taking into consideration such anomalous
instances, the total duration of the whelping period is sometimes put

108 In 1966, one white female pup with a firm hair coat was detected as late as April 26
(M.Ya. Yakovenko).

309

309

412

at about two months (Smirnov, 1935) and even extended to 2.5 months
(Sivertsen, 1941). However, it is not correct to include in this period the
extreme, absolutely atypical instances.

The exact moment of birth is usually beyond observation although
one such instance has been described (Sivertsen, 1941). Some members
of our hunting expeditions affirm unanimously that births occur very
rapidly and, as a rule, with no particular birth pangs. Nevertheless, an
instance is known of a dead, full-term pregnant female on the ice floes in
the White Sea in 1959 (M.Ya. Yakovenko). Instances are more common
of stillborn pups or those that perished soon after birth.

The birth of twins has not been established although, a century ago,
it was stated that the females produce one pup, often two or possibly
three, based on the fact that hunters sometimes found up to three pups
on the stretch of an ice floe around a single female (Brown,!” 1868).
Similarly, rare instances of two pups cared for by a female equally well
could point to such a possibility but only indirectly (M.Ya. Yakovenko).

Growth, development, and той. The newborn is covered with a long
dense, silky hair coat that is yellowish-white with a faint green tinge;
hence a just-born pup is called a greenling by local hunters. Its length at
birth averages 83 cm (in a straight line, Lcv) and along the dorsal surface

Fig. 168. White pup of the harp seal. White Sea (photograph by A.V. Yablokov).

109 Significantly, however, this author pointed out that not a single hunter saw more than
two fetuses in one womb.

311

413

(Lc) about 90 cm. The subcutaneous fat layer in a newborn is negligible
or almost non-existent. The weight is generally about 7 to 8 kg.

In the first few days after whelping the females are particularly
solicitous of their pups and remain with them almost constantly on the
ice floes. The milk composition of the White Sea mothers, according
to the latest data (Khuzin, 1970), reveals considerable variation: fat
from 13.5-40.2% (average 29.35%), nitrogen 1.21-2.95% (average
1.76%), and protein content from 7.7-18.8% (average 11.17%) [see
also Table 17]. Suckling is frequent, not only in the day but even at
night. The suckling mother lies on her side and the pup alongside her
stomach finds one or the other teat without difficulty. At times, even an
approaching ship cannot coerce a suckling mother to abandon her pup.
Sensing danger, she will crawl away attempting to carry the pup with her,
then abandon it to dive into an open water pool or through an air hole
in the ice but immediately emerge on the ice again. Having overcome
her fright at the ship, she comes close to the pup and again dives only to
reappear on the surface merely a few tens of seconds later. Confronted
with imminent danger, she will not leave the pup out of her sight.

Even in the early period of its growth, the pup exhibits quite good
mobility and traverses considerable distances, evidently not so much in
search of its mother, as to find protection from wind, to which, lacking
adequate fat reserves, it is initially very sensitive (Dorofeev, 1939). Per-
haps, too, there is a natural need for movement. At favored sites, well
protected from winds by an icy projection or the cornice of a hummock,
the pups remain for long periods. Their prolonged resting at one place,
due to the effect of body heat, forms a basket-like oval depression on
the ice. Inside the depression the pup is always dry.

A few days after birth the green tinge of the pup’s coat disappears
and it turns white; hence hunters refer to it as a white pup. It gains
weight rapidly and by early in the second week after birth has accumu-
lated 9-11 kg of subcutaneous fat; its total body weight now goes up
to 17-18 kg. The body length at the end of the white-pup stage aver-
ages 95-96 cm (Lcv) ог 102-103 cm (Lc). The firm white hair coat is
sported for roughly a week or one to three days longer (Sivertsen, 1941;
Chapskii, 1964; M.Ya. Yakovenko, Yu.I. Nazarenko); the assumption of
a long duration of this period, from 9 to 14 days (Dorofeev, 1936), is
hardly correct.

The commencement of a perceptible loosening of the neonatal hair
coat in the White Sea seal occurs roughly on March 8-9. Initially, the
hair begins to weaken slightly, loses its original brightness and purity
of color, and acquires a gray bloom. The gray bloom is often due to
the thinning of the white hair coat (as a result of the pup’s growth and

310

414

Fig. 169. Well-fed white pup of the harp seal. White Sea, early middle 10-day
period of March, 1965 (photograph by A.V. Yablokov).

its dark skin showing through) and the growing new, already pigmented
coat. Often the guard hair of the white coat turns gray at the base and
the pigment penetrates there as long as the hair bulb is not atrophied
(M.Ya. Yakovenko).

This state extends for a week, after which the next stage of molt sets
in (Fig. 170). By this time the white cover has become even sparser and
the growing stubs of new hairs can easily be seen through it. A few days
later, molting becomes even more evident: the white hair that has turned
gray falls out in large clumps almost simultaneously from the head, hind
and fore flippers, exposing a darker, short, and rigid hair coat. Following
this and almost simultaneously, rapidly growing bald patches on the back
and tail appear. The last hair to molt is that of the ventral side and body
flanks. The better fed the pup, the more rapid the molting of the white
coat. In underfed pups, who have lost their mothers early, molting is
not only impeded or even halted, but acquires a different sequence: the
primary coat is retained on the flippers, tail, and head longer than on
other parts of the body.

Lactation ceases in a normally molting pup at three weeks of age
(Fig. 171). The normally fed molting pup at this time is, on average,
106-108 cm (Lev) or 114-115 cm (Lc). Its total body weight averages
32-33 kg, and some even 38-40 kg or more; the subcutaneous fat with
the skin (blubber) weighs 22-23 kg. The fully molted pup is called a

311

415

Fig. 170. Normally fed molting pup and a pygmy (sick pup of the harp seal).
White Sea (photograph by M.Ya. Yakovenko).

312 gray pup. The hair coat is extremely similar in color in yearlings and

two- or three-year-olds but differs in greater density and softness; the
older juveniles also differ in body proportions: relatively larger head and
longer flippers. The main gray background of the skin among gray pups
is darker on the dorsum with brown angular, predominantly small spots
scattered here and there. On attaining roughly one month of age, the
pup under normal conditions enters the water for the first time and
commences an independent life.

The whelped females no longer spend as much time on the ice flow
as necessitated before. Their residence outside water follows a definite
pattern ordered by periodic lactation, depending on the time elapsed
from the moment of parturition, and the weather factors.!!? On pleas-
ant, quiet, and sunny days, 85, 90, and even 94% of the total mother
population can be seen on the ice surface in some sections of the rook-
ery. '!! On the contrary, in stormy weather, this index drops to 36-45%

110 Detailed information on the periodicity of lactation and the general regime in nurseries
was provided by the floating ice stations organized from the end of 1966 directly in the
rookeries. Groups of investigators landed by helicopter on the White Sea ice with tents
and the requisite equipment and carried out extremely interesting studies (L. Popov, 1966,
1967; Khuzin, 1970; Yakovenko, 1970; Yakovenko and Nazarenko, 1971”).

111 According to others (L. Popov, 1966), however, on such fine days, 45-55% of the
whelped females were seen on the ice during the day and up to 70-80% in the evening.

312.

313

416

Fig. 171. Normally molting pup of the harp seal (photograph by M.Ya. Yakovenko).

and on March 7 even further, to 11% (Yakovenko and Nazarenko, 1971*;
L. Popov, 1966).

Even in the early period of lactation, not all the whelped and
lactating females are seen simultaneously in the nursery although they
maintain a definite feeding cycle. The maximum number on the ice
floe varies at different times: evening, midday, or morning hours, but
more often peaks at midday and evening. The maximum percentage
rarely approaches as high as 85-90%; it is usually lower, around 70%.
It decreases in the course of time and the residence of the suckling
mothers on the ice floe dwindles. In the first few days immediately
after parturition, they remain on the ice floe for long periods and feed
the pups most often at 3-3.5 hr intervals even during the night. At
this time of lactation, the pup receives roughly 0.75 liter of milk in
one feed. Later, as the pup grows, feeding becomes less frequent (4-5
times) and the pup then suckles a larger amount of milk per feed (about
1.2 liters). At this time the lactation schedule in most cases is as follows:
(1) at 6:00-7:00 a.m., (2) 10:00-12:00 Noon, (3) 4:00-6:00 p.m., and
(4) 9:00 - 11:00 p.m. Further, suckling continues even after midnight until
the morning feed, as evidenced by the cries of pups and their quiescence
after sometime, evidently after satiation of hunger (Yakovenko and
Nazarenko, 1971*; Popov, 1966). The daily requirement for milk in

314

313

417

the first few days is about 4.5 liters, which later goes up to 5.8 liters
(Yakovenko and Nazarenko, 1971*; Popov, 1966).

The milk of the harp seal contains 42 -44.5% fat and 8.4- 12% pro-
tein (Sivertsen, 1941; Dorofeev, 1960). Table 17 shows the composition
of milk (10 samples) according to the more accurate data of Khuzin
(1970).

With such a high level of well-being, the daily growth of the pup in
a short period of time—from greenling to white pup beginning to molt
intensely—averages 1.4-2.3 kg (Yakovenko and Nazarenko, 1971*).

Pups which have completed normal lactation and molt grow heavy
and round like dumplings; they remain for one more week and sometimes
longer on the ice floe until the white wool is completely shed and the
gray coat begins to grow. During this period of starvation, extending
for about one-and-a-half weeks, the normally fed pups lose, on average,
about 5 kg of body weight, roughly averaging 0.5 kg (Chapskii, 1964) or
0.6-0.7 kg (Yakovenko and Nazarenko, 1971*) per day. Such a weight
loss in no way affects the future well-being of the young one on its taking
to independent living in water.

However, not all pups receive normal feeding; pups are encountered
which have been abandoned early by the suckling mothers or have lost
their mother for other reasons (Fig. 172). If at the age of one week, the
fat deposition in the pup does not exceed 6-7 kg (and their total weight
not over 10-13 kg or slightly more), cessation of lactation inevitably
leads to emaciation and later to the pup’s death.

The subsequent growth of the animals proceeds more slowly but con-
tinues not only in the first few years, but right up to the time of attaining
sexual maturity though at a very slow tempo (Chapskii, 1952; Yakovenko,
Nazarenko and Timoshenko, 1964*; Yakovenko and Nazarenko, 1967;
Khuzin, 1967; and others).

Earlier there were no reliable morphological age criteria and the
period of attaining sexual maturity was reported variously as: not before
two years of age (Nansen, 1924, 1939), at three years (N.A. Smirnov,
1927), not before four years (Bartlett, 1927*; Nikol’skii, 1933), and even
five years (Degerbol and Freuchen, 1935). At present, when the age of

Table 17. Composition of the milk of the harp seal, % (Khuzin, 1970)

Fat Nitrogen Protein Ash

Min Max Mean Min Max Mean Min Max Mean Min Max Mean
13:53) 13922400 2935 [22:95 176 ty Le Ol 1117: 01147 O82

314

418

Fig. 172. A typical underfed pup (dwarf) of the White Sea harp seal with remnants
of embryonal (white) hair coat on the head and flippers. White Sea (photograph
by M.Ya. Yakovenko).

a given animal can be accurately established within a year, the age at
which reproduction commences has been well substantiated.

The period of maturity among the female harp seals extends for
several years. In the White Sea population some females ovulate for the
first time even at the age of three years, with the percentage of such
early maturing females reported variously as 5-12 to 15.5 (Yakovenko
and Nazarenko, 1967) or 30% (Chapskii, 1963). According to these
same data, about 50% mature by the age of four years but slightly
less (36%) according to other authors (Yu.K. Timoshenko). The rest
of the females mature at five or even six years (Khuzin and Timoshenko,
1968* ) but some, however, at seven years (Fisher, 1954; Rasmussen, 1957;
Sergeant and Fisher, 1960; Sergeant, 1966; Yakovenko and Nazarenko,
1967; R.Sh. Khuzin). The average age of the onset of sexual maturity
among the female White Sea population in the second half of the 1960s
was established roughly at 4.5 years (R.Sh. Khuzin). Evidently, how-
ever, the average age of females that have attained maturity varies within
certain limits, depending on the intensity of hunting, especially on the
replenishment of the mother population by those of the younger gener-
ation that have escaped the hunters. [t is significant that over the decade
from 1953 through 1962, the average age of the Newfoundland females
that had attained sexual maturity fell from 5.5 to four years (Sergeant,

315

419

1966). Males develop somewhat more slowly and their maturity sets in,
in most cases, at five years of age (Chapskii, 1963). According to some
other data (Yakovenko and Nazarenko, 1967), even four-year-olds are
mature. On the other hand, Newfoundland males become capable of
mating generally at seven to eight years of age (Fisher, 1954).

The uterus (without vagina) of the lactating females two months after
parturition weighed about 200 g. The weight of the paired ovaries (their
average) in such females varied markedly, from 6 to 10 g; the testes of
adult males in the breeding season without the spermatic cord weighed
135-300 g and 200-370 g with it; their length varied from 10- 14.5 cm.
The growth intensity of the White Sea harp seal (males) is shown in
Table 18.

It can be seen from Table 18 that the increment in the first five
years is quite marked (sharp variations are undoubtedly due to the small
number of animals measured). Later, however, it is very small and can
be perceived only in a series of animals. Similar age variations are seen
among females but their growth tapers off slightly sooner (roughly by a
year), after which their length increase is very small.

Molting in animals of all later generations, commencing from year-
lings, is an extremely perceptible periodic phenomenon in the life of the
harp seal; at the same time, it has played an extremely important role in
hunting since it serves as a no less powerful impulse than reproduction
for the concentration of large numbers of animals in certain periods in
relatively small sections of drifting ice floes. Hunters attempt to reach
such dense molting rookeries as well as the nurseries. Chronologically,
molting follows the periods of parturition, lactation, and mating, extend-
ing among the White Sea seals from the last 10 days of March up to
mid-May. This period does not cover the duration of individual shed-
ding of the hair coat (which in some animals is much shorter) but the
duration of the overall period of molt among animals in a population.
The animals setting out to molt initially concentrate in nearly the same
areas (or in their proximity) where the nurseries existed quite recently.
In the 1960s, the molting nurseries were quite often encountered in the
neck region of the central basin of the White Sea. Quite often, espe-
cially when access to the basin was delayed, the animals migrated to the
northeastern regions of the sea into Mezensk Bay and into the inlet (or
collected there). By mid-May the animals departed (or drifted away) from
there to the ice floe fringes in the Barents Sea. Sometimes the rookeries
were formed opposite the western sections of Tersk coast. In 1964, a
large rookery was detected even close to the Karelia coast, in the region
of Gridin (M.Ya. Yakovenko).

420

315 Table 18. Increase т body length (Lc) of the White Sea harp seal (males) in relation to
age (Yakovenko ef al., 1964*)

Age, years Number Range, cm Average, Annual
measured cm increment,
cm
1 8 130-150 138.1
2 4 152-169 161.0 22.9
3 5) 156-172 165.0 4.0
4 6 166-182 174.7 Od
5 16 170-191 180.9 6.2
6 57 162-196 184.7 3.8
7 70 170-207 186.5 1.8
8 42 162 - 201 188.6 2.1
9 45 161-205 189.3 0.7
10 14 181 - 203 192.9 3.6
т 11 185 - 198 193.4 0.5
12 9 178-200 189.6 3.2
13 3 196-202 199.0 9.4
14 2 200 - 203 201.5 25
15 2 185 - 190 187.6 —4

The seals select various types of ice floes for molting rookeries with
316 the only proviso that they should not drift rapidly toward the open fringes
and at the same time provide access to the water. The animals lie along
the edges of large- and medium-sized ice floes forming a live border with
their bodies lying close together. Such ice floes are separated from each
other by narrow or sometimes broad open water pools or partitions or
gapes; the animals lie on either side of these, presenting a picture of com-
plex branched strips or bands. Such a deposition is highly characteristic
of the early male rookeries. The animals are disposed more haphazardly
on the so-called conglomerate ice, i.e., predominantly on small broken
ice pressed together by winds or currents and held together by frost.

The density of disposition, the size of the rookeries, their number,
and composition vary widely. The animals are sometimes concentrated
in a single huge rookery or sometimes divided into several rookeries
of much smaller size. Under favorable conditions, a rookery can exist
continuously for quite a long time.

An extremely characteristic feature of the molting rookeries is the
fluid state of their age-sex composition. The first (excluding pups) to molt
are the mature males and the first molting rookeries consist almost exclu-
sively of animals with a wing pattern. At the commencement of April
immature animals of both sexes except under-yearlings (gray animals)
join them. After shedding the embryonic white coat, the under-yearlings

317

421

molt no more in the first year. However, at this time, young animals (aged
one to four-five years still preserving infantile coloration) constitute no
more than 15%. The proportion of mature females in the rookeries at
this time runs into a few percentages but in any case does not exceed
10%.

In the last 10 days of April, the immature animals increase in num-
bers up to equal those of animals with a wing pattern and the percentage
of the adult females slightly rises.

According to the situation prevailing in the 1920s and the 1930s
and later (Dorofeev and Freiman, 1928; Surkov, 1957), at the end of
April and in the first few days of May, the percentage ratios existing in
the molting rookeries between the main constituent animals somewhat
reflect the potential ratio characteristic of the herd: adult males and adult
females 25% each and immature animals 50%. These figures can hardly
be regarded as factual since females predominate among the animals
caught. There is reason to believe that the actual position is considerably
more complex.

The above age-sex ratios in the molting rookeries at April end to
early May represent a regular process observable even at present. It is
graphically depicted as the crossing of curves showing the different peri-
ods of molt among the different constituent animals of the herd. This
is strikingly demonstrated, especially by the data for the early 1960s,
when no one doubted the distinct disproportion between the number
of productive males and mother animals (Yakovenko and Nazarenko,
1962; Yakovenko, Nazarenko and Timoshenko, 1963; Yablokov, 1962;
and others). Consequently, from the equilibrium (or similarity) of the
percentage ratios between males and females in the late molting rook-
eries, no conclusion whatsoever can be drawn about the actual balance
between the different groups in a herd.

The final period of the molting rookeries of the White Sea herd
was studied less fully since the seals desert the White Sea and hunting
ceases. It is usually assumed that the very late rookeries consist mainly of
immature animals of both sexes and adult females. However, even in the
first of the molting animal concentrations along the fringes of the western
part of the White Sea inlet, young ones were in a majority from April 5
(Khuzin, 1970) and observations of the migrating animals point out no
perceptible division among them into age groups as the animals advance
toward the White Sea (Beloborodov, 1969). Some direct observations
(Sivertsen, 1941) also point to a large variation of the composition of
the rookeries even in the last 10 days of May.

The process of molting is quite prolonged. The old hair coat con-
tinues to be sported until the new hairs grow to half the length of those

422

shed. However, the implantation strength of the older hairs gradually
decreases. During molt, whiskers too are shed along with the hairs; evi-
dently claws commence growth at this time. The horny layer of the epi-
dermis is also shed in small and large strips simultaneously.

Some new data available only recently suggest that molting among
the various groups of seals entering the White Sea occurs at different
times in different regions. It has been found that among an overwhelming
majority of the animals residing in the open sea, especially the adult
males, molting was at its peak on April 27 while molting of animals
arriving later in the inlet region had only just commenced (Khuzin, 1970).

Enemies, diseases, parasites, mortality, and competitors. From among
the vertebrates, the potential enemies of the harp seal in water are the
Greenland shark (Somniosus microcephalus) and the killer whale (Orci-
nus orca) and on ice floes, the polar bear. Raven and arctic fox can
also be listed among them. The destructive role of all these animals is
extremely variable. The Greenland shark and the killer whale pose the
greatest danger to the White Sea herd. However, the shark is confined
mainly to the deep sections although it is capable of surfacing and, what
is more important, it is generally rare in the White Sea. Seal remnants
were detected in the stomach of Greenland sharks caught on Murman
and on Kanin Island (Breitfus, 1906; Smirnov, 1935; Andriyashev, 1954).
The role of the killer whale has not been supported by actual data since
it is not actually caught in the Barents Sea; yet it cannot be ruled out
as a potential enemy. In fact, in the eastern part of the sea, especially
along the coasts, this whale is encountered extremely rarely during the
migration of the White Sea harp seal.

The polar bear, by and large, poses no real danger to the seals of
our herd for several reasons. It has not been sighted, as far as people
can remember, in the zone of reproduction and is extremely rare in the
high latitudes of our western Arctic. Further, the bear has access only
to seals on ice floes where the harp seal is almost nowhere seen in the
summer.

Foxes can probably attack the newborn but they are hardly even
seen in the nursery zones; by the time the rookeries drift toward Kanin
Nos, the young one has grown considerably and molted and the fox is
no longer capable of attacking it.!!? The fox, if it strays onto an ice floe,
has to satisfy itself under the best of circumstances with only the frozen
placenta.

In this respect, ravens cause serious damage and are often encoun-
tered in the nurseries. They feed mainly on the placenta or the dead

112 See: V.G. Heptner et al. Mammals of the Soviet Union, vol. 2, pt. 1.

318

423

pups but, in rare cases, may even attack live pups, primarily undergrown,
sick, underfed, and hence incapacitated ones. Regardless, the skeletons
of pups with a pecked head have been encountered from time to time
in the White Sea rookeries. Sometimes, large gulls also indulge in this
activity. In some years the number of dead pups with pecked heads has
reached 25% of the total dead pups recorded in a given section of the
nursery (Rudakov, 1936). In other years, however, as in 1947, no such
instances were reported (K.K. Chapskii). An instance of attack even by
ermines has been recorded (L. Popov, 1955).

Among the other more common factors for pup mortality are: (1)
defective births (birth with the amniotic sac and placenta) and other
instances of still births; (2) freezing and death due to emaciation; (3)
crushing by hummocking ice floes; and (4) washing away by waves. Pups
in the White Sea perish relatively more often due to the second and
third factors though sometimes the first factor as well as the second play
a predominant role (Yakovenko and Nazarenko, 1962). The emaciation
of a pup transformed into a starveling is caused by the cessation of
lactation, due to the loss of its suckling mother. If the pup ceases to
receive milk at a weight of less than 15 kg, it cannot survive (Yakovenko
and Nazarenko, 1962). Pups whose lactation has been interrupted even
somewhat later do not attain normal growth.

The destructive effect of the hummocking of ice floes, which crushes
the pups, is very difficult to estimate since the dead ones buried under
the vast ice escape attention. Such causes of death can be judged only
from instances when the crushed pup is partly visible among the broken
ice (or the tracks of the animal are lost under it). Such finds are very
rare and do not reflect the true scale of mortality, which is perhaps
relatively high since hummocking in the White Sea is a fairly frequent
phenomenon.

The washing away by waves of pups which have yet to complete lac-
tation (or their falling into the water while negotiating on the ice floes),
causing excessive cooling of the body, is no mean factor in their mor-
tality. However, at present, when the nurseries are formed in the open
waters of the White Sea and not close to the fringes (as, for example, in
the 1920s), very few pups perish for this reason. In the other locations of
the White Sea rookeries, however, such deaths of pups represent one of
the most significant factors (N. Smirnov, 1927). Yet situations that prove
fatal to the pups do arise from time to time in the White Sea nurseries.
One such is when the young ice breaks up intensely during compres-
sion and later greatly expands, as happened in 1967 (Г. Popov, 1971*).
This factor is of equally great relevance to the infant mortality of the

319

424

5
Jan Mayen and eastern Newfoundland (in the “front” zone) populations
(Nansen, 1924, and others).

There is yet one more, albeit episodic, very intensely manifest natural
factor of infant mortality. In years of anomalous wind directions in the
White Sea, when the persistent northeastern and even northern winds
prevent the drifting of ice floes into the inlet zone in spring, the well-fed
young find themselves carried away by the young ice floes in the opposite
direction, i.e., into the western part of the White Sea (Fig. 173). As soon
as the ice floes break up and thaw, pups scatter in small groups, at times
all along the periphery of the sea, right from the uppermost part of
Kandalakshsk Bay to the Onega River and Arkhangel’sk. The weakened
and emaciated pups, not finding their natural food, become incapable of
independent survival under the unusual conditions and often even crawl
to the coasts, thus becoming easy prey to any land predator, even man.

ZONE OF RECOVERY

® ARKHANGELSK

Fig. 173. Some sites of tagged White Sea harp seal pups in 1966 scattered along
the periphery of the White Sea (data of the Polar Research Institute of Sea
Fisheries and Oceanography, PINRO).

319

425

Such a situation prevailed in particular in 1966 and probably led
to the large-scale death of pups (Tambovtsev, 1966; Bianki and Kar-
povich, 1968; Beloborodov and Potelov, 1968). Such instances undoubt-
edly existed even in earlier times although not very often.

The overall magnitude of the natural mortality of White Sea pups
during their residence in the nurseries is not amenable to precise compu-
tation since the newborn can be washed off, crushed during hummocking,
fall into water, and the dead buried under snow. The figures cited below
should therefore be regarded as extremely approximate. For the 1930s,
the figures cited at 10 to 11% are slightly exaggerated (Rudakov, 1936)
and for the 1940s, on the contrary, the figure of less than 5% is an
underestimate (K.K. Chapskii). The actual figure is obviously 5 to 7.5%.
The overall mortality of the pups in their first year for the White Sea
herd was taken at roughly 20% in the calculations of population dynam-
ics (М. Smirnov, 1928; Р.А. Rudakov, К.К. Chapskil). The mortality in
the subsequent age groups has been roughly assumed at 9% in the sec-
ond year, 6% in the third year, and 5% each in the fourth and fifth
years; the average is 7.5% for the mother population and the produc-
tive males (K.K. Chapskii). These same indices were used in the latest
calculations of the population dynamics of the White Sea harp seal for
1970 (Yakovenko, 1967). For seals of the western Newfoundland herds,
the Canadian investigators put the approximate extent of natural mor-
tality in the first year at 50% of the generation (Rasmussen, 1957). The
average index of natural mortality for the entire population reproducing
in the northeast of the Strait of Belle Isle has been assumed at roughly
15% (Fisher, 1952).

There are no clear references to the diseases they cause but seals
do represent a source of disease for the people around them (called
“chingi”). When skinning the killed animal to remove blubber, infec-
tion of any wound produces an extremely serious inflammatory purulent
process [seal finger] leading ultimately to abnormal bone deformation
(or damage to the other parts of the limbs).

The helminth fauna of the harp seal has not been studied equally
well in all parts of its range. Evidently the helminth fauna was better
studied for the harp seal inhabiting the White Sea and in the region
of Jan Mayen (Vagin, 1933*; Skryabin, 1948*; Mozgovoi, 1953; Delya-
mure, 1955; Delyamure, A. Skryabin and Alekseev, 1964*; Delyamure,
A. Skryabin and Treshchev, 1965*; Khest, 1932*; Zhuar, 1935*; Stankard
and Shoenborn, 1936*). Ten species of helminths and two larval forms
are known among harp seals. The trematodes Orthosplanchnus arcticus
and Pseudamphistomum truncatum infect the gall bladder and the bile

320

426

ducts of the liver. The cestodes Diphyllobothrium cordatum, D. schis-
tochilus, Diplogonoporos tetrapterus, Diphyllobothriidae g. sp., and others
are sometimes parasites in the intestine. The nematodes Contracaecum
osculatum, Phocascaris phocae, Ph. cystophorae, Anisakidae g. sp., and
others infect the stomach and intestine, with as many as a thousand
present at one time; Terranova decipiens is encountered only in the intes-
tine. The acanthocephalan Corynosoma strumosum infects the intestine
(V.V. Treshchev).

A comparison of the helminth fauna of the White Sea and Jan Mayen
harp seals showed that acute differences along with features of similarity
exist (Table 19). Of the 121 Jan Mayen animals studied (V.V. Treshchev),
113 (84.8%) were infected. Newborns (seven) were free from helminths;
however, the yearlings as also almost all animals of other age groups were
100% infected. The most infected body parts were the stomach (in 91.7%
of the animals) and the small intestine (87.6%), and less frequently the
duodenum (29.7%). It was established that five- to eight-year-old White
Sea harp seals are infected more severely than other age groups. An
average of 131 helminths were found in the infected animals.

According to the data of investigations covering 169 White Sea harp
seals (V.V. Treshchev), 95 (56.2%) were infected. Newborns with white
hair coat and normally molting pups were not dissected for helmintholog-
ical studies but two of the 72 moited pups (gray ones) aged 1.5 -2 months
were found to be infected with immature nematodes. All animals older
than one year (97), except for two, were infected (97.7%). The seals of
the White Sea herd were more severely infected with helminths at the
age of 13-16 years. Often the stomach (in 56.2% of the animals) and the
small intestine (52.0%), and less frequently the duodenum (44.4%) were
infected. An average of 341 helminths was found, indicating the severity
of the invasion, which led to the formation of innumerable unhealed
sores up to 20 mm in diameter on the stomach walls.'!°

There are no serious competitors of the harp seal in the White, Bar-
ents, and Kara seas with regard to food. In the pelagic regions in sum-
mer, stray ringed seals are sometimes found along with this species on
the polar fringes. In autumn, in the coastal regions, especially on Novaya
Zemlya where large masses of polar cod gather for spawning, ringed seal
and bearded seal appear along with the harp seal. However, they rep-
resent no competition since fish food is abundantly available there. A
similar situation prevails later on the northern coasts of the White Sea.
In the pelagic sections, however, in winter and early spring, other seals

ПЗ A short review of the helminths of harp seals was made by the staff of the helmintho-
logical laboratory of Crimea State University under the guidance of Prof. S.L. Delyamure.

427

320 Table 19. Comparison of the helminth fauna of the harp seals of Jan Mayen and White

321

Sea herds

Helminth From Jan Mayen From White Sea
region region

Orthosplanchnus arcticus ar
Pseudamphistomum truncatum
Diphyllobothrium cordatum
Diphyllobothrium schistochilus
Diplogonoporus tetrapterus
Diphyllobothriidae g. sp.

++++

Contracaecum osculatum
Phocascaris phocae

+++

Phocascaris cystophorae
Terranova decipiens
Anisakidae g. sp.
Corynosoma strumosum

—
—>

+ +

are not found near the nurseries and molting rookeries. Only the white
whale can be seen rather frequently in the large open water pools but
predominantly in the central basin; these whales are not, however, seen
right within the rookeries.

Population dynamics. Man’s hunting activity is almost the lone factor
responsible for the population dynamics of this seal. Until the beginning
of this century, hunting in the White Sea was relatively low, the herd
reduction was much less than the births, and the growth of the herd was
therefore good.

In the first 15 years of this century, hunting intensified noticeably. and
the growth of the herd slowed down as a consequence. From the 1920s,
the population began declining sharply and continued to fall almost up
to the early 1940s (see pp. 391, 430). During the Great Patriotic War
(especially in 1942, 1943, and 1944), further decline in the population
was arrested and the herd slightly improved but the 3- or 4-year break
was inadequate. The subsequent fresh hunting spree, although not so
intense as before 1940, caused further population reduction. From 1965,
when the White Sea herds quite clearly became very lean, the use of ships
for hunting in the White Sea was banned for five years. An exception was
permitted only in favor of the coastal collectives who were assigned the
right to hunt for 20,000 pups a year. Additionally, nearly one-half this
number was killed outside the White Sea boundaries by the Norwegian
hunting fleets.

322

428

The sharp reduction in hunting activity stopped the degradation of
the herd but the measures did not produce the expected results; the
population rose at a slow tempo. By the end of five years’ restriction,
however, the White Sea population had improved considerably and began
to rise; nonetheless, by 1968 the mother seals only slightly exceeded the
1963 strength (M.Ya. Yakovenko). However, the accumulated reserves
should promote the growth of the herd. The seal population is regularly
controlled by the scientific institutions of the USSR and especially the
Soviet-Norwegian Commission for Seal Research in the Northeastern
Subarctic and Arctic Atlantic.

The status of the other populations of harp seals, the Jan Mayen
and Newfoundland seals, is not so fortunate. The population dynamics
are adverse in the case of the Jan Mayen herd, geographically closest
to the territorial waters of the USSR and even representing a hunting
base for our fleets (see pp. 389-391). The steadily declining numbers
of animals caught per ship reflect the significant reduction of reserves.
A century before (1866-1870), the average catch per ship was 4,341; at
the beginning of this century (1901-1905) 1,400; in 1921-1925, 1,046;
and in 1931-1935, fell to 862. Hunting practically ceased in 1941 - 1945.
In the postwar years, in spite of the ban and improved fleet, the catch
tended to decline.!4

Judging from the results of aerial photographic surveys of the harp
seal nurseries in the Newfoundland region, even the Canadian population
underwent sharp reduction, with the mother seals in it decreasing by
nearly one-half (see p. 391).

Field characteristics. The adult animals in the final phase of the hair
coat coloration are easily recognized from the intensely dark wing-shaped
patterns on the body standing out in contrast to the light-colored back-
ground and also from the head which is as dark as the pattern. Among
the animals of transitional type coloration, these dark patches are quite
distinct along the body flanks. They are diffuse toward the outer fringe
although the wing-shaped patches are of smaller dimensions. The imma-
ture animals of both sexes and the very young (infantile type of col-
oration) gestating or whelping females have fairly distinct contoured,
small, dark patches that are sparsely scattered on a gray background.

These seals usually live in herds on the coasts and do not emerge
onto stationary coastal ice (fast ice). (K.Ch.)

14 The numerical data were borrowed from R.Sh. Khuzin who processed the hunt-
ing statistics published by Sivertsen (1941), Iversen (1927*), and the Norwegian Fishing
Directorate.

323

429

Economic Importance

The harp seal is of primary importance to hunting. Its potential resources
оп restoration and subsequent rational utilization would ensure a catch
in future (for the population as a whole) of at least 400,000 animals per
year for a very long time.

Unfortunately, due to mindless plundering of reserves even in the
relatively recent past under pressure of international competition and
markets, the reserves of this species of seals had been rather depleted by
the middle of the present century. This situation was largely due to the
long absence of a reliable census. The numerical data given in Table 20
show the volume of hunting in the past and at present.

The advance of the harp seal to first place among the seals hunted
was promoted not so much by the very high potential of its population
increase (the total actual reserves of the ringed seal are slightly more than
the present-day strength of the harp seal), as by the favorable conditions
for its hunting. These conditions are: the high degree of concentration
in localized sections of the range where these seals form massive and
fairly dense groups in rookeries on the ice and are quite accessible even
to poorly equipped hunters. With the present organization and hunting
techniques, however (powerful icebreakers, better hunting ships, trans-
port and survey helicopters and planes, radio communications, etc.), the
high concentration of these seals makes for highly efficient hunting.

The harp seal is caught at present mainly for the fur of the juveniles.
An additional raw material is the subcutaneous fat used as medicinal,
edible, and commercial fat. The meat portion is used as feed in the
farms of fur animals.

The harp seal has long been hunted in the White Sea. The origin
of such hunting runs into the prehistoric period. There was regular seal
hunting in the Neolithic period, mainly (judging from kitchen remnants)
of the harp seal, even in the basin of the contemporary Baltic Sea. In
the remote past the seals played a very significant role in the life of
prehistoric man at many places by providing him not only with hides
and fat, but also edible meat. Until recently, the coastal Eskimos and
Chukchis depended almost wholly for their existence on the hunting of
the walrus and various seals.

Even at present for people from the south who settled along the
White Sea coasts and took to raising cattle and at places cultivation and
fishing, sealing (in which the hunting of the harp seal was of exclusive
importance) is not the least of their activities. Right up to the beginning
of this century (especially in 1901), earnings from hunting the White
Sea harp seal in Mezensk district were put at 27,000 rubles and exceeded

322

430

Table 20. Average number of White Sea harp seal* killed per annum at five-year
intervals from 1875 through 1945 (in thousands)

Years Killed by Russian Killed by Norwegian Total
hunters hunters average

ее ВРК pos ee ted res kel tee for the

Min. Max. Average Min. Max. Average period

1875 - 1880 19.3 48.3 30.7 5.6 9.1 73 38.1
1881 - 1885 8.9 58.3 26.6 gril 19.8 9.6 36.2
1886 - 1890 12.9 33.0 19.1 15.8 22.7 19.6 34.8
1891 - 1895 13.3 40.7 23.6 24.1 33.0 28.5 S251
1896 - 1900 37.0 74.5 57.1 27.1 38.0 34.7 91.7
1901 - 1905 19.3 64.6 33.9 32.5 79.8 57.9. 92.0
1906 - 1910 19.2 25.6 21.8 42.6 107.0 76.8 98.7
19111915 27.7 50.2 46.3 84.5 118.4 99.2 125.6
1916 - 1920 28.0 49.6 37.8 74.9 154.2 106.6 144.4
1921 - 1925 36.4 124.6 70.4 69.0 343.0 188.8 259.3
1926 -1930 92.5 182.1 140.5 90.7 РЗ 167.2 316.7
1931-1935 102.1 197.8 141.1 47.1 150.5 108.1 245.1
1936 - 1940 10.1 168.9 95.3 34.1 42.7 37.1 124.9
1941-1945 Poot 131.8 55.0 — — = 55.0

*Calculation based on material compiled by R.Sh. Khuzin, who used such primary sources
as Iort and Knipovich (1907) and Siverstsen (1941*), whose data included other species as
well (their quantum does not exceed 1%); data for the postwar Norwegian statistics (Fish
Catch, 1949 to 1959) and Russian statistics.

earnings from salmon fishing (23,000 rubles). A higher income came only
from forestry (wood processing and sawing) at 100,000 rubles, agriculture
77,000 rubles, cattle breeding 47,000 rubles, and navaga fishing 34,000
rubles. The income was less from deer farming, river and lake fisheries,
hunting in forests, and transport.

The primitive seal hunting on the ice floes in the White Sea was cum-
bersome and risky in the past and was organized by groups of hunters.
The people on the coasts and in Mezensk district called such groups
“bursas”. For the most part, a group consisted of five to seven hunters
in a spacious boat. On the Tersk and Karela coasts, these groups (called
“romshas”) comprised only a few people. When the rookeries began
forming, the “bursas” set out with their paraphernalia for the sea ice.
Dragging heavy boats, experiencing quite some hardships, and exposed
to danger, the hunters wandered in search of the rookeries for many
days. It was most risky to land on drifting ice floes singly or in twos
or even threes (without boats) while chasing the seals sighted from the
coasts. It was necessary to hurry back, towing the hides and the blubber,
to negotiate the island before the ice broke apart excessively.

The first Russian ships began being detailed for seal hunting
in the White Sea right at the beginning of this century but group

324

431

hunting continued until the Revolution and after. From the 1920s to
the 1930s, most of the hunters from the coastal collective farms engaged
icebreakers which, in the 1920s - 1930s as well as in the postwar years,
carried out state hunting operations in the White Sea. Apart from the
icebreakers, special hunting ships were recruited. On the whole, hunting,
regularly assisted by reconnaisance planes, was more intense.

The White Sea harp seal was killed in the largest numbers (exceeding
300,000 annually) for five years from 1924 through 1928. In the next five-
year period (from 1929 through 1933), the average annual kill dropped to
250,000, and still later from 1934 through 1938, fell further to 188,800 on
average. There is no doubt that hunting in the 1920s was extremely intense.
The average annual kill in the postwar years steadily fell as follows:

О. 166,300
Spy О оное 151,700
OSD Об: а, 115,000
О A), Su res aa) ot 81,900

Norwegian hunting ships also cruised along with our hunting opera-
tions in the White Sea region and in the adjoining southeastern sections
of the Barents Sea. Before the end of the 1930s, the Norwegian hunters
were given a concessional right to hunt for the harp seal in the north-
ern part of the White Sea (north of the line joining Capes Orlov and
Konushin). At the end of the Great Patriotic War, the agreement was not
renewed and at present the Norwegian hunters hunt outside the White
Sea limits and in the adjoining areas in a few dozen small hunting boats.
Hunting is regulated by agreements.

From the second half of the 1960s, the organization of our hunting
activity in the White Sea underwent radical change. In the five years
ending with 1969, state hunting ceased because of the depleted reserves.
Neither icebreakers nor special hunting ships carried out hunting in the
White Sea. An exception was made only for the local collective farms,
which were given the right to hunt 20,000 pups a year. The collectives
rented helicopters to reach the hunters on the ice and transport the
killed animals to the coastal base. No more than 10 days were spent in
the entire hunting operations under favorable conditions.

Until recently, well-fed normally molting young animals were
brought onto the coasts where they were allowed to complete molt in
special enclosures on land.

The Norwegian hunters in the immediate proximity of the White
Sea inlet area caught no more than 14,500 animals per annum in the
1960s (Table 21), mainly the older juveniles and adults. It was thought
prudent to maintain the hunting level as at the end of the 1960s in

325

432

the next decade to promote a more rapid population recovery. After
a short period of relative restriction, the population had not reverted
to the level of the 1920s and 1930s (the year 1925 was a record for
the White Sea region; Soviet hunters caught 124,600 animals and the
Norwegians 343,000 animals, 1.е., slightly less than half a million seals).
The ban imposed in 1963 on killing adult females in the nurseries had
an extremely salutary effect in restoring the White Sea population.

The technique of hunting in the nurseries was extremely simple in
the past. The hapless white pups could not escape from the hunter and
were killed directly using clubs. Since suckling mothers usually do not
leave their pups, “hunting” them also posed no problem. Nevertheless,
rifles were used to kill them. In the past, hunting in the molting rookeries
was more complicated. Hunters with harpoons [rifles] approached quietly
in white masks and aimed from behind cover. The shots frightened the
sleeping animals and soon only dead animals remained on the ice floe.
In some cases, when the ice floes became compact (usually in high tide),
open water pools disappeared and the ice lumps filling them froze and
the animals were thus cut off from the water. Hunters could then kill
them even with clubs. Usually, however, even in compact frozen areas,
the animals could ultimately get into the water by pushing aside the ice
lump or pressing it with their bodies.

Table 21. Harp seals (White Sea population) killed in the postwar years (in thousands)
(after R.Sh. Khuzin)

Year Killed by Killed by Total

Soviet hunters Norwegian hunters killed
1946 79.1 8.5 87.6
1947 161.1 6.6 167.7
1948 146.3 8.9 155.2
1949 183.4 25.1 208.5
1950 194.7 LT, 212.4
OSH 192:2 33.8 226.0
1952 131.4 19.1 150.5
1953 88.3 12.4 100.7
1954 152.8 11.6 164.4
1955 97.8 191 116.9
1956 68.0 25:1 93.1
1957 107.9 22,3 130.2
1958 119.6 15.1 134.7
1959 101.4 8.5 109.9
1960 95.9 10.7 106.6
1961 93.8 1:2 105.0
1962 106.9 8.3 15.2.
1963 69.5 13.3 82.8
1964 62.7 14.6 77.3
1965 20.1 6.4 26.5

1966 20.0 12.2 32.2

326

433

The first of the denser rookeries of males with a “winged” pattern
and disposed in rows along the edges of open water pools and fissures
(Fig. 174) are the least accessible to hunters. It is difficult to approach
the animals within the required distance as they abandon the ice floe at
the first shot and dive pell-mell into the water.

The killed animals are processed very quickly: first a cut is made
along the abdomen from head to tail, next circular incisions made through
the skin with the fat layer around the base of the flippers, and then
the subcutaneous fat separated from the muscles. The skins with the fat
upward are spread out on the ice and stacked for loading in a vessel
that comes later. Nowadays, the extensive used of airplanes and killing
pups exclusively has made the job quite simple and fast. At the coastal
base the fat is separated from the skin, the skins degreased to the max-
imum possible extent, and salted. In the natural or even dyed form,
the white embryonic pelage and now, more so, the skins of the molted
gray pups are in great demand in the internal and well as international
markets.

Drifting ice floes in the region of Jan Mayen Island represent another
region of hunting. Exploitation of this international hunting zone began
over 200 years ago. The maximum intensity of hunting in Jan Mayen
was recorded in the 1870s (200,000 seals were killed there in 1874). The
decline in killing began subsequently, touching the lowest level at the
end of the first 10 years of this century. World War I interrupted the

Fig. 174. Early spring rookery of adult males. White Sea, April 18, 1966 (photo-
graph by Уи.Г. Nazarenko).

326

327

326

434

hunting activity and this helped the herd to restore itself to some extent.
After the war, hunting resumed and reached a fairly high level.

World War II again interrupted hunting activity but it was rapidly
resumed thereafter and, in the first half of the 1950s, exceeded the prewar
level. On average, the kill in the first five years of the 1950s was 39,300
seals with a very large number of modernized vessels operating. However,
the success was short-lived and unreliable. The pressure on the animal
population was excessive and it rapidly declined, an index of this decline
was the reduction in the number of animals killed per ship (see Table 22).
The lowest annual yield for the entire hunting duration in this century
occurred in the first half of the 1960s (average kill per ship fell to 467
seals in one expedition although the vessels had been equipped with new
diesel motors) (R.Sh. Khuzin).

Soviet ships began hunting in the ice floes of Jan Mayen Island
region in 1955 and continued hunting there until 1965 (see Table 23).
Hunting was discontinued as it was unprofitable.

The commercial exploitation of the Newfoundland populations by
the local people, in which the USSR became somewhat interested in
the 1960s, began almost concurrently with hunting of the White Sea
resources. Hunting of harp seals using ships had begun there at the end
of the eighteenth century and reached an immense scale of 500,000 a year
in the 1820s. Hunting continued at this level for forty years (Allen, 1880).

Table 22. Norwegian hunting of the harp seal in the Jan Mayen Island region from the
18805 (after R.Sh. Khuzin)

Years Average annual Average number of Average kill per
kill, thousands expeditions per ship expedition,
ship per annum numbers
1881 - 1885 82.8 17.5 5,241
1886 - 1890 33.3 22.8 1,469
1891 - 1895 45.8 19.6 2,338
1896 - 1900 24.3 12.4 1,960
1901 - 1905 15.1 10.8 1,400
1906 - 1910 14.0 21.6 648
1911-1915 17.4 29:2 596
1916-1920 32.3 59.7 542
1921 - 1925 19.5 18.6 1,046
1926 - 1930 32.6 33.5 969
1931 - 1935 32.3 37.4 862
1936 - 1940 36.7 45.6 803
1941 -1945 — = —
1946 - 1950 36.1 37.0 976
1951-1955 39.2 55.6 859
1956-1960 25.0 39.8 585

1961 - 1965 18.6 39.8 467

435

Table 23. Jan Mayen harp seals killed by Soviet hunting ships in the 1960s (after

R.Sh. Khuzin)
Year Number of Total killed by all ships,
hunting ships number

Total Pups
1960 8 3,356 420
1961 8 6,043 755
1962 8 2,423 303
1963 i, LOG, 222
1964 8 4,483 560
1965 8

6,273 896

One can well conjecture the original population level of the local
harp seals to be able to sustain such a prolonged and intense rate of
hunting. By the end of the nineteenth century, the number of animals
killed had fallen to half in spite of maintaining the same intensity of
hunting; in the first few decades of this century, the number decreased
even further, to 160,000 a year.

World War II served as a fortuitous interlude that extended for
sometime after the war. The continuous interruption of hunting using
ships undoubtedly promoted a partial recovery of the animal population.
But hunting resumed in the 1950s and exceeded the prewar level, with
the annual kill reaching 262,000 (Fisher and Sergeant, 1960*). At the
present level of the animal population, given such large kills the animal
population can hardly restore itself to the original level.

At the beginning of the 1960s, hunting was carried out in the New-
foundland region twice on a trial basis by Soviet hunters. As it proved
unprofitable and Canada extended its coastal waters to 12 miles, the
Soviet Union ceased hunting in the Newfoundland region.

At the end of the last century and in the first thirty years of this
century, this seal was also caught in nets. This was mainly practiced by the
locals off Murman but here and there the White Sea harp seal was caught
even in the White Sea before total icing set it. Catching in nets was quite
profitable off Murman. For example, at Kharlovka about 1,000 seals were
thus caught from the end of winter to early spring in 1898 - 1899.1 This
hunting practice flourished even in the 1920s (Skvortsov, 1927). Nets
were no longer used in subsequent years off Murman mainly because the
seals changed their approach to the Murman coast.

Sighting the harp seal on the Murman coasts, people unjustifiably
assumed its adverse effect on the catch of cod there. Even very old reports

15 See: Proceedings of the Murman Scientific-Hunting Expedition, St. Petersburg, 1912.

328

436

contain frequent references to the fatal consequences of the presence of
seals on the Murman fishing industry. Thus, in 1801, the seals report-
edly “chased away the entire fish from the Murman coast” right up to
Rybach’ii Peninsula. In 1803, the “catch was good from the spring but
kozha!!© arrived insummer and chased away the entire cod”. In 1807,
“tight from the spring, kozha spread all along the coast like tree stumps
in a forest, as a result of which the fishermen could not catch even a single
fish”. Similar complaints were registered in subsequent years (Reineke,
1830*). In fact, the White Sea harp seal continued to be accused for
quite some years, often with even greater exaggeration.

The accusations that the harp seal consumed large quantities of com-
mercial fish, especially cod, or chased it away were never confirmed.
On the contrary, the arrival of cod was usually not accompanied by the
arrival of seals; instead the absence of cod coincided with the arrival of
the White Sea harp seal. Thus the cod per se holds no attraction for the
seal (N. Smirnov, 1924); this is explained by hydrological conditions. The
harp seal is a distinct element of the arctic fauna; it is sensitive to the
thermal regime of the water (even early in this century it served as an
indicator of the cooling of the water or of the invasions of arctic waters
(Linko, 1912)). Thus, instances of the temporary disappearance of the
relatively thermophilic cod on the Murman coast and the simultaneous
appearance there of the White Sea harp seal were actually impelled by
cooling or ingress of arctic waters.

The harp seal of the White Sea herd is one of the most harmless
species of seals in relation to the fishing industry, coming in third place
after the bearded seal and the walrus among the Pinnipedia of the North
Atlantic. (K.Ch.)

Subgenus of Ribbon Seals
Subgenus Histriophoca

RIBBON (BANDED) SEAL!”
Phoca (Histriophoca) fasciata Zimmermann, 1783

1783. Phoca fasciata. Zimmermann. Geographische gesch. des Men-
schen u. der algemein. verbeit vierfussigen Thiere, 3, p. 277. Kuril
Islands.

116 «“Kozha” is a local term used by the coastal people for the harp seal.
17 Krylatka—adult animal; belek—newborn pup; and serka—under-yearling after casting
the juvenile hair coat.

437

1831. Phoca equestris. Pallas. Zoogr. Rosso-Asiat., I, р. Ш. Kuril Islands,
Sea of Okhotsk (“Rarissime in mari Ochotensi, frequentius circa
Curilorum insulas. . .”). (V.H.)

Diagnosis

Four light-colored transverse bands run along the generally dark back-
ground of the trunk (Fig. 175). The labial whiskers number 41-42 and
orbital whiskers 6-8. The whiskers at the tip of the snout have a wavy
surface (Fig. 176). The first digit on the fore flippers is longer than the
rest. The paired air sac joined with the trachea is well developed among
males (Fig. 177). (V.A.)

Description

These are seals of moderate dimensions [up to nearly 2 m in length]; the
males are usually larger than the females.

Body long, streamlined. The color of the adults is typical: four light-
colored bands run along the dark background (black or brown among
males; brown or brownish-gray among females). One band encircles the
neck like a collar, covering also the rear portion of the head, and another

==,

92

Ра >

Sa к. <
2 — = о ems A =
ее —= a) О i

329 Fig. 175. Color of the adult ribbon seal (banded seal), Phoca (Histriophoca)
fasciata (figure by N.N. Kondakov).

329 Fig. 176. Whiskers of the ribbon seal (banded seal), Phoca (Histriophoca) fasciata
(figure by N.N. Kondakov).

438

329 Fig. 177. Air sac of the ribbon seal, Phoca (Histriophoca) fasciata (figure by
N.N. Kondakov). A—general view of air sac filled with air (seen from the right);
B—sketch of respiratory tract in the ribbon seal. 1—cervical section of air sac;
2—trunk section; 3—trachea; 4—enlarged section of lower trachea; 5—lungs;

6—thoracic cage (after M.M. Sleptsov).

encircles the body at the base of the hind flippers; two symmetrical bands
mark the outline of the base of the fore flippers. From a distance, these

330 flippers look like the attached wings of a bird. The width of each band
is 5.5-15 cm and its color varies from pure white to yellowish. Fine
mottling is sometimes seen on the light-colored bands of young animals.
There is no seasonal variation in coloration. (Age-related color variations
are described under “Growth, Development, and Molt”.)

The skull is shortened and broad, the cranial portion and zygoma
broad, the facial portion short and narrow (Fig. 178). The nasals are
short, on average 36.6 mm. The bony nasal septum does not reach the
posterior margin of the bony palate. The length of the palatine bones
is usually less than their maximum width. The tympanic bullae are large
(about 33.5% of the condylobasal length), their width considerably less
than their length, and set sharply inclined toward the longitudinal axis
of the skull. The alveolar margin of the upper jaw is curved; the lower
jaw is curved with a downward convexity corresponding to the curvature
of the upper tooth row (Fig. 179).

The teeth are small and number 32-36. The canines are small and
obtuse, the incisors weak, slightly curved, almost vertical. The molars
and premolars are small, set wide apart, their roots tending to fuse. The

330

331

439

Ри

iS)
<0 cree \ = Si a My
у md у %
Ip AD \ :

\

LX i WR
Sy чи y PS Е
y a 7 wo Г) 5 . я in,
\F2 ( ib inane SY,
: oct iat Ve Reo

Fig. 178. Skull of the ribbon seal (banded seal) Phoca (Histriophoca) fasciata
(figure by N.N. Kondakov).

only accessory cusp, located posterior to the base of the last tooth, is
most often in-the form of a faintly recognizable denticle.

The average body length varies from 155 to 165 cm depending on
the age composition of the sample). The maximum length of the male
(Bering Sea) is 192 cm, of the female 198 cm (males, however, are gener-
ally larger than females). The length of the os penis may reach 150 mm.

The condylobasal length of the skull in males averages 191.7 mm,
in females 190.7 mm; mastoid width in males 125.1 mm, in females
121.4 mm; length of the upper tooth row in males 57.5 mm, in females
59.0 mm (Ogney, 1935; $. Naumov and М. Smirnov, 1936; Chapskii, 1955,
1963; Shustov and Yablokov, 1967).

The maximum weight of these seals is 150 kg, the average being
55 kg. The average weight of the blubber (fat with hide) is 20 kg and
the subcutaneous fat layer is 2- 4.5 cm thick. The weight of the os penis
is about 20 g. The weight of the internal organs of mature animals is as
follows (g): heart 499-513, lungs 978 - 1,030, liver 1,378 - 1,403, stomach
401-441, intestines 1,351 - 1,433, and kidneys 123-144 (A.S. Sokolov et
al., 1969). The mode of life of the ribbon seal outside the icy season is
not known and hence there is no information on the seasonal changes
in weight. (V.A.)

330

332

440

Fig. 179. Bony palate and teeth of the ribbon seal, Phoca (Histriophoca) fasciata
(figure by N.N. Kondakov).

Taxonomy

The ribbon seal is distinguished in the family by its specialized pelagic
mode of life, which has led to some characteristic structural features.
Some authors (Ognev, 1935; Simpson, 1945; Chapskii, 1963) regard
the ribbon seal as a monotypical genus, while others (S. Naumov
and N. Smirnov, 1936) combine it in the same genus as the harp seal,
with which the ribbon seal shares several common morphological and
ecological features and evidently has a common ancestor, although the
genesis of the latter is not clearly known. (V.A.)

Geographic Distribution

Seas of the northern Pacific Ocean—Sea of Japan, Sea of Okhotsk,
Bering Sea and Chukchi Sea. Endemic to this section of the world oceans.

Geographic Range in the USSR

Divided into two isolated portions, 1.е., Sea of Okhotsk and Bering Sea
(Fig. 180). In the Sea of Okhotsk this seal is encountered on the spring
ice floes from La Perouse Strait to the Shantarsk archipelago and She-
likhov Bay. the southern boundary runs along the northern part of Tatar
Strait (Dorofeev, 1936) and into the coastal waters of the northeastern

331

441

250 500 750 1000 km
—

Fig. 180. Distribution of the ribbon seal (banded seal), Phoca (Histriophoca)
fasciata in the USSR (V.A. Arsen’ev).

coast of Hokkaido Island (Nishiwaki and Nagasaki, 1960; Nishiwaki,
1966). Distribution in the Bering Sea covers the coastal waters of Koryak
Land from Olyutor Gulf in the north, Gulf of Anadyr, Bering Strait, and
the Chukchi Sea to Kolyuchin Bay in the west (Shustov, 1965).

Geographic Range outside the USSR

Eastern part of the Bering Sea from Bristol Bay to Bering Strait and
waters of the Chukchi Sea to Cape Barrow in the east (Scheffer, 1958;
Shustov, 1965).

332

442

The southern boundary in the Bering Sea runs from Olyutor Gulf
northeast roughly up to 60°N lat., from there eastward and then south-
east up to Bristol Bay (Fig. 181). The southern limit does not cross the
islands of the Aleutian range (Allen, 1880). The northern boundary runs
at places between the Chukchi Peninsula, Alaska, and Wrangel Island
from Kolyuchin Bay to Cape Barrow. (V.A.)

Geographic Variation

The southwestern boundary of distribution of the ribbon seal in the
Bering Sea lies more than a thousand miles away from the northeastern
boundary in the Sea of Okhotsk. The existence of two local populations
is thus highly probable. All the same, geographic variation has not been
established. (V.A.)

Biology

The mode of life of the ribbon seal is known for a comparatively small
duration of the year, 1.е., at the time of spring-summer rookeries on the

Fig. 181. Species distribution of the ribbon seal (banded seal), Phoca
(Histriophoca) fasciata. Dots represent sites of records of the ribbon seal
(V.A. Arsen’ev).

333

443

ice floes. From this viewpoint, И is one of the poorly studied species of
Pinnipedia of the Northern hemisphere.

Population. An idea of the population of the ribbon seal in the Sea of
Okhotsk was provided by the data of the annual catch, aerial surveys, and
observations from ships used for hunting. The total population was thus
put at a few tens of thousands. No special census was undertaken. The
abundant population of the Bering and Chukchi seas began attracting
hunters only in 1961. Calculations based on the areas of ice floes, density
of disposition of seals on various types of ice floes, and aerial and visual
observations put the total strength of this population at 80,000 to 90,000
(Shustov, 1969).

Habitat. These seals generally select firm, clean, white hummocky ice
blocks invariably with an even surface for their spring-summer rookeries.
Instances of finding these seals on dirty ice blocks are very rare and are
possible at the end of the icy period when the area of the ice floes has
greatly diminished. The height of the ice floes is not of much consequence
as the seal can jump onto an ice block 1 m or more in height. The main
rookeries are disposed on drifting ice floes far away from the coasts but
usually overlying depths not exceeding 200 т. The seals are also sighted
in coastal waters but only in the case of early thawing of ice floes. Outside
the icy period, the seals evidently lead a pelagic mode of life since they
never emerge onto the coasts (barring extremely rare cases). The present
sites of distribution of the seals have not been established.

Food. Data on the food of the ribbon seal have been collected exclu-
sively for the period of their residence in the rookeries on the ice floes
with no information at all available outside this period. Most of the
dissected stomachs in all the regions of study were found to be empty.
Thus, 91.6% of 443 stomachs dissected in the Sea of Okhotsk were empty
(Pikharev, 1941); in another case, 82% of 398 stomachs were devoid of
food (Arsen’ev, 1941); in the Bering Sea, 97.4% of 1,175 stomachs dis-
sected were empty (Shustov, 1965). Based on these data, it was earlier
thought that the ribbon seal did not feed in the icy period but thorough
investigations established that the intestine of many seals contained food
remnants while the rectum was packed with fecal matter. Very rapid food
digestion is evidently characteristic of this seal.

In addition to the items listed in Table 24, the stomach of seals in
the Sea of Okhotsk at one time contained the bone remains of navaga
(Freiman, 1936b) and mysids (Nikulin, 1937b). In the Sea of Okhotsk
as well as in the Bering Sea, some stomachs revealed the feathers of sea
birds, sand, and stones.

The geographic variation of the food of ribbon seals was quite signif-
icant but data for the Sea of Okhotsk (V.A. Arsen’ev, 1941) are clearly

334

44+

inadequate. The age-related food variations are: young animals (primar-
ily the under-yearlings) feed mainly on pelagic invertebrates (amphipods,
mysids and isopods) while adults feed on benthic and pelagic forms in
spite of the fact that the depths in the regions of their habitat reach 50
to 100 m or more.

Home range. The ribbon seal does not form large rookeries on ice
floes but lives singly or sometimes in twos or threes. There are no distinct
male or female rookeries. These seals are seen only on ice floes with a
density of predominantly four to six points.

Hideouts and shelters. These seals do not make air holes or snow
shelters.

Table 24. Food items of ribbon seals (Shustov, 1965a)
Food item Number of cases Percentage of
number of stomachs
with food remains
Seaof Bering Seaof Bering
Okhotsk Sea Okhotsk Sea
CRUSTACEANS
Crangon dalli 1 — 2.0 —
Nectocrangon lar = 3 = 9.4
Pandolopsis sp. = 6 = 18.7
Pandalus borealis — 8 = 25.0
Pandalus goniurus 5 1 9.8 3:1
Eualus ватагай т 6 = 18.7
Spirintocaris murdochi os 1 = 3.1
Lebbeus sp. = 3 — 9.4
Temisto sp. (2 species) = 6 = 18.7
Stilomysis grandis = 2 = 6.2
Amphipods =a 4 — 12.5
Cephalopod mollusks 24 5 47.0 15.6
FISHES
Polar cod, Boreogadus saida = 4 — —
Atlantic herring, Clupea harengus = 2 — 6.2
Pacific navaga, Eleginus gracilis — 2 — 6.2
European smelt, Osmerus eperlanus — 1 — 3.1
Stout eelblenny, Lumpenus medius = 4 = 12.5
Alaska pollock, Theragra chalcogramma 47 = 92.1 —
Capelin, Mallotus villosus 1 2 2.0 6.2
Common sand eel, Ammodytes hexapterus = 1 = 3:1
Smooth lumpsucker, Aptocyclus ventricosus 1 = 2.0 —

Pacific cod, Gadus morrhua macrocephalus 4 — 7.8 =

334

335

445

Daily activity and behavior. The number of stomachs with food rem-
nants was 20% more among seals caught during the night, suggesting a
more active feeding at night (Shustov, 1965). No other data are available
on daily activity.

In cloudy weather, many seals are seen resting on ice floes in the
morning and evening hours while most of them remain in water at noon.
In rainy weather most seals prefer to be in water and rarely venture onto
the ice. On quiet, sunny days the seals almost do not enter the water in the
day but gather there in the evening hours and at night. A large number of
seals remain in water at night, roughly between 8:00 p.m. and 4:00 a.m.,
regardless of weather conditions (Pikharev, 1941; Shustov, 1965b).

The ribbon seal very easily negotiates ice floes with long leaps,
without touching the edge of the floe. It lies quite close to the edge,
quite often on a level surface at the foot of a hummock with its head
invariably toward the water. It is extremely sensitive and approaching it
without detection is very difficult. If, however, a boat with the motor
switched off runs straight toward the animal and the people in the boat
remain quiet, the seal carefully surveys the boat and permits its approach
within rifle shot without attempting to escape into the water. Probably
its auditory faculty is poorly developed compared to vision. An injured
seal attempting to reach the water quickly, will invariably stop and howl
sharply, which the hunter promptly takes advantage of. The animal moves
rapidly on the ice floe, contorting its body like a snake, dives almost
noiselessly, without a splash, moves far away in the water, and never
surfaces immediately. A badly wounded animal attempts to return to a
block of ice while a slightly injured one can fiercely attack the hunter
(Pikharev, 1939, 1941). Sometimes, for no reason at all, the seal begins to
move rapidly on the ice floe from one side to the other, rolling and turn-
ing from one side to the other, then rushes suddenly into the sea, only
to return immediately to the ice floe with a leap. The motive for these
actions is not clearly understood; they resemble playful acts although
done singly, without a companion.

Seasonal migrations and transgressions. Information on migrations is
practically not available. In the rookeries on the ice floes, the seals are
mainly passive and the movements they do make can hardly be regarded
as migratory. On the spring ice floes in the Sea of Okhotsk, the largest
concentrations are noticed south of Tauisk Bay, in the region of lony
Island, north and east of Sakhalin. At the end of the icy season, herds
of these seals migrate into the region of Shantarsk Islands and into the
Gulf of Sakhalin. They are distributed on the ice floes, in patches. In
relatively nearby regions on the ice floes, appearing almost identical
externally, some will be occupied by the animals and others vacant, the

446

occupied and vacant floes alternating. Possibly, this is associated with
the depths of the region and hence the chances of finding food (Shustov,
1965b). With the thawing of ice floes in the Sea of Okhotsk and the
Bering Sea, the ribbon seal disappears. Its summer residence has not
been established. Only some stray observations have been reported on
the migrations of some seals through the Bering Strait in spring to the
north and in autumn to the south (Shustov, 1965b; Tikhomirov, 1966).

Some distant transgressions of the ribbon seal are known. Early
September, 1927, a seal was killed in Vladimir Bay in the Sea of Japan
(about 44°N lat.) (Pikharev, 1941). In another case, an adult ribbon seal
was noticed on June 17, 1944, in Tsushima Strait (35°10’N lat. and
130°34’ E long.) 45 miles from the coast (Vedenskii, 1950). Transgres-
sion of the seal was noticed into the western part of the East Siberian Sea
(Ognev, 1935) and toward Wrangel Island. A male ribbon seal, 131 cm
long, was caught on November 16, 1962, close to Morro Bay in Cali-
fornia. [ts body was devoid of a hair coat except for the vertical surface
of the fore and hind flippers, head, and upper portion of the neck. The
animal was kept in an aquarium where it died a month later (Aryan
[Roset], 1964).

335 Fig. 182. Female ribbon seal with a newborn. Bering Sea (photograph by А.Р. Shustov).

336

447

Reproduction. The period of whelping extends from March end to
early May. In 1962, mating was noticed for the first time in the Bering
Sea on March 29 while newborns were encountered on Мау 2-3, 1963.
Mating takes place soon after parturition and thus gestation extends for
about a year. Most of the females are capable of producing an offspring
every year. Females attain sexual maturity at two to three years of age
and males a year later at three to four years. Females older than five
years are the most productive. The upper age limit for productivity has
not been established nor has any climacteric period been detected among
females. The intensity of reproduction is quite high and the number of
gestating females in some years varies from 30 to 60% or more of the
total number of females in the population (Shustov, 1965b).

Growth, development, and molt. The ribbon seal is characterized by
a latent period of embryonic growth [delayed implantation], roughly cal-
culated as 2.5 months. The newborn averages 85 cm in length, i.e., about
one-half the length of the mother. It reaches this length roughly after
nine months of uterine development. The average weight of the pups is
8.6 kg. The pups are covered with a long, soft, silvery-white embryonic
coat with smoky spots; this coat is sported for about two weeks. In this
period the pups feed on the mother’s milk. At 25 to 30 days of age, the
molted pups (normally molting pups and gray pups) measure an average
length of 112 cm and weigh 28.3 kg. The hair coat is short and coarse,
slaty on the back, gray on the flanks, and light gray on the belly. After
the first molt a dark band with sharp outlines appears on the generally
monochromatic dorsal background. At two years of age the white bands
characteristic of adults are seen for the first time (Fig. 183). At this age
the female acquires the adult coloration while the male of two (or some-
times three) years of age most often is very dull colored like the adult
female. Males acquire the adult vividness at 3-4 years of age.

The intense growth of seals in the first few years slows down after
maturity. Growth cessation among females occurs evidently at seven to
eight years of age and among males at 7-9 years. By this time the seals
are fully grown. Females lag behind the males in growth though not very
significantly (Table 25).

The longevity of the ribbon seal has not been established. The oldest
male studied was 27 years while two females were 26 years old. These
animals did not give the impression of senility (Shustov, 1965b; Shustov
and Yablokov, 1967; Tikhomirov, 1968).

Shedding of the embryonic hair coat constitutes the first molt, the
signs of which are noticed 7-10 days after birth. The hairs are initially
shed from the head and the hind flippers and later from the other sections

336

337

Fig. 183. Young ribbon seal. Bering Sea (photograph by А.Р. Shustov).

Table 25. Variation in body length of seals with age (measured along the dorsal
surface) (Tikhomirov, 1968)

Age, years Males
No. of animals
measured

1 16

2 20

3 29

4 33

5) 1

6 20

7. 17

8 19

9 16
10 21
ее 13
12 14
13-14 22
15-17 26
18 апа абоуе 15

Average
length (cm)

132.0
145.0
155.0
156.6
165.0
158.9
160.1
161.0
168.1
163.8
165.0
167.7
165.2
168.7
163.1

Females

No. of animals
measured

Average
length (cm)

129.0
145.0
152.0
153.9
160.2
161.5
168.0
169.0
163.9
163.0

163.9

168.8
168.1

of the body. The juvenile coat is preserved for quite sometime in the
armpits of the fore flippers. Molting ceases roughly at two weeks of age.

The molting of adult seals extends for quite a long period. The sec-
ond half of April can be regarded as the beginning of the molting period
but the cessation of this period has not been established. A ribbon seal
caught in the Sea of Okhotsk on May 16, 1939, was at the peak of

338

449

molt while another one caught on May 20, 1938, was found to be fully
molted. In mid-July, molting animals were seen among those that had
already completed molt. The duration of molting can be roughly put at
three months. During molt, not only a change of the hair coat occurs,
but often a simultaneous and intense peeling of the epidermal layer
(Pikharev, 1939, 1941).

Enemies, diseases, parasites, mortality, and competitors. The ribbon
seal has practically no enemies in the icy period. Only on occasion does it
fall prey to the killer whale, polar bear, or even a large Greenland shark.
No information is available on the subject outside this period although
there is undoubtedly greater possibility of death being inflicted by the
killer whale or shark. Diseases suffered by ribbon seals have not been
studied but sometimes animals with skin diseases are encountered. Such
animals are partly or even wholly devoid of hair coat, their epidermis is
peeled, and bleeding cracks are seen in the affected sections of the skin (a
seal in this state was caught in California). The mobility of such animals
is greatly reduced and they do not fight man. A blood analysis of three
sick seals gave an erythrocyte sedimentation rate of 18 to 25 mm/hr versus
3-4 mm/hr for healthy animals. Such seals, though few, are nevertheless
seen every year.

The following ten species of helminths have been detected among the
ribbon seals of the Bering Sea. Cestodes Diphyllobothrium sp. and Diplo-
gonoporus sp. have been found in the intestine. Nematodes of Anisakidae
gen. sp. have been found in the intestine and stomach; Contracaecum
osculatum Rud., detected in many species of Pinnipedia in the Northern
and Southern hemispheres, localizes in the intestine (not detected among
ribbon seals of the Sea of Okhotsk). Phocascaris phocae Host, known
only among harp seals as well as ribbon seals, parasitizes predominantly
the stomach.

Terranova decipiens Krabbe was detected only once in the stomach
of ribbon seals; it is known among walruses, sea lions, fur seals, elephant
seals, and many species of true seals. Another species of this genus, Ter-
ranova azarasi Yamaguti and Arima, was found in the intestine of ribbon
seals from the Sea of Okhotsk but not the seals of the Bering Sea. From
among the acanthocephalans, Bolbosoma nipponicum Yamaguti was also
found among many marine mammals of the Pacific Ocean basin. It local-
izes in the intestine, as does Corynosoma strumosum Rudolphi, known
among a large number of species of Pinnipedia and also among some land
mammals and birds. Corynosoma semerme Forssel, widely distributed
among Pinnipedia and cetaceans of the Northern hemisphere, is also
found among ribbon seals (Delyamure, 1955; Shustov, 1965c).

450

Natural mortality of this seal has not been studied. As in other
species, mortality is maximum in the younger age groups. Death caused -
by enemies is evidently insignificant but parasites can cause death, albeit
to a small extent. Maximum mortality arises from unfavorable ice con-
ditions and other abiotic factors.

The scattered disposition on drifting ice floes suggests that there
is по competition with other species for a place on them. Only at the
end of the icy period, in the years of early thawing, when many seals of
four species gather on the rapidly diminishing ice massifs, is competition
possible between the bearded seal, common seal, ringed seal, and ribbon
seal. There is hardly any competition in relation to food since the food
zones of all these species are distinctly “delineated”.

Population dynamics. The natural population dynamics of this seal
have not been studied but its fluctuations cannot be significant. In recent
decades this situation has undergone significant changes as a result ot
hunting. Before hunting with the help of ships was organized (1932), the
ribbon seal was caught by the local people in very limited numbers since
it essentially stayed far away from the coasts. This level of hunting had
hardly any impact on the seal reserves and the population maintained
a natural equilibrium. In the first two decades, hunting was carried out
only in the Sea of Okhotsk where the comparatively low level of killing
did not alter the natural state of the population. As the hunting fleet
enlarged in the early 1950s, and the catch of all species of seals increased,
some 20,000 ribbon seals were caught annually. An analysis of the age
composition of the seals caught revealed a gradual “rejuvenation” of
the population but pointed to a disturbance of the natural ratio of the
age groups and a reduction in the reproductive capacity of the popula-
tion. The results of these investigations formed the basis for suggestions
regarding the need to restrict the killing of the ribbon seal in the Sea of
Okhotsk.

Hunting of the Bering Sea population commenced only in 1961;
until then the population had remained undisturbed. An analysis of the
hunting activity of ships and the age composition of the animals caught
in 1961-1963 showed that the ribbon seals killed per day decreased con-
siderably irrespective of hunting conditions. The specific proportion of
the larga, which is the most difficult seal to kill, increased in the catch of
the ships; larga becomes the target only when seals of other species are
not available. The age composition of the catch also changed. In 1961,
the average age of the ribbon seals caught was 9.8 years; it fell to 6.9
years in 1962, and further dropped to 4.9 years in 1963. This points to
a reduction of the total population and its reproductive capacity since
females older than five years are the most productive (Shustov, 1965).

339

339

451

With expanded hunting in subsequent years for newborn pups because of
their valuable fur, the age composition of the animals killed somewhat
balanced but the state of the population still causes anxiety.

Field characteristics. Characteristic coloration is an unfailing feature
for recognizing the ribbon seal (Fig. 184). The seals are found only on
white, clean, and stable ice floes drifting far away from the coasts; they
do not form groups and are usually seen singly or in twos and threes.
In the summer months these seals adopt a pelagic mode of life; sites of
summer gatherings have not been established. (V.A.)

Economic Importance

The ribbon seal is of little importance to the local people who hunt in
the coastal zone where its population is very small. The proportion of
the ribbon seal in the ship catches in the Far East is not less than 30%
of the total catch (over 80% in the Bering Sea) and thus its importance
here is quite substantial. It should be borne in mind that the ringed seal,
which represents the largest number of seals caught, is a much smaller
animal and provides a smaller quantity of useful products compared to
the ribbon seal.

Fig. 184. Adult male ribbon seal on an ice floe. Bering Sea (photograph by
A.P. Shustov).

340

452

Seals of all the species are killed in the Far East using firearms from
motor boats dropped in the water from a schooner in the hunting region.
Seals resting on ice floes represent the predominant target since killing
the animals in the water inevitably results in great losses as most of these
killed in the icy period drown.

The products of hunting are the hide, fat recovered from the subcu-
taneous blubber, and the carcass. The blubber is removed at the site of
the kill (on the ice floe). Its further processing is carried out either on
some Suitably equipped ship or at the coastal base where the ships deliver
the raw products for processing. Salt is used as a temporary preservative
of raw skins on the ships. The hides of adult seals are mostly used as
raw leather while those of white pups and the better quality skins of
adult animals serve as valuable raw material for the fur industry. The
fat recovered is used for commercial purposes in tanning, metal, and
other industries. The carcass is a valuable product for the animal breed-
ing farms extensively organized in the Far East; the meat and bones
are used as feed for fur animals. The local population fully utilizes the
animals killed.

Both the populations of the ribbon seal are exposed to heavy hunting
(see “Population Dynamics’’). The results of investigations indicate the
need for an extremely careful approach to utilizing these populations.
Hunting norms should be fixed annually for each population on a ratio-
nal basis depending on its condition and strict adherence to the norms
enforced. (V.A.)

Genus of Gray Seals
Genus Halichoerus Nilsson, 1820

1820. Halichoerus. Nilsson. Skand. Fauna. Dagg. Djur., I, p. 376. Hali-
choerus griseus Nilsson = Phoca grypus Fabricius.

These are relatively large seals, the adults measuring about 2 m or
more (Lc). The facial portion of the head is noticeably elongated (more
than twice the distance from the eyes to the ear opening) and elevated.
Its height in front is almost the same as at any other point while the
upper contour of the profile draws an even or slightly convex line at the
bridge of the nose. The whiskers are flattened, with wavy edges. The first
and the second digits on the fore flippers are longer than the others.

The profile of the skull is seen as an almost perfect straight upper
contour from the occipital crests to the anterior margin of the nasal
bones; the height of the facial portion on the anterior margin of the
nasal bones is no less than the height of the cranium.

341

453

The anterior nasal opening is extremely elongated and opens upward;
the maximum width occurs in its posterior one-fourth. The nasal bones
are relatively short (not more than one-fifth the condylobasal length)
and broad; their total width at the anterior margin is about one-half
or more their length. On the posterior (orbital) side of the zygomatic
processes of the maxillary bones lies a horizontal carinate crest which
conceals the suborbital aperture when viewed from above. The length of
the zygomatic bones without process is more than double their smallest
width; the lower posterior process of these bones is considerably longer
than the upper. The features of the tympanic bullae, viewed from below,
appear as a triangle with rounded apices and resemble those in seals of
the subgenus Phoca s. str. The bony lobe of the external auditory meatus
is simple in design, flattened and broad.

The premolars and molars, except for the first molar, are massive;
the second and third premolars and molars of the upper jaw and also
the first three premolars of the lower jaw have conical crowns and lack
accessory Cusps; accessory cusps are seen on the fourth premolar of the
upper and lower jaws and on the lower molars; highly reduced acces-
ОГУ cusps are sometimes seen even on the initial premolars. A second
molar is sometimes seen in the upper jaw. At least the three initial
premolars of the upper and lower jaws among adults have a single root.
True molars, however, invariably have two isolated roots set in separate
alveoli. Incisors have laterally flattened roots. The dental formula is:

SA ЩАС. 1 2
5, Ст, Ру, M; = 34 of М = 36.

The neonatal pelage is white with a creamy tone; after the first molt
the hair coat is spotted.

The females are somewhat smaller than the males.

There is one pair of teats.

Ecologically, the genus is represented by two forms. One form inhab-
its the Baltic Sea and also partly the Canadian-Newfoundland waters and
is biologically associated with ice floes. These seals reproduce, suckle
the pups, and molt on ice floes. The herd instinct and polygamy are less
characteristic of this form. The population of the second form, inhabiting
unfrozen waters, exhibits no affinity for ice floes, spends part of the time
on the coasts, and forms rookeries of a distinct harem type. Whelping,
lactation, and partly even molt occur on the coast.

The distribution of the genus is restricted to the coastal waters of
the boreal belt of the North Atlantic and is now mainly confined to
three isolated portions of this belt: western Atlantic (American), east-
ern Atlantic, and Baltic. The American section is bound by waters from
the southern boundary of the Gulf of Maine almost up to northern

454

Labrador, including the Gulf of St. Lawrence and Newfoundland coasts.
The European section comprises the waters surrounding Iceland, the
Faeroe Islands, Ireland, and Great Britain, and also the coastal waters
from Bretagne and La Mancha to eastern Murman, including the North
and Baltic seas, and also the inlet zone of the White Sea. The boundaries
of distribution are mainly determined by the lines of the polar front and
heavy pack ice, continental shelf zone, and boundaries of the temperate
boreal waters. The distribution as a whole is characterized by interrup-
tions, being broken into fairly distinct isolated sections. Thus one of
these, the Baltic, is isolated from the eastern Atlantic which, in turn, is
wholly isolated from the western Atlantic.

The origin of the genus has been traced to the Pliocene as genus
Gryphoca van Bened. Deeper roots of the genus have not been detected.
It is possible that the Miocene seal, Miophoca vetusa Zapfe, represents
only one of the much older ancestral branches of this phylogenetic group.

These seals exhibit the most proximate contemporary genetic asso-
ciations with seals of the genus Phoca (especially of subgenera Phoca
s. Str. and Pusa) together with which they form the subtribe Phocina
(Chapskii, 1955).

The genus consists of a single species, the gray seal or “tevyak,” Най-
choerus grypus Fabricius, 1791, which constitutes 5.5% of the number of
species in the family. There is no basis for assuming that the genus con-
sisted of a larger number of species in the geological past. The economic
importance of gray seals is not significant.

The USSR is host to this single species, or 7.7% of the number
of pinnipeds in our fauna and 0.3% of the number of species in our
mammalian fauna. Its distribution is restricted to the Baltic Sea and
southwestern part of the Barents Sea. The economic importance of the
genus in our waters is altogether negligible. At places, some probably
damage the fish reserves by consuming particularly valuable fishes, i.e.,
salmon and eels. (K.Ch.)

GRAY SEAL!'®
Halichoerus grypus (Fabricius, 1791)

1791. Phoca grypus. Fabricius. Scrivter of Naturhist.-Selskabet, Kjoben-
havn 1, p. 167, pl. 13. Greenland.

118 “ТеууаК” is a name used by the coastal people but is not less common. Known as

“zhirovets” at places in Murman. Other names—“long-snouted,” “gray,” “hump-nosed,”
“pig,” and other artificial, bookish, often translated names—are almost out of use since the
coastal inhabitants are little acquainted with these names or this species.

342

343

455

1820. Halichoerus griseus. Nilsson. Scand. Fauna, Dagg. Djur., I, р. 377.
Greenland.

1851. Halichoerus macrorhynchus. Hornschuh et Schilling. Arch. Natur-
gesch., 17, p. 28: Baltic Sea.

1851. Halichoerus pachyrhynchus. Hornschuh et Schilling. [bid. Baltic
Sea.

1886. Halichoerus grypus var. atlantica. Nehring. Sitz.-Ber. Ges. Natur-
forsch. Freunde, Berlin, p. 122. Western coast of Norway.

1886. Halichoerus grypus vat. baltica. Nehring. [bid. Baltic Sea. (V.H.)

Diagnosis
Only species of the genus.
Description

General build of body massive, without distinctive features, while pre-
serving all the typical features of the subfamily of true seals (Fig. 185).
However, the snout is elongated, high; the profile at the eye level shows
no curvature. Some skin #014511? are seen posterior to the ear opening.

The whiskers are set in six rows on the upper lip; the first to the third

rows from below have 7-9 each. On the fore flippers, the first digit is
the longest (with the claw), the second slightly shorter, while the length
of the others decreases markedly toward the last.

The color of the hair coat reveals considerable individual variation
depending on age and sex and, further, is subject to some geographic
variation. The coloration essentially consists of two elements: the gen-
eral background and the fairly dark spots dispersed on it. The background
and partly the spots exhibit a wide range of gray tones and shades, from
extremely light silvery-white to, at places, almost white to very dark, and
even totally black. The main background in most cases is considerably
lighter than the spots and is usually different on the dorsal and ventral
sides, the latter almost invariably being perceptibly lighter. The spots are
of diverse sizes and shapes; the sharpness of their contours, number, and
disposition also vary. From very early times, many authors (Millais, 1904;
Collett, 1911-1912; Ognev, 1935; and others) attempted to group all the

19 This formation was described by Pocock (1933) who regarded it as a rudiment of the
outer ear shell. However, these skin prominences, measuring 3 to 15 mm (Mohr, 1952),
cannot be regarded as typical of the species. There is even less justification to view them
as analogous to the rudiments of outer ear shells. These skin folds are altogether lacking
in the Baltic gray seals.

342

456

Fig. 185. Gray seal (‘ЧеууаК”), Halichoerus grypus (figure Бу М.М. Kondakov).

diverse colors into two main types. One type covered animals with a rel-
atively lighter, contrasting spotted coloration and the other type animals
with a much darker, dull-spotted coloration. According to this scheme,
the light gray main background on the dorsal side of animals with light
coloration is somewhat darker and covered with innumerable, predomi-
nantly dark gray, blackish spots, sometimes scattered singly though quite
densely and sometimes gathered in clusters which are often fused. Some
spots have distinct contours while others are quite diffuse. The angular,
sometimes complexly contoured spots on the sides of the neck, on the
chest, and on the shoulder blades exhibit the utmost contrast. The spots
at these sites are also the most vividly pigmented, at places almost pure
black. On the dorsal side, the spots are duller and generally lack sharp
outlines.

The dark-colored type animals are distinguished by a very dark gen-
eral shade that depends not only on the darkening of the main back-
ground, but also on the intensity of pigmentation of the spots covering
it and the increased number of spots or their sizes. Concurrently, the
contrast of the pattern either disappears or weakens due to the indis-
tinct, diffuse contours of the spots. When the background is particularly

344

457

darkened and the spots are fused, the animals appear from a distance
to be monochromatically dark-colored, almost black. On closer exam-
ination, the spots are definitely apparent; furthermore, small, sparsely
scattered, light-colored specks are discernible on the dark skin, either in
streaks or blotches.

Various transitional coloration forms are noticed between these two
extreme types. The following features are characteristic of males as well
as females among all the diverse types. The dorsal side is gray or bluish-
gray, with fairly innumerable almost black spots; the underside of the
body is almost white with highly contrasting black spots that are partic-
ularly prominent in the anterior portion of the body.

In spite of the extensive studies on the color variation of this species,
its earlier descriptions (Millais, 1904, and authors citing Millais; Ognev,
1935; and others) are no longer adequate, 1.е., attempts to differenti-
ate two main color types—light- and dark-colored animals, or at least
the greater number of them. The information collected in the last two
decades has provided data pertinent to the prevailing situation. There
is no doubt about the existence of predominant color types but these
can now be more conclusively interpreted as individual, age-related, sex-
related, and evidently geographic variations. The magnitude of this vari-
ation differs in different population groups. Evidently, its range is most
extensive among the land-loving (pagophobic) gray seals inhabiting the
European and American Atlantic. The Pagophilic Baltic seals are evi-
dently characterized by a relatively greater color stability.

Only the Murman gray seal can be regarded as a representative of the
Atlantic population in our waters. Its color is very similar to that of the
British-Irish populations and is perhaps indistinguishable from the color
of the latter. It is characterized by as much range in individual varia-
tion as in the first year after the initial neonatal molt (Fig. 186). Among
such molted pups, the color intensity of the main background as well
as the nature of spots (size, shape, and sharpness) and the intensity of
their color vary; the color may be very light or sometimes very dark “on
any general background”. Wholly monochromatic black-colored animals
are also encountered (Karpovich, Kokhanov and Tatarinkova, 1967). A
similar color variation is seen among the juveniles of British-Irish and
Norwegian populations. In these regions pups are encountered with and
without spots, with a bluish-gray main background, even greenish, some-
times creamy-white or dark-colored or pure black; some animals are
brownish or even reddish-brown (Millais, 1904; Collett, 1911-1912).

Color variation among the adults is even more diverse. At times, it
is difficult to find two animals with totally identical coloration. Never-
theless, the color of Murman gray seals does not go beyond the various

ааа
Tee ~
съ SRG

SR Ne RA SS

344 Fig. 186. Individual color variation of molted juvenile gray seal, Halichoerus
grypus. Great Ainov Island, Murman coast (after figures by V.D. Kokhanov).

shades of gray and varies from almost white to almost or even totally
black. This is true of the main background on which the spots are dis-
persed and of the spots themselves. The main background is more often
of light gray shades, sometimes whitish and at places almost totally white;
specimens with a very dark, almost black background are not very rare.
345 On the underside of the body, the background color is somewhat lighter

459

than on the upper side, but this difference can sometimes be evened out
by the significantly greater spottiness on the throat, chest, and belly,
compared to the dorsal side on which the spots are frequently duller.

The spots vary widely in shape, size, sharpness, color intensity, and
density. In most cases the spots are diverse, often of queer shapes and
extremely dissimilar in size even in the same animal. The larger spots
have sharp as well as highly diffuse contours. The spots on the flanks and
underside of the body are the most intensely pigmented and contrasting;
they are often totally black. The spots are usually 10-15 cm in size; innu-
merable much smaller spots, down to mere specks or dabs, are scattered
between very large ones as well as separately from them, descending onto
the throat, occiput, and the base of the fore flippers. Extremely matte
and comparatively small spots sharply predominate on the dorsal side.
Thus the entire dorsal surface very often appears from a distance light-
colored and monochromatic. At the same time, animals are encountered
with totally dark or black coloration, this being characteristic of males.
The main background as well as the spots never contain admixtures of
brown, cinnamon, rust, or yellow shades but these are encountered, albeit
very rarely, among the animals in British rookeries.!”°

A very similar sharply manifest sexual dimorphism points to the
genetic kinship of the Murman gray seals with the other gray seals of the
open Atlantic waters breeding on the land. Adult males are distinguished
from females by a very dark and fairly monochromatic color. Males are
not characterized by contrasting spots and their body underside is only
slightly in color than the upper side. On the contrary, brighter coloration
is characteristic of the females, among which the main background is
much lighter, especially on the flanks and ventral side, where it is very
light. The spots on this background are highly contrasting but vary in
degree of sharpness of contour, which is sometimes well defined and
sometimes highly diffuse. These spots are concentrated predominantly on
the front portion of the body where even contrasting specks stand out.'?!

The age-related color variation among the pagophobic and
pagophilic forms has not been thoroughly studied. Evidently, in the early
stages of postnatal growth, their color differs only as much as the groups
under comparison differ in the range of individual variation. Pups of

120 The above color characteristics of the Murman gray seal are largely based on the
reports of V.D. Kokhanov, a zoologist in the Kandalaksh sanctuary.

121 The contrasting fanciful patterns of bright black coloration of the Murman gray seals
renders them visible from a distance even in twilight hours. Thus, several tens of animals
lolling on the coast appear “surprisingly similar to a herd of spotted cows at rest. Their
white skins appear variegated with large and small black spots” (Belopol’skii, 1951).

346

460

different ecological groups form herds of unmolted white pups. In most
of the pups of the Atlantic population, the short hair coat after the
long embryonal hair has been shed is almost the same as among the
pups of the Baltic seals. Its typical features are as follows: The main
background is pale gray, usually somewhat darker on the dorsal surface
than on the ventral. Comparatively small but numerous dark gray or
even Olive-colored spots, often fused, are scattered fairly uniformly on
the upper as well as the underside of the body. The sharpness of the spot
edges varies in different sections of the body of even the same animal as
also in different animals; in the anterior portion of the body, however,
and in particular on the flanks, chest, and neck, the spots mainly have
sharp outlines and hence stand out contrastingly. Such a coat among the
juveniles usually appears brighter and somewhat “fresher” than among
the older seals.

The molted pups of Murman reveal up to 12 distinct color varia-
tions which can be grouped into the following six classes: (1) almost
black, monochromatic; (2) dark gray, with very dark spots only on the
trunk; (3) similarly dark gray but with spots on the head as well as on the
flippers; (4) close to black, with light-colored spots; (5) with light gray
main background and dark, almost black snout and extremities of fore
and hind flippers; and (6) with the same light-colored background and
slightly darker colored head, flippers, and spots of the same shade scat-
tered on the trunk (V.N. Karpovich, V.D. Kokhanov, I.P. Tatarinkova).

The pups from the Baltic Sea after shedding the embryonic pelage
appear more monotypical. They are then gray with innumerable blackish,
partly fusing small spots. Instances of black or almost black coloration
have not been seen among these pups. The spottiness covers the entire
head and extends to the occipital, temporal, and throat regions (Lillje-
borg, 1874; Allen, 1880; Schubart, 1929; Lonnberg, 1929; Freund, 1933;
Mohr, 1955; and others).

The subsequent color changes of the hair coat until the animal
attains maturity have not been adequately understood. Some authors
(Collett, 1881; Millais, 1904; Mohr, 1955; and others) believe that the
original vivid and contrasting spots (with sharp contours) are gradually
lost as the animals grow up, giving place ultimately to a uniformly gray
or very dark coloration. But this is not actually so. There is no single
pattern of age-related color variation that can be regarded as common to
all the animals or to different populations. This is indirectly supported
by the considerable diversity of the original coloration of the pups, espe-
cially the specimens that are found to be almost black from the moment
of casting the embryonic pelage. Although there is no possibility of their
turning blacker later, even these pups change color. Thus, one gray seal

347

461

that lived for several years in the basin of the Bergen Biological Station
was almost black with a brown tinge at the age of one month; it began to
lighten in color after two years (especially the main background between
the spots) and roughly ten years later again turned very dark (Collett,
1911-1912). Observations of two juveniles sporting different colors in
the Berlin Zoological Garden showed that the hair coat of the light-
colored animal became even lighter and spottier after each molt, while
that of the dark-colored animal, on the contrary, became darker so that
the spots became increasingly less distinguishable (Nehring, 1887).

Whatever the pattern, the pup at the transitional age is characterized
mainly by a transitional color with larger, more contrasting spots than
pups that have not yet attained the final pattern (or color type) char-
acteristic of the adults of the corresponding sex. Evidently the young
animals do possess some features reflecting their affinity to one or the
other ecological groups: Atlantic or Baltic. The growing juvenile from
the Baltic Sea is apparently characterized by a gradual loss of the con-
trast of spots from year to year; the spots either become less distinguish-
able on the generally darkened background (Ropelevskii, 1952), or they
disappear altogether (Mohr, 1965), or become diffuse or sparser. Among
the males in general, the spots are lost sometimes in a wholly dark back-
ground (Aul, Ling and Paaver, 1957). The spots are less sharp among
young animals of transitional age and among the Canadian population
(Mansfield, 1963). On the other hand, the view has been expressed that
the color and spottiness of at least part of the British-Irish gray seals
are as diverse among young animals as among adults (Millais, 1904). On
the whole, a clear picture of the age-related succession of color changes
among the various populations is not yet possible.

The process of color variation in the much later stages of age pro-
ceeds divergently, i.e., begins to show sex-related differences. Concomi-
tantly, changes in the configuration of the spots characteristic of the
males and females continue and many spots enlarge and lose their sharp-
ness of contour.

The color of the adult males of the Baltic seals is generally quite
uniform and falls in the category of spotted coloration described above,
being only somewhat duller. The main background of the dorsal side
of the body is Smoky-ash with dove-olive tones. It is noticeably darker
on the flanks and on the ventral side and the boundary of the back-
ground color is often quite distinct. Innumerable dark-colored spots are
scattered all over this general background. On the dorsal surface, they
are mostly small with diffuse outlines that are generally regular (less
often, highly sinuous) and not very contrasting. On the flanks and ven-
tral side the spots are denser, appear very dark (black), often fuse with

462

each other, acquire very complex outlines, and stand in sharp contrast
against the light-colored background. The spots are noticeably larger in
the anterior half of the body than in the posterior half; the spottiness
on the belly is paler and quite often looks like marble streaks. The spots
become smaller around the head and on the throat and look like dabs.
Fine specks are sometimes seen even on the occiput, sometimes extend-
ing also onto the sinciput; on the sides of the head, near the eyes and
ears, however, the spot-dabs are as numerous as оп the throat. The snout
and the upper portion of the head are also quite often monochromatic,
grayish dove-colored. Quite often, too, fine specks descend even onto the
fore flippers. On the lower side of the neck and on the chest, the spots
form a typical pattern, more often a longitudinally elongated garland of
fused dark gray spots with diffuse contours standing out sharply on a
very light-colored background. On the belly the spots are not only paler,
but their outlines more diffuse.

Adult female Baltic seals, compared with males, are generally lighter
in color and less spotted (Aul, Ling and Paaver, 1957). It is possible that
the spot pattern on the neck and chest is better developed among females
than males (brighter and more contrasting).

The hair coat fades somewhat in the course of the year. Among
the British-Irish and Norwegian populations, the skin turns very pale by
spring and the spots lighten; they turn a vague brown on the dorsal sur-
face while the main background between them fades to a monochromatic
straw color (Millais, 1904; Collett, 1911-1912).

Sex-related color differences are quite sharp among the Murman
gray seals. Adult males are usually dark, almost black, with spots that
appear indistinct from a distance. Only the underside of the body is
lighter with dark spots being fairly distinct. The females are lighter and
more spotted than the males although dark females do occuroccasionally
(V.D. Kokhanov). Seven basic color variations have been recognized
among the females. The males and females exhibit deviations (roughly
12%) wherein the males exhibit the color features characteristic of
females and vice versa. Sex-related color differences are seen among
Murman gray seals right from the earliest age, 1.е., as soon as the pup
begins to shed the original (embryonic) pelage.

The main body background in most young females is light gray with
scattered darker gray spots. Less frequently, the spots are black on a
light gray or even darker background. A very dark, almost black main
background (lighter on the underside) predominates among the males; in
the first case the scattered spots are black and in the second light colored
(lighter than the main background) (V.D. Kokhanov, V.N. Karpovich,
1.Р. Tatarinkova).

348

463

Sex-related color changes are also similar among the Norwegian
(Collett, 1911-1912) and British-Irish (Hewer and Backhouse, 1959)
populations of gray seals. This confirms the greater affinity of the Mur-
man gray seals with the European-Atlantic population than with the
Baltic. The adult males of the Norwegian population are dark-colored,
almost totally monochromatic blackish-brown; the lighter-colored ani-
mals show a dark gray main background alternating with large brownish-
black spots. The females are lighter in color: the main background is
either brownish-gray or silvery-white as in the juveniles and the spots
sometimes more, sometimes less vivid; sometimes spots occur only on
the shoulders; some animals are totally devoid of spots and appear fairly
monochromatically white from afar; the spots vary from light brown to
very dark (Collett, 1911-1912).

The sex-related color differences of the hair coat are similar among
the British-Irish populations. The main background in adult males is a
uniform dark color interrupted quite often by not very contrasting and
rather small clearances in the form of angular white spots and motleys,
sometimes with complicated patterns. In females, innumerable dark spots
are scattered on a very light-colored main background (usually differing
in vividness on the dorsal and ventral sides). These spots are invari-

ably darker than the main background and the contrast is quite good

in most females. Among the dark-colored males, the main background
is close to black with indistinct small and narrow white streaks show-
ing through. Among the dark-colored females, the main background
is dark gray and the spots are numerous, quite large, and deeply pig-
mented, almost black. Light-colored males are distinguished by a very
pale, brownish-gray main background covered with very bright, some-
times almost white streaks. Light-colored females have a light gray main
background covered with comparatively sparse and small spots (Hewer
and Backhouse, 1959).

Coloration is also similar among the Canadian gray seals. The males
sport a dark gray, almost black, background color patterned with tiny
dark gray spots on the flanks which may fuse into large groups. The
overall impression is a very dark coloration. The color of the females is
smoky-gray dorsally, turning silvery-gray or even white along the flanks
and on the belly. In much older animals, the dark gray or black spots may
have fused. The female in general is lighter in color and more spotted
than the male (Mansfield, 1963).

Among the females in the British section of the range, stray speci-
mens are encountered, albeit very rarely, with an orange-rust-red head,
neck, and lower surfaces of the flippers, and sometimes rust-colored
flanks; elsewhere the body color remains normal (Backhouse and Hewer,

349

464

1957, 1960). Animals with this type of coloration have not been reported
in our waters and it has been emphasized that even brown shades are
not found among the Murman gray seals (V.D. Kokhanov).

The skull is massive, the facial portion appears elongated, and the
rostral portion broad (Fig. 187). Among adults the width between the
zygomatic bones noticeably exceeds that of the skull between the mastoid
processes; the maximum lateral bulge of the zygomas falls close to the
posterior margin of the orbit, in the posterior half of the skull. The
interorbital space is moderately compressed (its width is not less than
15 mm among the adults).

The general appearance of the skull resembles more that of a large
larga or Atlantic common seal but differs in the structure of the anterior
nasal opening, nasal bones, massive rostral portion, nature of crests,
form of the lower jaw, and especially the sharp structure of the crowns
of the premolars. The broad and short nasal bones have a shortened
apex, forming more than half a right angle, with rectilinear sides. The
anterior margin of the nasal bones has three deeply incised denticles. The
nasal processes of the pre-maxillary bones reach the nasal bones but only
adjoin them (not invariably though) and do not wedge deeply between
them and the upper maxillary bones. The longitudinal bony septum in
the choanae falls far short of the posterior edge of the bony palate incised
in the form of a deep arc. The anterior palatine openings are large, with
a deep troughlike pit anterior to each. The width of the tympanic bullae
together with the bony lobe of the external auditory meatus exceeds their
length. The sagittal crest is straight, narrow, and relatively low.

Most of the premolars and the upper molars have a high conical
crown, without accessory cusps. The latter are developed, though not
intensely, only on four premolars and lower molars (and often also on the
lower initial premolars). These are in the form of small denticles disposed
one each anterior and posterior to the main cusp at its base. Intensely
reduced and barely visible denticles are seen even on some other pre-
molars at the base of the crowns. The canines are relatively large (the
longitudinal diameter of the alveolus is more than 10 mm); the extreme
lateral incisors in the upper jaw are somewhat larger than the rest.

Individual skull variations are significant in terms of absolute units
as well as ratios. The structure of the crown of premolars is variable.
Sex-related differences in the skull ar manifest only in sizes and some
ratios; the skull of males is larger.

Age-related skull variations are seen primarily in the elongation of
the skull due to the predominant growth of the facial portion which
becomes more massive and increases in height. Among young animals,
the facial portion falls much below the occipital and the line of the upper

349

350

465

Fig. 187. Skull of the gray seal, Halichoerus grypus (figure by М.М. Kondakov).

profile of the skull is intensely inclined toward the anterior margin of
the nasal bones. The width at the zygomatic arches is less developed
among young animals than adults; it does not exceed the width at the
mastoids in the young; the width of the rostrum at the level of the upper
canines too is relatively smaller than in adults. The contour elements of
the cranium (crests) are almost undeveloped among young animals. The
sagittal crest is not manifest: instead, only folds are seen to the right
and left of the median. These folds slightly diverge sideways and form
a small elongated area; occipital crests, seen as cornices in adults, are

466

practically absent in the young. As in other seals, the roots of the teeth
are initially hollow and thin-walled but intensely thicken over time and |
the pulp cavity fills in. Among many old seals, the roots are unusually
hypertrophied due to the cement layers while the crowns are worn down
almost to the base. Over time, all the skull bones thicken, as a result of
which the overall weight of the skull also increases.

The individual and partly the geographic variation of body size is
significant; females are generally smaller than males. The body length
of the adults measured from tip of nose to tip of tail along the dorsal
surface (Lc) varies from 170-250 cm. This length in the largest animals
can reach 3 m. However, various authors cite different values: 2-2.5 m
(Freund, 1933), 2.5-3 m and at places where these seals are killed most
intensely (southern Baltic), mostly 2-2.5 m (Mohr, 1952). According to
the old data (Lilljeborg, 1874), the body length of the Baltic male gray
seals is 195 cm and that of females 175 cm.

Data on the body length of male Murman gray seals are extremely
limited and not wholly reliable. According to some authors, the average
size of an adult animal is 2.5-3 m (Skvortsov, 1928*) and according
others (Belopol’skii, 1951), the males are 2.5 т long. A large male caught
on Franz Josef Land measured 263 cm along the dorsal surface (Lc)
(Tsalkin, 1936).

The body length of gray seals of the British and Irish populations
(from which the size of Murman gray seals can be judged to some extent)
varies from 245 to 260 cm in males (largest male measured 292.5 cm)
and 168 to 198 cm in females (Millais, 1904). The average length of adult
Canadian male gray seals is 235 cm, of females 200 cm (Mansfield, 1963).
The available information on weight is also in general terms. It ranges
from 130 to 320 kg for seals of both sexes. During lactation, the females
lose over 30 kg of weight and males in the mating period even more than
50 kg (Collett, 1911-1912).

The condylobasal length of the skull is 240-270 mm and its maxi-
mum length [?] 330 mm (Collett, 1911 - 1912). This length among females
is considerably less than among males: 264 mm in the Baltic male seals
and 240 mm in females (Lilljeborg, 1874). In the largest males of the
Atlantic region, it reaches 320 mm (Allen, 1880) and even 330 mm (Col-
lett, 1911-1912) white it hardly exceeds 260 mm in the largest of the
females. The width of the skull at the zygomatic arches reaches 200 mm.
(K. Ch.)

Taxonomy

Only species of the genus.

351

351

467

Geographic Distribution

Temperate latitudes, partly the cold belt of the North Atlantic from
Canada and the adjoining regions of the USA to the coasts of central
and Northern Europe and the Baltic Sea. The range is broken into fairly
isOlated sections.

Geographic Range in the USSR (Fig. 188)

This covers the eastermost part of the world distribution and includes
two distinct regions: coastal waters of Murman and the Baltic Sea.

The gray seal is distributed all along the coastal belt of Murman from
Varanger Fjord to St. Nos Strait and possibly farther in the southeast
up to the western threshold of the neck of the White Sea. However, the
find of the gray seal on the eastern extremity of the range is evidently
sporadic and it is only very rarely that this seal is found in the neck
region itself (Schrenk, 1955*; N.A. Smirnov, 1929; Kirpichnikov, 1932).
Details of the seal distribution and the disposition of all of its coastal

Fig. 188. Distribution of gray seal, Halichoerus grypus in the USSR (K.K. Chapskii).

352

468

rookeries have not yet been established. The gray seal reproduces for
certain or resides regularly on the Bolshoi (Large) and Malyi (Little)
Litskie, Veshnyak, Kuvshin, Malyi Zelenets, Kharlovskie (“baklyshes”),
Semiostrovnye (“ludy”), Ainov and other islands, and in the region of
Dal’nii Zelenets, lokan’gi, Teriberki, and Pala Bay. The gray seal whelps
presumably in the bays of Rakushnaya, Voyak, Fedotovsk, and Kora-
bel’naya (Karpovich, Kokhanov and Tatarinkova, 1967).122

The view that the eastern boundary of distribution extends up to
Novaya Zemlya, based on the record of this seal by K. Ber on the west
coast of Novaya Zemlya, is erroneous. This is an exceptionally rare event
of the straying of an animal far beyond the range of its distribution. Very
similarly, a lone gray seal was also caught in Franz Josef Land archipelago
(Tsalkin, 1956*). The same is the situation with regard to the White Sea,
which was included in the distribution of this species (Schrenk, 1855*),
although episodic finds of lone animals have been reported from eastern
Murman in the inlet region and the adjoining neck regions (N. Smirnov,
1929; Kirpichnikov, 1932; A.G. Beloborodov; and others). It is quite
likely that lone animals even more rarely can penetrate into the northern
part of the neck region too.

In our Baltic waters the gray seal is encountered from time to time in
the northern coastal belt of the Gulf of Finland from the boundary with
Finland to Nevsk Bay, while it is quite common in the southern coastal
zone commencing from Kopor Bay and farther west. It is encountered on
the Estonian coasts all along the Gulf of Finland but is mainly confined
to regions far removed from the mainland coast, with a preference for
the tiny islets scattered there (Aul, Ling and Paaver, 1957). It inhabits
Irben Strait and the Gulf of Riga (its Estonian as well as Latvian sec-
tions) including Rukhnu Island. Evidently, from time to time, especially
in water, it is seen in the Latvian marine coastal waters. Rare encounters
in the coastal belt of the sea within Lithuania and Kaliningrad districts
are possible since gray seals were found in the past on the coasts of the
former eastern Prussia, Pomerania and Ryugen Island, as also on the
coasts of the former Polish corridor.

122 The first reference to the distribution of the gray seal beyond the Polar Circle in
our territory was made by I. Lepekhin (1805). Concrete evidence of the catch of gray seal
in Varanger Fjord (Pleske, 1887), on the coasts of eastern Murman (N. Smirnov, 1903),
and on Litskie Island (Telegraph of Murman Research and Hunting Expedition, 1906)
became available much later. It took an even longer time for more detailed information to
become available (Skvortsov, 1928*; Sdobnikov, 1933*; Belopol’skii, 1941, 1951; У. Uspen-
skii, 1941*). New data have been collected in recent years by the workers of the Kandalaksh
Sanctuary.

353

469
Geographic Range outside ше USSR (Fig. 189)

In the Baltic Sea, the range covers the Finland portion of the Gulf of Fin-
land, Gulf of Bothnia, northern and partly central regions of the Baltic
Sea itself (from the west, its Swedish belt). To the south of Gotland
Island, it is encountered more rarely, in the south up to Eresunn Strait;
it is rare in the southern and southwestern parts of the sea, including the
coastal waters of the Polish People’s Republic, the German Democratic
Republic, on the adjoining coasts of Denmark, and the Federal Repub-
lic, on the adjoining coasts of Denmark, and the Federal Republic of
Germany where it is encountered sporadically. The gray seal is evidently
even more rare now in Denmark Strait although the breeding of this seal
in the past on Anholt Island in Kattegat has been recorded.

Outside the Baltic Sea, the population in the eastern Atlantic section
is concentrated mainly in three regions where the seals breed and
reside permanently: 1) region of Great Britain and Ireland including
the Hebrides, Orkney, Shetland, and Faeroe islands; 2) coastal waters of
Iceland (excluding evidently the northeastern section); and 3) Norwegian
coast, predominantly its middle portion where the most important
(though not many) breeding sites are concentrated. In general, however,
the boundaries of distribution in the European part include the coastal
mainland from Bretagne (sporadic), La Mancha, Pa-de-Kal along the
southern coast of the North Sea up to the Jutland Peninsula, joining
there with the Baltic and Norwegian sections of distribution extending up
to Nordkapp. The presence of this species on the coasts of Spitsbergen
is rather doubtful in spite of positive references to it (Saemundsson,
1939).

There are no settled populations on the coasts of Greenland but stray
animals and probably even small groups wander there from time to time
(Brown, 1868; Allen, 1880; Winge, 1902; and others). The American part
of the distribution includes the Gulf of St. Lawrence and the adjoining
regions of open coastal waters in the north from the Strait of Belle Isle
along Labrador roughly up to the latitude of Hebron (about 68.5° N
lat.) and in the south partly along the west coast of Newfoundland, to
the south and southwest of this island and Cabot Strait, including Sable
Island, coastal waters of Nova Scotia, Gulf of Maine, and up to Cape
Cod inclusive. (K.Ch.)

Geographic Variation

The geographic variation of the gray seal has not been adequately studied.
Nonetheless, the species distribution is quite fragmented into distinct

352

470

Fig. 189. Species distribution of the gray seal, Halichoerus grypus (К.К. Chapskii).

sections with extremely incomplete contacts between them. First of all,
the western Atlantic (American) zone of habitat of the species is practi-
cally isolated from the eastern Atlantic (European) zone since gray seals
do not reside regularly in the waters of southern Greenland and are
sighted rarely and irregularly.

The Baltic zone of distribution is largely isolated from the Atlantic
Ocean zone. In the remote past, gray seals resided undoubtedly even in
the Denmark Strait and a contact between the populations of the Baltic
and North seas did exist though not wholly constant. A real interruption
of distribution evidently prevailed for some centuries (for more details,
see “Geographic Distribution”).

Two subspecies of gray seal can be recognized in our waters at
present.

1. Baltic gray seal or “tevyak,” H. g. macrorhynchus Hornschuh et
Schilling (syns. pachyrhynchus, baltica) (Fig. 190).

The body and skull dimensions are comparatively smaller than those
of the nominal form. Sexual dimorphism in dimensions is not very sig-
nificant. The color is generally very light with fairly bright, though often
diffuse, contrasting spots on the chest and neck with relatively less vari-
ation. There are practically no animals in the population with intensely
dark, almost black coloration. Sex-related color differences are relatively
insignificant.

It occupies Gulf of Finland and the waters of Estonia, Latvia, Lithua-
nia, and Kaliningrad district.

353

354

471

Fig. 190. Baltic gray seal. Leningrad Zoological Garden (photograph by К.К. Chapskii).

Outside the USSR, the Baltic gray seal is found in the rest of the
Baltic Sea regions. The form differs not only in morphological, but also in
its biological relation with ice floes on which it breeds (pagophilic form).
2. Atlantic gray seal or “tevyak,” НЯ. в. grypus Erxleben, 1777 (syns. gris-
seus, gryphus, halichoerus, gris, thienemanni, atlantica) (Fig. 191).

The body and skull dimensions on average are slightly larger than in
the preceding form and sex-related size differences are more significant.

The color on average is darker with somewhat less sharp and con-
trasting but extremely variable spottiness. Animals are encountered with
very dark, almost black coloration. Sex-related color differences are more
intense than in the preceding form.

It occupies the Murman coast.

Outside the USSR, this subspecies is found in the rest of the area of
distribution of the species, at least in the European waters in addition
to the Baltic Sea.

Seals of this form exhibit no affinity for ice floes during the period
of breeding (littoral form).

Assigning the nominal name of the subspecies to Murman and gener-
ally to the European seals of the oceanic waters is presently still tentative
since the waters of southern Greenland represent the terra typica of this
form. It may be assumed that the animals existing there are identical with
the American form. However, a comparative systematic analysis of the
European and American gray seals has not been attempted to date. Yet,
while taking into consideration the quite complete (though not absolute)

354

472

Fig. 191. Murman gray seal in water. Great Ainov Island, Murman coast, Мау,
1964 (photograph by V.D. Kokhanov).

isolation of these two large populations, it must be considered that they
are not identical at the subspecies level. If this is so demonstrated, the
name grypus should be given to the American gray seals while their Euro-
pean counterpart should be designated by the closest recent synonym.

Nevertheless, the systematic homogeneity of the American form may
give rise to some doubts since its composition includes pagophilic as well
as aegialoid [littoral] populations. It is possible that some differences do
exist between the various populations of seals in European waters.!79

Outside our waters, these animals are not classified into particular
subspecies.

Biology

Population. Even an approximate census of the population has not been
possibie in our Baltic Sea waters. It was difficult to establish even the
relative sizes of the local populations. Evidently, these seals are least
numerous in the easternmost portion of the Gulf of Finland in the region
of Vyborg (Freund, 1933). In Nevsk Bay only rarely are stray young

123 According to the latest data (К.К. Chapskii), fairly stable craniological differences do
exist between the Atlantic and Baltic subspecies.

355

473

seals encountered and that, too, only in the summer-autumn season;
the population increases westward. This seal is regarded as common in
Estonian waters.

In the Gulf of Riga it is relatively more numerous around autumn,
especially during the formation of ice floes, in search of which the seals
traverse Irben Strait (Aul, Ling, and Paaver, 1957). South of the inlet
into the gulf their number decreases sharply down to rare single ani-
mals for many kilometers of the coastal zone. The nature of the coastal
zone, especially the ruggedness of the coastline, and also the extent of
human habitation and hunting activity—all these greatly influence the
population of this seal.

The numerical strength of the local seal population changes sharply
in different seasons of the year; thus a greater concentration of the seals
occurs in the period of breeding and a sparse population in the feeding
season. Factors such as the formation of ice floes, their distribution and
suitability for whelping, as also the availability of food play a decisive
role.

While conducting a census of the world reserves of the gray seal, the
strength of the Baltic Sea population was also evaluated. This evaluation
was mainly based on information regarding the number of animals killed.
The statistics were incomplete because of the nonavailability of data for
our waters. In the first evaluation of the world reserves of gray seals
(Lockley, 1954), a figure of 5,000 was cited for the Baltic population,
exclusively for the Gulf of Bothnia. The view was later expressed that the
total reserves in the Baltic Sea should be in the tens of thousands (Davies,
1957). Practically, however, in subsequent evaluations the total Baltic
population was roughly put at 10,000 (Haglund, 1961) ог 5,000 - 10,000
(Hook, 1964*; Curry-Lindahl, 1965). A more realistic figure lies between
7,000 - 10,000.

With this background, it is as yet difficult to reckon the population
in the waters of the USSR. According to preliminary data (1972), the
mother population of our gray seal in the main region of its winter-spring
concentration, i.e., in Irben Strait with adjoining sections of the Gulf of
Riga, was roughly put at 350-400. In our zone of the Gulf of Finland,
however, the strength is a mere 50 (V.A. Zheglov).

The total strength of our Murman population was put at 300 in
foreign compilations (Smith, 1966*). A direct count of the newborn on
all the islands of the archipelago of Semi Islands and on the Ainov
Islands in the early 1960s identified a total of 200 seals. By applying the
method of extrapolation to other regions in which breeding is wholly
likely, the total population of annually reproducing females of gray seals
has been put at 500-800 (Kandalaksh Sanctuary; Karpovich, Kokhanov

356

357

474

and Tatarinkova, 1967). The main population (without pups) of Murman
seals constitutes roughly 1,500.

The population of the gray seal in other sections of its range is as
follows: 34,000 in the waters of Great Britain of which 5,000 to 8,500 seals
are concentrated on the Orkney Islands, Northern Rona, Outer Hebrides,
and Farne Island; 2,000 in Irish waters; and 2,000 on the Norwegian
coasts. The population of Iceland has been put at the same number as
on the Norwegian coasts and on the Faeroe Islands at 3,000. On the
whole, for the eastern Atlantic part of the area of distribution there
are 42,500 seals (not counting the Baltic Sea and Murman population).
In the western Atlantic (American) part of the distribution, the total
population runs roughly into 5,000, of which 3,000 are in the Gulf of St.
Lawrence and 2,000 east of Nova Scotia (Smith, 1966*). The total world
reserves Of gray seals thus come to 55,000 - 60,000.

Habitat. The habitat is closely associated with ecology, which for
this species is of a twofold nature, as in the case of seals of the subgenus
Phoca s. str. (common seal and larga). In our, as well as in international
waters, there are two ecological forms of the gray seal. One of these is
biologically associated with the ice floes on which they breed and molt.
These are typical pagophilic (ice-loving) animals. This form includes seals
of the Baltic Sea and the Gulf of St. Lawrence. The seals in the rest of
the European region, including our Murman population as also some
populations of the Nova Scotia Islands, reproduce on the coasts. In this
season they form fairly cohesive rookeries at definite places on the coast
year after year and can be regarded as wholly pagophobic. It is usually
assumed, although not always justified, that the pagophobic populations
of gray seals are more stationary and reside longer on the same sec-
tions of the coastal zone, usually not straying far into the open sea. For
rookeries, they select uninhabited, predominantly small islets, inaccessi-
ble to man, and made up of smoothly rolled rocky or more rarely pebbly
coasts with a flat and fairly level surface. They are disposed right on the
coast, slightly above the littoral zone as well as away from it in the more
elevated sections, including those overgrown with grasses.

Usually, the flat rocky coasts of islands slightly above the line of high
tide serve as the site of whelping on Murman but at times the semisub-
merged littoral sections are also used (Fig. 192); a distinct preference is
shown for smoothly- rolled boulders. Sometimes the ascent of the coast
is quite steep (up to 30° or more) but matters not if it is smooth. It is
not important how much of the slope is covered with a thick layer of
snow. Beaches with boulders or pebbles are rarely selected for whelping
(Uspenskii, 1941; Karpovich, Kokhanov and Tatarinkova, 1967). On the

356

475

Great Ainov Island (in the region of Varanger Fjord), the sites of whelp-
ing fall predominantly in the littoral zone toward the open sea but are
almost wholly absent on the southwestern side turned to the mainland
coast. Often, they are disposed there in sections with peat clumps among
tall grasses (lyme grass) (V.D. Kokhanov).

Our Baltic pagophilic populations during the breeding season are
disposed close to the outer rim of drifting ice floes, abounding in fissures,

Cape Polenyi

©) Lake Severnoe

Lake Zapadnoe
O Lake Nedupoynoe

AONIV. LVayy

Sy
1S)
(oop

О
—
о
en
=
ise}
—
Go}

>

axv Isl

О Lake Srednee
Lake Maloe
Cape Gagachiya

Littoral

Disposition of Gray
Seal pups

Cape Robinson
Hut

Fig. 192. Figure showing the disposition of newborn gray seal pups on Great
Ainov Island, Murman coast, November - December, 1963 (after V.D. Kokhanov).

476

open water pools, and partings, which are essential for keeping contact
with the water. Only in very rare cases, when the winter is extremely
mild, do some animals use the coastal rocks for whelping (Curry-Lindahl,
1965). Under the influence of ice floe movements, the fissures develop-
ing among them, and the expansions and contractions of the open water
pools, considerable changes occur in the original distribution of the new-
borns over the course of time: some pups and their suckling mothers drift
with the ice floes to the very edge or sometimes deep into the ice massif,
thereby being islanded in the dense ice “conglomerate”.

The Baltic seals are usually regarded as more pelagic (Curry-Lindahl,
1965) than the British animals whose coastal rookeries in summer are
very specifically fixed.

Food. In our waters this aspect has been thoroughly studied only
recently. There are practically no definitive data on the food of the Mur-
man gray seal. Judging from observations, however, lumpfish and cod
assume a prominent position in its food (V.D. Kokhanov). Only gen-
eral information is available on the food of the Baltic gray seal. On the
coasts of Estonia, it feeds on Baltic herring, cod, viviparous blenny, eel,
salmon, and bream (Aul, Ling, and Paaver, 1957). The animals held in
the Leningrad Zoological Garden (caught in the Gulf of Riga) consume
diverse fresh fishes from the Gulf of Finland and Lake Ladoga including
such freshwater fishes as roach, ide, dace, rudd, bream, etc. In nature, at
February end to early March, 1972, only the remnants of benthic fish, 1.е.,
viviparous blenny and flounder, were found in the stomach of whelped
females of the Baltic population (V.A. Zheglov).

The gray seal of the west European and Canadian populations is
usually regarded as a consumer of predominantly a wide variety of fishes.
The instance of catching a seal on a baited hook dropped to a depth of
145 m and also the repeatedly reported facts of the emergence of these
seals with deep-water fish between their teeth (Collett, 1881), point to
the possibility of these seals “fishing” at considerable depths. In the
European seas the gray seals consume, in addition to the above-listed
fishes, halibut, other flounders, herring, ling, whiting, and other cods and
pike. Apart from fish, they catch cephalopod mollusks (squids—Loligo
forbesi and Eledone cinerosa were noticed in the Irish Sea) and at times
consume even deep-sea crustaceans (Collett, 1911-1912; Freund, 1933;
Duncan, 1956; Lockie, 1962). In the stomach of some tens of gray
seals caught in the Gulf of St. Lawrence, the following were identified:
among fish—mackerel (?), cod, herring, salmon, hake, small winter
flounders, unidentified species of flounders, skate, and shark; among
invertebrates—squids, crabs, and shrimps (Fisher and Mackenzie, 1955;
Mansfield, 1966).

358

477

The diet of the gray seals varies considerably depending on age,
season, and local conditions. These factors are also responsible for the
magnitude of losses suffered by the fishing industry (see p. 495). Accord-
ing to some sources, the gray seal is such a voracious eater that it can
consume its own weight in fish in a single day in captivity (Legendre,
1947). The daily ration of caged gray seals abroad is 6.8 kg (Steven, 1934,
1936). They receive a slightly larger ration in the zoological gardens of
Leningrad and Kaliningrad where the seals enjoy good health.

The stomach of one animal shot contained 27 completely whole
eels that had evidently been swallowed in a very short period of time
(Curry-Lindahl, 1965). In another animal 54 common perch and one
pike, in addition to several herrings, were found in the stomach; it was
said that an adult seal can consume 60 to 80 herrings daily (Friedel,
1882). However, .it should be noted that, in nature, it feeds at inter-
vals during the periods of reproduction and molt. Evidently lactating
females feed sporadically, with intervals of starvation occurring during
the summer-autumn season.

When the fish is large (not in the case of an eel), the gray seal tears
it apart with the claws of its fore flippers and consumes it in bits. Smaller
fish, such as the herring as well as the eel are swallowed whole (Nehring,
1887). These habits have been recorded during the food intake of seals
at the Leningrad and Kaliningrad Zoological Gardens.

Home range. Historically, each local population has maintained a
fairly stable home range. This is primarily and especially true of pago-
phobic populations. Observations of the British-Irish populations have
established that the bulk of the mother population whelps year after year
on the same coastal section. Nevertheless, the animals are not confined
to the same site year round. Even the most settled among them abandon
the rookery at the end of reproduction, lactation, and molt, wandering
from place to place along the coastal belt. Therefore, establishing the
dimensions of water bodies covered by a given group or a Single animal
is a difficult task.

The Murman seals lead a more stationary life. They are distributed in
different regions of a comparatively narrow coastal strip as fairly distinct
isolated local groups living in herds. Such conditions make it wholly
possible to determine the home range areas of the seals in future. Stray
animals are, however, encountered close to Mezensk Bay.

The Baltic gray seals, on the other hand, reveal a varying affinity for
given sections of their habitat. In the winter-spring period, when they
are confined to ice floes, mainly close to the edges, the area occupied
by them often varies in accordance with ice floe drift. Thus the area of
occupancy is sometimes more, sometimes less. Under these conditions,

359

478

the density of disposition of the seals is highly variable not only from
year to year, but also in the relatively short icy season.

During the periods of reproduction or molt the seals on ice floes
generally do not gather into groups; instead they scatter, sometimes at
considerable distances from each other, although not as far apart as in
the case of the larga at this time in the Bering Sea (see p. 353-354). But
sometimes the animals do gather in a small section of the ice floe or even
in groups forming small rookeries. At this time the concentration of gray
seals is maximum. According to recent observations, gestating females
are disposed quite often in small groups of two-five or large groups of
20-40 animals; sometimes they group in even larger numbers or remain
singly (V.A. Zheglov).

At the end of the reproduction and lactation period, the adult seals
that have mated do not evidently go into the water nor do they abandon
the region of whelping; the seals begin to molt. The well-fed pups also
remain on the ice floe for some time more until the embryonic hair coat
has been completely shed. After the disappearance of ice floes in the
summer-autumn season, the seals scatter in a wide expanse and keep
moving in search of food for much of the time.

Hideouts and shelters. The gray seal organizes no special protection
for the pups born even on ice floes or on snowbound land. The newborns
lie in the open and only sometimes, as for example on the islands along
the Murman coast, especially on Great Ainov, enjoy the protection of a
projecting stone or a peat clump (V.D. Kokhanov). Information on the
disposition of pups on ice floes in the Baltic Sea has only begun to be
collected (V.A. Zheglov).

For a long time there was no unanimity of opinion regarding the
ability of the pagophilic gray seals to make air holes in ice floes for
communicating with the water. According to some (Holm, 1921, Aul,
Ling, and Paaver, 1957), the seals do make air holes and keep them
open. One such hole is sometimes used by many animals. Curry-Lindahl’s
(1965) reports do not confirm this view. There is no need for the gray
seal to make air holes as the whelping site in the fringe zone of drifting
ice floes already has innumerable open water pools. Evidently, they set
out to make air holes only when needed, i.e., when the open water pools
or an enlarged fissure is covered by a recently formed ice crust.

Daily activity and behavior. The daily cycle of activity and the behavior
of the Baltic gray seals have not yet been studied thoroughly.

In general, the gray seal is a fairly gregarious animal but its herding
instinct is not equally well developed among the ecologically different
populations; among the pagophilics, especially the Baltic seals, this ten-
dency is weaker than among the land seals. The herding reflex evidently

360

479

peaks by the time of reproduction. However, some British rookeries exist
long before or after the breeding period or there are those in which the
animals do not reproduce at all. The same is noticed among Murman
gray seals, which quite often are bound in herds on their favorite sites
on islands after the breeding season.

The regime of the coastal rookeries also varies. Some exist for a long
time and even almost uninterruptedly round the year, as for example
on the small islets of St. Kilda to the extreme west of the Hebrides
(Waters, 1965). Others, however, function for a much shorter period or
are characterized by almost uninterrupted traffic for the reason that all
the females do not arrive at the same time and remain on land for 2-3
weeks after whelping, their pups for an average of 4-5 weeks (Fitter,
1961), and later leave the rookery altogether. The coastal collections of
animals also vary widely in size: in the British Isles the sizes vary from
small groups of a few tens to huge herds of 2,000 or even up to 5,000 seals.
During the periods of reproduction and feeding, the animals of different
ecological groups are active during the day as well as at night except
for hours of sleep, for which there is probably no fixed time schedule.
These seals can sleep in water as well. Activity and ability of orientation
and finding food at night are characteristic of the Murman gray seals
who have lived long under conditions of the polar night, coinciding with
their period of reproduction. Activity of the British gray seals right up
to their successfully catching salmon in the dark has been confirmed by
observations on the east coast of England in the estuary of Tweed River
(Rae and Shearer, 1965).

During their residence on the coastal rookeries for reproduction in
our Murman region as well as in British waters, the seals do not generally
feed. During lactation the whelped females of the Murman population
remain silent in the water not far from the pups lying on the coast, always
ready to respond to the pup’s call. Less often, they spend the intervals
between sucklings on land near the pups. The nurseries there, as at other
places, break up as the well-fed pups begin to enter the water. A part of
the population abandons the reproductive section while others remain
for a longer period, until spring, and a very small number even up to
summer; at this time, the seals remain in groups in the water. In spring,
coastal rookeries are often formed there again. A characteristic features
of these rookeries is that the seals are disposed in clusters, quite close
to each other (Karpovich, Kokhanov, and Tatarinkova, 1967).

Seasonal migrations and transgressions. The seasonal migrations of
the gray seals do not exhibit similarity in magnitude or regularity with
those of the harp seal, hooded seal, or fur seal. Some remote similarity
of these migrations is seen only with the Baltic gray seal.

480

The magnitude of migrations and the duration of residence in the
sea and the density of animals of different age groups and in different
sections of distribution (with varying extent of food availability) differ
among the pagophilics and land seals. It is known that the young of the
British population remain continuously at sea for the first two years,
often quite far from the site of their birth (Davies, 1956*).

In the winter-spring period, like the harp seal, but on an extremely
small scale, the bulk of the gray seals concentrate in relatively limited
sections of distribution where the ice conditions are most favorable for
reproduction and later even for molt; at the end of this season, however,
they return to the extensive feeding expanses. Thus, on our Baltic coasts,
in the winter-spring season, the gray seal is usually confined mainly to
Irben Strait and the western regions of the Gulf of Riga. In the ice-free
period, however, these seals are more widely scattered.

The main impetus for the spring-winter migrations is the need for
finding suitable ice floes that ensure dependable conditions for raising
the offspring. The seals seek out stable, white, coarsely broken ice floes
that are not intensely hummocky and with a fair amount of open water
pools. The selection of Irben Strait for this purpose is wholly justified
since suitable conditions required for whelping and lactation are gener-
ally available there. The seals find similar conditions also in the central
eastern section of the southern strip of the Gulf of Finland. Regions of
their breeding are also known outside our territorial waters. The seals
begin to move to the breeding zones by late autumn, the younger animals
migrating somewhat earlier (Holm, 1921). In the wintering site, they are
seen in clusters by December, sometimes even in November.

By March end to early April, at the end of the breeding season, the
adult animals leave the suckled pups on the ice to complete molt and
begin to return to their summer habitat. Initially, this process is passive
with the drift of ice floes but later becomes more active.

The Murman gray seal, having lost its biological link with ice floes,
has become a more settled animal. For three-fourths of the year, from
August end, the whole of autumn, winter, and spring months, its popula-
tion, with some minor changes, is confined to the Semi and Ainov islands.
In the former region the seals may be seen even in spring (before June)
in herds of 20-70 animals or more on Litskie, Veshnyak, and Kuvshin
islands and also in some sections of the mainland coast where they form
fairly dense rookeries in spring. Such groups of gray seals are seen at this
time on the Ainov Islands also. But most of the animals depart in June
and only rare strays continue to remain until the end of August. The
summer habitat of the bulk of the seals is not known for certain. They

481

361 begin to return to the sites on the Semi and Ainov islands by autumn,
from the end of August (Karpovich, Kokhanov, and Tatarinkova, 1967).
In the context of seasonal migrations of Murman gray seals, the
corresponding process among their kin, especially the British popula-
tions, is of interest. There are no essential differences in this respect
between many British and our Murman populations. The seals remain
throughout the year in the waters of western Wales, migrations here
being only of local interest (Lockley, 1954). The British populations are
sometimes classified as nonmigratory (Davies, 1956*). However, massive
tagging of pups, commencing in 1954, has shown their great capacity
to migrate and for distant transgressions. Although 85% of a batch of
pups marked on Farne Islands did not leave the area of their birth,
some were caught on the coasts of Holland, the Federal Republic of
Germany, and Denmark, while some reached Norway and the Faeroe
Islands (Hickling and Ennion, 1959*). New data (tagging at the com-
mencement of the 1960s on the Orkney Islands) confirmed the extensive
scattering of juveniles, including even Norway where some twenty tagged
animals were recovered from 1960 through 1962 (Hickling, Rasmussen,
and Smith, 1962). While these do not represent systematic distant migra-
tions, they confirm the great mobility of the juveniles and their ability
to negotiate long distances. This has been convincingly demonstrated by
the fairly extensive actual exchange even between isolated populations,
which appear extremely remote at first glance (see p. 488).

Reproduction. The period of mating among this species extends on
the whole for almost three-fourths of the year, from July end to March
end or early April while some animals mate even somewhat later. Such
a wide mating season is explained partly by the fact that the animals
belonging to the ecologically different groups have their own periods of
mating, partly due to extensive population variations at these periods
among the seals in the British part of the distribution.

The Baltic seals reproducing on ice floes mate in early spring, mostly
in March (predominantly at the end of this month) or in April; it is pos-
sible that mating occurs at a much earlier or even at a later period,
however, in February (Lilljeborg, 1874; Freund, 1933), May (Nehring,
1886), and even in June (Neif, 1757*). It has further been pointed out
that in the more northern Baltic regions, especially in the Gulf of Both-
nia, the seals mate later (in June) than in the Gulf of Finland. According
to the latest data (V.A. Zheglov), the period of mating in Irben Strait
extends from around March 10 to mid-April.

In the Baltic Sea the seals concentrate for reproduction in a fairly
restricted strip of coarsely broken fringes of ice floes, usually tending in
March to south of the Aland Islands, mainly in a latitudinal direction,

362

482

forming а deep loop in the Gulf of Finland. In the southern part. of
the Gulf of Bothnia extensive air holes become available by this time
but whether the gray seals remain in the gulf or reproduce along its
periphery is not known. In general, however, the disposition of the mas-
sive drifting ice floes suitable for whelping is highly variable not only
in a given season, but also in different years. In such situations there
is little justification for assuming fixed dispositions of productive males
and the mother population with pups or the existence of sharply iso-
lated populations. The affirmation that there are no definite places in
the Baltic Sea where these seals can be found for certain in the season
of reproduction or molt is not mere speculation (Smith, 1966*). Never-
theless, judging from the data of recent years (V.A. Zheglov), the stable
regions of reproduction in our waters are Irben Strait as well as the
predominantly western part of the Gulf of Riga. A very small portion of
the population breeds in the central and eastern regions of the southern
part of the Gulf of Finland.

Not much is known about the nature of the mating period. Never-
theless, in spite of statements to the contrary in the literature (Curry-
Lindahl, 1965), harems are not a characteristic of the Baltic seal. The
seals mate two weeks after whelping. Copulation occurs in water as well
as оп ice floes but more often evidently in water. Among the British gray
seals, there is a definite relation between the nature of coitus and the
extent of manifestation of the harem regime in the rookery. On land,
these seals mate at places where the males hold firmly to their territorial
sections, as for example in the largest rookeries on the northern Rona
and Farne Islands. But even under these conditions, when fairly deep
erosion “tanks” or “channels” are available, the animals eagerly take
advantage of them for coitus. When, however, the males are confined
preferentially to the sea in front of the whelping site, mating proceeds in
water (Hewer, 1960). Under the prevailing harem regime of the rookery,
the males holding onto a definite territory fertilize many females. Evi-
dently, in water too, the more powerful males stand in an advantageous
position by chasing away the much younger and weaker animals.

This aspect of the life of Murman gray seals has not been clearly
understood. In the Norwegian waters in the remote past, when the seal
population was much larger, there were 2 to 4 or 5 females to every bull
(Collett, 1881). Murman gray seals mate late in the autumn and early
winter, 2 to 3 weeks after whelping or slightly later. This period essen-
tially falls at the end of November to the end of December. Any overture
to mating before this period (long before completion of lactation of the
female) or male proximity to her pup is decisively resisted by the suck-
ling female, even to the extent of biting the neck of the intruder. Mating

363

483

is completed mainly on land. In any case, mating in water could not be
observed. Coitus is preceded by conjugal play in water or on the coast
when the animals gently bite each other. Signs of distinct harem forma-
tion were not seen among the Murman population nor were there more
females than males among the adult animals. It is nevertheless possible
that both monogamy and polygamy prevail on Murman. Polygamy was
suggested by the disposition of pups in groups of 2-4 together (Kar-
povich, Kokhanov, and Tatarinkova, 1967).

Conforming to the extended period for mating, the whelping season
too reveals wide variations geographically as well as among the various
seal groups in the same region. Thus, in the Great Britain-Ireland region,
some main groups in the population have a much extended whelping
period while others have a short whelping period. In Wales, the Irish Sea,
and on the western coast of Ireland, whelping extends from July through
October; among Scottish-Hebrides populations, pups are encountered
in September-October and on the Farne Islands (northeastern coast of
England) from the second half of October to the middle of December
(Coulson and Hickling, 1964). However, the bulk of the females whelp
within a very short period of about a month. The view that the eastern
Atlantic seals whelp increasingly later toward the south (Mohr, 1952)
has not been fully substantiated.

The Murman seals are distinguished by a somewhat greater stabil-
ity in this respect: the period of reproduction among them extends for
not more than 1.5 months in the late autumn. Even according to older
information (Prigorovskii and Breitfus, 1912, 1915), newborn pups of
gray seals were caught on Murman throughout December but, according
to much later information, they were seen on the Semi Islands in early
November (S.M. Uspenskii, 1941), from the end of November to the
end of December (Sdobnikov, 1933*); they were caught on November
16 and 17, December 6 (Belopol’skii, 1941) or in November and Decem-
ber (Belopol’skii, 1951). The above whelping periods in eastern as well
as western Murman have been confirmed by the latest observations; they
are identical and the total duration of whelping extends for 30 to 40 days.
The earliest pups are born in the first ten days of November and the last
ones toward mid-December. In 1963, on Great Ainov Island, two peaks
were noticed in the birth of pups, one from November 10 to 15 and the
second, a sharper peak, at November end to early December (Karpovich,
Kokhanov and Tatarinkova, 1967).

Though neither the whelping nor mating season of the Baltic seal as
reported in the literature is very accurate, there are no particular dispar-
ities regarding these aspects. In most cases the pups appear by February
to March, the main whelping season falling in March (Lilljeborg, 1874;

484

Nehring, 1886; Aul, Ling, and Раауег, 1957) ог February end to early
March (Holm, 1921) or between February end and mid-April (Smith,
1966*). Recent observations have shown that the total whelping period
among the Baltic gray seals extends for one month from around Febru-
ary 20; most of the pups are born from February end to March 4-5
(V.A. Zheglov).

The gestation period has been put variously at 8.5 to 12 months
because of the lack of accurately established mating and whelping peri-
ods. In the middle of this century, it was assumed as 11 months, as in
the case of other seals (Mohr, 1952). Obviously, however, the entire
period from mating to parturition stretches two weeks longer, to 11.5
months. But considering that a prolonged delay of zygote implantation is
characteristic of gray seals (Lockley, 1954; Backhouse and Hewer, 1956),
the actual embryogeny extends for roughly nine months or so.

The birth process is extremely brief, being accomplished in a few
seconds; the placenta is discarded in an hour and many animals eat it
(Matthews, 1952). Twins are very rare among gray seals (Collett, 1881;
Curry-Lindahl, 1965) although according to some authors (Аш, Ling,
and Paaver, 1957), Baltic seals do produce twins.

Births in the normal course are an annual feature. The overall bar-
renness has not been established accurately for any of the populations
of these seals though the quantum of females not participating in repro-
duction, on analogy with fur seals, was put at 20% among the British
population (Hewer, 1964).

The British female gray seals become capable of reproducing partly
(50%) at five years of age and wholly (100%) at six years. By this time,
depending on the state of ovogenesis and ovulation, the length of the
ovaries increases on average by 32-35 mm and the weight goes up to
5-8 g. Thus the female births her first pup at six years of age. The males
become mature at seven years of age but stake their claim at the site of
reproduction only on attaining 10 years; they cease mating at 20 years of
age. Among mature males, in the mating season, the weight of the testes
together with the spermatic cord varies from 100 to 290 g, on average
166 g. The weight decreases by the end of the mating season roughly to
100 g; the length of the os penis is not less than 10 cm.

The females remain productive for a very long time; although rare,
gestating females older than 30 years (31-32 years) have been recorded.
The oldest female encountered in a breeding site was 33-34 years
(Hewer, 1964).

Similar data are still not available for Murman gray seals. The oldest
suckling Baltic seal seen in our waters was aged 21 years (V.A. Zheglov).

364

365

485

Growth and development'**. There is very little concrete information
on the size of newborn pups of Baltic seals in the earlier works. The
following figures are cited: 95 cm (Lilljeborg, 1874) and 60 cm (Priemel,
1909). The first figure is wholly reliable (evidently, the pup was measured
along the dorsal surface) while the latter figure is clearly an undermea-
surement. According to the current data (V. A. Zheglov), the body length
in a Straight line (i.e., Lcv) varies from 75 to 85 cm and along the dorsal
surface (Lc) from 82 to 92 cm; weight ranges from 6.1 to 9.5 kg, on
average 8.15 kg. The length of the suckling pup averages 100-110 cm
or 110-120 cm (Lc) and its weight 40-50 kg (the lower limit pertains
to females and the upper limit to males). The body length of a newborn
Murman female gray seal in a straight line (Lcv) is 107 cm, axillary girth
64 cm, thickness of the skin with the subcutaneous fat layer (51) 0.5 cm;
total weight 15.2 kg (Uspenskii, 1941). A slightly older male pup, roughly
three days old, was 110 cm long (Lcv) with an axillary girth of 66 cm,
thickness of skin (St) 0.9 cm, and weight 20.5 kg (Fig. 193). Pups born
with a body length (evidently, Lcv) of 80-107 cm add 15-25 cm in the
period of suckling, the most intense growth occurring in the first two
weeks. After 5-7 days the canines have cut through and the rest of the
permanent teeth are cut immediately afterward (Karpovich, Kokhanov,
and Tatarinkova, 1967).

The newborn British seals weigh 11.5-17.5 kg but most 14-16 kg,
with a body length of 91-108 cm; the body weight increases during the
period of lactation to 40-50 kg (Matthews, 1952; Coulson and Hickling;
1964). The duration of lactation has been recorded by various authors as
two weeks (Darling, 1948) and four weeks (Lilljeborg, 1874). Evidently
suckling most often ceases in the middle of the third week (captive pups
were weaned on the 18th day, and on Farne Islands, England on the
16th day) (Coulson and Hickling, 1964). During the first three days of
lactation the daily weight increment averages 1.5 to 1.8 kg. This rapid
increment in weight is ensured by the high fat (52% or more) and protein
(11%) contents of the mother’s milk (Amoroso and Matthews, 1951).

The frequency of suckling (Fig. 194) changes with time. The newborn
receives its first nourishment 30 min after birth and later at 2-hr inter-
vals; in the latter days this interval increases and the number of feeds
decreases to a few times a day (Smith, 1962). Among the Baltic seals, the

124 The data available in the literature on the size of the newborn, its growth during the
period of lactation, and in the subsequent period of its independent living, at least in the
first year, as well as the sizes of adults are difficult to utilize since it is usually not known
how the animals were measured: along the dorsal curvature or in a straight line to the tip
of the tail, or to the end of the extended hind flippers.

364

486

Fig. 193. Newborn pup of Murman gray seal in the first few days after birth. Great
Ainov Island, Murman coast, December, 1963 (photograph by V.D. Kokhanov).

pups suckle initially at 3 - 4 hour intervals, receiving 400 - 650 р or more of
milk in each feed. The daily weight increment varies from 1.2 to 2.5 kg but
most pups double their weight in 8- 10 days and reach 33 - 40 kg or more
by the end of the lactation period (V.A. Zheglov). The suckling dura-
tion increases as the pup grows and the milk intake likewise increases.
According to observations on Great Ainov Island (western Murman),
in the second half of the lactation period suckling extends for almost
15 min. The pups suckle the two teats alternately, taking 3-4 mouthfuls
at a time from each (Karpovich, Kokhanov, and Tatarinkova, 1967).
The behavior of pups of the Murman population in the period of
suckling has been described as follows: “The newborn pup moves little
and permits fondling with the hand with no resistance. Just a few days
later, it becomes active and attempts to shy away from man or protect
itself, sometimes by trying to attack”. The pups usually remain within
the coastal belt, up to 100 m from the line of high tide. Sometimes,
however, they do stray up to 300-400 m beyond the coast. The pups
spend much of the first two weeks sleeping, mostly on the right side or
on the back. With the commencement of intense molting, they become
more active, mainly on the rocks, often descending into the pools in the
sublittoral and littoral zones. Some pups still not completely molted go
into the sea (Karpovich, Kokhanov, and Tatarinkova, 1967). Completely
molted pups, however, continue to remain on land for a few more days,

366

367

365

487

often moving away for long distances from the place of birth, or migrate
from one island to another in the vicinity (Karpovich, Kokhanov, and
Tatarinkova, 1967).

The pup retains its white embryonic coat for the same short duration
on the ice in the Baltic Sea as well as on the Murman islands, Norway or
Great Britain. This coat loses strength in seven to ten days (Neif, 1957*;
Collett, 1881; and others); among Baltic Sea newborn males and females,
this happens in 5 -6 and 8 - 10 days respectively. In 17 - 20 days hair begins
to appear again on the trunk (V.A. Zheglov). Among Murman gray seal
juveniles too molting is over by the end of the first week. The first molt
follows a fairly common pattern for all the animals of the family. Weak-
ening and shedding of the embryonic hair coat usually occurs first on the
head, the hind and fore flippers, and the tail (Fig. 195). On the Sth- 8th
day the embryonic coat thins out on the snout and on the flippers; dark
spots show through the embryonic coat here and there on the back. On
the 9th- 12th day dark spots are seen all over the body, which fuse into
a continuous gray field, signifying the commencement of intense molting
(Fig. 196). By the 13th - 16th day the head, back, and sections around the
flippers have completely molted and the embryonic coat elsewhere on
the body scales off intensely. On the 17th-22nd day hair remnants are
seen only on the flanks and on the abdomen but these too are weak; on

Fig. 194. Posture of gray seal during suckling. Great Ainov Island, Murman coast,
December, 1963 (photograph by V.D. Kokhanov).

488

the 23г4 - 24th day the remnants are rare; on the 25th- 27th day there is
nothing left of the embryonic coat and the pups appear in the new coat
with smooth short hair (Karpovich, Kokhanov, and Tatarinkova, 1967)
(Fig. 197). Thus the entire molting process in Murman takes roughly
three weeks. Little is known in general about the further growth of the
juvenile. There is altogether no information on its life and growth tempo
in our territorial waters. On taking to independent living in water, the
pups are usually scattered widely, quite often straying very far from the
place of their birth (see p. 481). The subsequent growth of the juvenile
has been studied only among the British populations; by the end of the
first year, the males begin to outstrip the females in growth (Table 26).

The yearlings are practically no different from the well-fed pups (at
the time of first molt of their embryonic coat) and are even lighter in

366 Fig. 195. Some growing gray seal pups; commencement of molt. Great Ainov
Island, Murman coast, December, 1963 (photograph by V.D. Kokhanov).

367 Table 26. Average body weight (after Hewer, 1964)
Age, Average body weight (approximate), kg
in years

Female Male

1 41 42
2 49 59
3 55 74
4 65 90
5) 72 105

366 Fig. 196. Molting gray seal pup. Great Ainov Island, Murman coast, December,
1963 (photograph by V.D. Kokhanov).

367 Fig. 197. Molted gray seal pup. Great Ainov Island, Murman coast, December
(photograph by V.D. Kokhanov).

weight than the latter. This phenomenon (Coulson, 1959*; Coulson and
Hickling, 1964) is explained by the characteristics of the further growth
conditions of a pup in the first few months of its independent existence.
While still on the ice floes, after lactation ceases, the pup suffers a weight
loss which continues for sometime even in the water as the youngster

368

490

has to acclimatize and become accustomed to finding its own food. This
transitional period is evidently a vital factor in its life (as among other
seals) and the pup is often forced to merely survive by expending the
reserves accumulated during lactation. The pup can replenish its body
losses only after total adaptation to the water and taking to intensive
feeding.

The growth of the animals, judging from the growth curve of the
British seals (Hewer, 1964), continues with gradual attenuation until
roughly 10 years among females and up to 15 years among males, or
perhaps somewhat later. The average difference in body length between
males and females after total growth cessation is roughly 35 cm. Growth
is most intense among females for 6-7 years and among males for 8-9
years. Increase in skull length proceeds roughly in this same range.

The known longevity in nature is 35 years for females and 25 years for
males (Hewer, 1964). In zoological gardens some females have survived
for 28 and males 41 years (Matheson, 1950; Mohr, 1952).

Information on the molting of adult Baltic seals is very scant. At the
end of the period of reproduction, after some nourishment, the Baltic
seals return to ice floes and form fairly large molting rookeries on them
(Holm, 1921). According to the earlier data (Neif, 1757*), this process
commences by March 25; the present information (Curry-Lindahl, 1965)
puts the molting of seals in April-May. There is no information at all
on the molting of Murman seals.

In British waters the molting period proceeds 1.5 to 2 months after
the mating period; females molt much earlier (mainly at January end to
early February) than males (mainly in March) (Backhouse, 1960; Smith,
1962).

Enemies, diseases, parasites, mortality, and competitors. These aspects
have been little studied in our waters. In the Baltic Sea the eagle may be
an enemy of the pups (Aul, Ling, and Paaver, 1957) but these birds are
very rare in the pelagic regions of the sea. The pups are probably threat-
ened by large gulls, especially the great black-backed gull (Larus marinus
L.), which pecks at the placenta and the newborn (V.A. Zheglov). Evi-
dently the pups also suffer from unfavorable ice conditions. Instances of
early thawing or hummocking of ice floes can pose mortal consequences
for the pups which are yet weak and have not had sufficient time to
accumulate the subcutaneous fat reserves requisite for independent sur-
vival in water. Ravens pose some danger to the newborn on Murman
(Uspenskii, 1941). Recent observations on the breeding sites of the gray
seal on Murman have shown that the losses among pups are very few.
On Great Ainov Island in 1963, of 93 pups, only two were found dead.
Two dead pups were found in the winter of 1960/61 on Veshnyak Island

369

491

(Karpovich, Kokhanov, and Tatarinkova, 1967). In the British Isles pups
born on the rocks in high-tide areas perish almost wholly; on northern
Rona Island, in two weeks of October, pup mortality was 16% (Boyd,
Lockie, and Hewer, 1962*). The mortality among newborns in 1957 on
the Farne Islands was in the same range (England; Hickling, 1959*).

In calculations of the age-sex structure of the British populations,
mortality in the first year was put at 60% of the total population of pups,
30% in the second year, and 12% in the third year; it was 6.7% in each
of the subsequent years (Hewer, 1964).

The helminth fauna of gray seals inhabiting the USSR waters has
yet to be adequately studied. Gray seals are infected by eight species
of helminths, among which not a single cestode appears (Delyamure,
1955); among the trematodes, Metorchis albidus, Opistorchis tenuicollis,
and Pseudamphistomum truncatum infect the gall bladder and bile ducts;
among the nematodes, Anisakis similis infects the intestiae, and Contra-
caecum osculatum and Terranova decipiens infect the stomach and the
intestine; among the acanthocephalans, Corynosoma strumosum and С.
semerme infect the intestine.

The gray seal has no competitor in the Baltic Sea. This is the largest
of the marine life normally inhabiting this water body and has its own
ecological niche, preferentially confined to the more pelagic regions, and
feeding at depth. The common seal in the southern part of the Baltic is
much smaller than the gray seal. The distribution of both these species
in Murman represents the boundary of the range and both evidently live
under different biotopic conditions.

Population dynamics. In the southern parts of the Gulf of Finland the
population is insignificant and apparently is continuing to decline. The
total population of the gray seal in the Baltic Sea too continues to drop
with no remission, especially in the gulfs of Finland and Bothnia. Thus,
the average annual catch of both species of seals in Finland compared
with the pre-Revolution period, dropped in the second half of the 1920s
roughly by 30% and in the early half of the 1930s by as much as 60%.
The situation was similar in Swedish waters. The gray seal in the Baltic
Sea is largely unprotected from foreign hunters. Encouraged by mone-
tary rewards, they kill it wherever and whenever convenient. Apart from
the fact that the seal is intensely hunted in general, significant qualitative
changes have occurred in the structure of its population due predomi-
nantly to killing of pups in the past as well as now, as they are quite
easy to catch (Bergman, 1956). Hence the herd is relatively overloaded
with older animals (Curry-Lindahl, 1965) whose natural death cannot be
replenished. That the Baltic seal population, which showed some recov-
ery after the war years, continues to fall is supported by the magnitude of

369

492

the continuously declining kill in spite of the alluring monetary rewards
granted by the Swedish and Finnish governments for every animal killed
(Table 27).

Apprehensive of the extermination of the seals, Swedish organiza-
tions for environmental conservation have turned to the government with
proposals for the abolition of rewards for each killed animal, a ban on
hunting during the breeding period, and the establishment of a number
of sanctuaries (Curry-Lindahl, 1965).

The small population of Murman gray seals underwent changes in
the current century in relation to hunting activity. In the pre-Revolution
period and the initial post-Revolution years, the population was evidently
at maximum. In the 1930s, as a result of excessive killing of pups, the
population fell but was again restored after the years of the Patriotic
War. The establishment of Kandalaksh sanctuary played a positive role
in sustaining and raising the population. At present, “the population of
gray seals in the sanctuary has restored to the levels prevailing in the
1920s” (Karpovich, Kokhanov, and Tatarinkova, 1967).

Field characteristics. The gray seal can be distinguished from other
seals by the straight (upper) profile of the head, long snout, and light
coloration of the upper portion of the head (Fig. 198). The last feature
unmistakably distinguishes from a distance a swimming gray seal from
the common seal whose head is invariably dark (Millais, 1904; Collett,
1911-1912). In the Baltic Sea the gray seal can be recognized from a dis-
tance by its large size, elongated snout, and coloration, i.e., innumerable
dark spots scattered haphazardly on a much lighter gray background; a
gray seal protruding above the water can be recognized from the con-
trasting color of the throat, neck, and chest with a pattern of larger spots
seen prominently on a light background. Murman gray seals can also be
identified by these features as well as by the dark, almost monochromatic
color of the males. (K.Ch.)

Table 27. Dynamics of catch during 1943 to 1953 (Lockley, 1954)
Years Catch
Average per year Maximum and minimum
(number of animals) per year
1943-1945 940 1,345; 625
1946-1949 710 783; 615
1950-1952 528 707; 397

1953-1954 246 290; 203

370

370

493

Е

Fig. 198. Semiadult gray seal. Northern spits in the White Sea, September, 1970
(photograph by A.G. Beloborodov).

Economic Importance

The gray seal commands little economic importance in our animal hunt-
ing activities. Its reserves in our waters are insignificant and hence there
are no special hunting prospects here. Before the Patriotic War, its kill in
Murman did not exceed several dozens (mainly young animals) per year.
To increase the population to a level capable of supporting profitable
hunting, hunting should be banned everywhere for 10 years and then a
thorough study of its ecology done, especially of its feeding habits, to
determine whether this seal damages the salmon population.!” If such
damage should prove negligible, the gray seal could become a regular
source of small numbers of valuable fur. The other ancillary products
that could be utilized are the fat (a weaned juvenile can yield about
20 kg of fat and an adult 50 kg or more) and edible meat for animal
farms (10 to 12 kg from a young animal and roughly 50 kg or more from
an adult seal).

In the Gulf of Finland in the Leningrad region there is neither spe-
cial hunting of this seal nor is it pursued as a sport. On the Estonian
coasts in the prewar years, seals were caught fairly regularly but the gray

125 The available general references indicate that in Murman this seal causes no significant
damage to the fishing industry.

371

371

494

seal was not а frequent catch. Outside ше USSR, in the Baltic Sea, the
gray seal was caught mainly in the winter-spring in the region of the
Aland Islands and the adjoining expanses in the zone of ice fringes and
drifting ice floes, in the Baltic Sea itself, the gulfs of Finland and Bothnia,
and in smalier numbers, On occasion, in summer and autumn at various
other places. The total catch by Sweden and Finland (encouraged by
monetary rewards) in the 1940s and 1950s reached an average of 1,000
or slightly more per annum (Table 28).

Outside the Baltic Sea, the killing of these seals is rewarded on the
Faeroe Islands (some 800 animals including 500 pups were killed in two-
and-a-half years) and in Canada. Great Britain has imposed a prolonged
seasonal ban during the breeding and suckling periods.

Hunting in the Baltic Sea is mainly undertaken by fishing vessels cruis-
ing in the period of whelping and molting of seals along the fringes and
among fairly thin ice floes. Most of the catch there at this time is of the
young; quite a few suckling mothers are also caught. Sweden and Finland
have no rules whatsoever pertaining to sealing, nor are there any agree-
ments between them regarding it. In the summer-autumn months stray
seals close to the coast are killed quite often from launches and boats.

Lack of supervision and totally unrestrained killing of the Baltic gray
seal population has arisen because of the commonly held belief that this
animal is a plunderer of fish. Without doubt, the gray seal does consume
fish. But the fish consumed by it is nowhere, and never has been, of much
economic importance and its rapacity is not significantly reflected in the
fish reserves and hardly so in the catch. No perceptible damage caused
by this seal has been recorded either in Murman or the Gulf of Finland.
It has been said that it does not generally damage the fishing industry in
the Baltic and North Seas (Freund, 1933) although it was earlier blamed
as the most dangerous of seals in the Baltic Sea (lort and Knipovich,
1907* ). Furthermore, it has played no perceptible role as a destroyer of
economic fish in some Canadian regions (Mansfield, 1966) despite an
increase in its population through controls imposed along the lines of
experience gained in Great Britain.

Table 28. Number of gray seals caught in the Baltic Sea from 1930-1959
(Curry-Lindahl, 1965)
Years Caught in Total
Finland Sweden
Adults Pups Total
1930 - 1939 8,995 17,265 26,260 11,952 38,212
1940 - 1949 3,800 2.715 6,515 6,792 13,307

1950-1959 3,743 2,743 6,465 3,597 10,062

372

495

Nevertheless, the presence of gray seals is undesirable in the larger
economic fishing areas, especially in the estuarine sections of seas
through which salmon pass into rivers to spawning sites and back.
Here their adverse role on the fishing industry is obvious and controls
necessitated. The damage caused by the seals to man is not restricted
to the consumption of economic fish, i.e., to the adverse effect on fish
reserves, but is also manifest in their consumption or partial damage of
fish caught in nets, especially salmon, and the damage they do to the
nets per Se.

In the coastal waters of England and Scotland the damage caused
by gray seals is quite significant at places. According to rough but well-
argued calculations, gray seals (about 47 head) in 1957 could have con-
sumed roughly 10 tons of true salmon and sea trout in the estuary of the
Thames River during the entire fishing season covering 182 days. The
damage extending to other fish (cod) as well as squids would account
for about 3% of the total catch of salmon (the calculations assumed the
following daily ration: adult seals 6.8 kg; immature seals aged two to
five years 5.5 kg; and yearlings 2.3 kg) (Lockie, 1962). The total dam-
age inflicted by the seals to the fishing industry on the Scottish coasts
has been put at 67,000 pound sterling per annum (Rae and Shearer,
1965). Fishermen of other countries too (Sweden and the Faeroe Islands)
deplore these salmon losses. Nevertheless, there is no unanimous view on
this subject. The quantum of damage in some other parts of its range is
insignificant (in Norway and along the coasts of the two German states)
and the gray seal is a protected animal there. (K.Ch.)

Subfamily of Monk Seals or 8-incisored Seals

Subfamily MONACHINAE Trouessart, 1904125

These seals are large in size; their body length with tail along the dorsal
surface is not less than 200 cm.

The hind flippers have a fairly deep median notch. The first digit
on the fore flippers is the longest and the rest gradually decrease in
size toward the fifth; the claws are well developed on the fore flippers
but those on the hind flippers are highly reduced and sometimes almost
altogether invisible. The nostrils are mainly disposed on the dorsal side

126 The present structure of the subfamily was established by Gray (1850) long before
Trouessart, but Gray named the subfamily Stenorhynchinae from Stenorhynchus, already
used in the systematics of marine invertebrates. (K.Ch.)

The year 1897 is perhaps more correct as this was the year in which volume I of the
Catalogus was published. (V.H.)

496

of the snout. A proboscis on the anterodorsal portion of the snout is
lacking. The whiskers are fairly flattened, or smooth, or with slightly
wavy edges.

There are two pairs of teats.

The skull is essentially massive, with thick bones, and there is usually
a broad gap between the orbits. The maxillaries anterior to the orbits
are not swollen (not convex). The zygomatic arches project laterally to
different degrees. The preorbital processes are prominent in most cases.
The external opening of the bony auditory meatus is simple, without
prominent lobes. The nasal aperture is less open upward and its upper
posterior edge does not extend beyond the anterior margin of the orbits
at the back; in most cases, in fact, the nasal aperture does not reach the
orbits (genus Ommatophoca is an exception). The intermaxillary pro-
cesses are usually contiguous with the nasal or almost so.

There are two incisors on each side in the upper and lower jaws.
The dental formula is:

мы. 1
т, Ст, Pa Ms —32.

The cheek teeth, except for the first premolar, have two roots.

There are no sharp differences in the size and body build and partly
in the coloration between males and females. In general, the body build
is similar to that of seals of the subfamily Phocinae. The hair coat among
newborns is dark toned.

Biological affinity for a hard substratum differs: the seals of one
genus (Monachus) form rookeries and reproduce on land and the rest of
the genera on ice floes.

The distribution of the subfamily is broken into different sections
(Fig. 199). These seals are partly distributed in the Northern hemisphere:
one genus (Monachus) occurs in the subtropical and tropical belt of the
Atlantic and Pacific oceans and the rest in subantarctic and antarctic
waters, mainly in the zone of drifting ice floes.

Monachinae undoubtedly represent a product of the further devel-
opment of Phocidae. They deviated from the branch of the 10-incisored
seals (Phocinae), lost one each of the upper incisors, and acquired the
features of high adaptation to pelagic living. This is manifest particularly
in the reduction of claws on the hind flippers, in the llrge notch on the
posterior margin of the latter, in the much sharper reduction in length
of the digits of the wrist from the first to the fifth, etc.

The evolution of the subfamily has been traced to the Miocene of
western and southeastern Europe by which time the members of this
phyletic branch had not only fully evolved, but differed in some respects
by an even greater specialization (Pontophoca). It is highly probable that

497

‘(uysdeyD yy) эвищовецой ‘$[еэ$ pasosioul-g jo Апшезап$ ay) JO UONNNGUISIG ‘66т ‘9 ELE

SSS та
ary 0056 0007 0008 0005 60% 0 0007

498

parts of the Tethys or its tributaries served as the birth place of the
primary forms of the subfamily.

The subfamily has not yet been conclusively systematized. Disagree-
ments exist regarding its volume and constituent genera. The old view
that the subfamily Monachinae includes only one genus, Monachus, is
still quite prevalent (Simpson, 1945; and others); the rest of the (antarc-
tic) 8-incisored seals constitute a special subfamily, Lobodontinae Hay,
1930 (= Lobodoninae Kellogg, 1922).

However, it would seem more correct to adopt the structure of the
subfamily to include a large volume in the form of a single group com-
bining all the 8-incisored seals (Chapskii, 1961, 1969). At the same time,
differentiation of Monachinae at a much lower level, at the rank of tribes
Monachini and Lobodontini (Scheffer, 1948*), is entirely reasonable
and rational (Chapskii, 1971*). But the spelling should be Lobodon-
inae.

Insofar as the composition of the genus is concerned, there is every
justification (Chapskii, 1969) that it should be left alone in the same
manner as adopted at the beginning of this century, 1.е., not include
the genus Mirounga in spite of a recent suggestion to this effect (King,
1966). The subfamily would then contain nine genera (about 41% of
all the genera of the family), of which four are extinct and five extant
(50% of the contemporary genera of the family): monk seals or white-
bellied seals, Monachus Flemming, and the four monotypical genera of
seals of the Southern hemisphere, i.e., crab-eater seal Lobodon Gray;
Weddell’s seal, Leptonychotes Gill; Ross’ seal, Ommatophoca Gray, and
leopard seal, Hydrurga Gistel, 1848. The total number of species is seven,
which constitutes about 39% of the total number of species of the
family.'’

The species of the subfamily have no economic importance anywhere
in the Northern hemisphere because of their negligible population. Until
the end of the 1960s, these seals were practically untouched in antarctic
waters. The population of some species there, especially of the crab-
eater seal, is quite large and the idea of their commercial exploitation is
gaining ground.

There is only one genus within the USSR, the monk seal, Monachus
Flemming, 1822 (20% of the genera of the subfamily), with a single,
practically extinct species (roughly 15% of the species of the subfamily).

_ Encountered in the USSR waters as a very rare, straying single ani-
mal. (K.Ch.)

127 According to the system adopted in this publication, the family consists of 18 species.

Si).

499

Genus of Monk Seals

Genus Monachus Flemming, 1822

1822. Monachus. Flemming. Philosophy of Zoology, 11, p. 187. Phoca
monachus Hermann.

1824. Pelagios'** Е. Cuvier. Mem. Mus. Hist. Nat., 11, р. 196. Phoca
monachus Hermann.

1841. Pelagocyon. Gloger. Gemeinn. Naturgesch., vol. 1, no. 34, p. 163.
Pelagocyon monachus = Phoca monachus Hermann.

1848. Rigoon. Gistel. Nat. Thierr. flr hoh. Schulen, Х, р. 32. Renamed
Pelagios Cuv.

1854. Heliophoca. Gray. Ann. Mag. Nat. Hist., 13, p. 201. Heliophoca
atlantica Gray = Phoca monachus Hermann. (V.H.)

Seals of large dimensions. Body length from tip of snout to tip of
tail 240-295 cm.

Similar in body proportions to the species of the subfamily Monachi-
nae. There are no sharp external differences between males and females.

The skull is quite massive; the much older animals have fairly
intensely projecting zygomatic arches. The sagittal crest is quite distinct
though not high; the occipital crests are massive and sharply displaced
posteriorly on both sides of the sagittal crest along the linea nuchalis
terminalis. The squamose-mastoid processes are very well developed;
the posterior margin of the mastoid itself projects slightly posterior
to these processes when viewed upward. The nasal processes of the
premaxillary bones adjoin the nasal bones. The facial portion (apex)
of the nasals does not exceed one-half their total length. The anterior
margin of their suture is bisected without forming a median angular
projection. The petrosal bone projects into the foramen lacerum
posterius.

The upper incisors posteriorly have a prominent transverse groove
merging into a frill.

The hair coat is not high but has an extremely prominent nape; the
hairs adhere very closely to the skin.

The color of the upper side of the body is a dark gray, blackish-
brown, turning very light on the underside. Some sex- and age-related
differences exist. The skin around the nostrils and in the gap between
them is covered with hair. The whiskers are smooth and oval in cross
section. The neonatal coat is dense, quite high and smooth, and very dark.
The claws on the fore flippers are well developed but highly reduced on

128 Various authors have spelled it differently, viz., Pelagius, Pelagus, or Pelagias.

Tif

500

the hind flippers; nonetheless, their narrow and short tips are visible on
the latter.

The seals of this genus are thermophilic and, among all the pin-
nipeds, are the most capable of surviving in subtropical and tropical
conditions. They are biologically associated with the coast on which they
whelp, suckle the pups, molt, and form rookeries, their size depending
on the total population and the nature of the coast. They do not form
harems nor do they undertake any significant or regular migrations, being
predominantly confined to the coastal zone.

The distribution of the genus is highly interrupted (Fig. 200).
Its species are distributed in three isolated regions of the subtropics
and tropics of the Northern hemisphere. One is the basin of the
Mediterranean Sea (up to the Black Sea inclusive) with the adjoining
sections of the eastern Atlantic, in the southwest from Gibralter to the
Canary Islands and the coasts of northwestern Africa. Another is the
now practically uninhabited westernmost part of the Atlantic: the Gulf
of Mexico and the Caribbean Sea. The third region of distribution is
even more isolated, lying in the Pacific Ocean and covering only the
archipelago of the Hawaiian Islands.

This genus has evolved through intermediate links from Miocene
ancestors found in western and southeastern Europe, including the Sar-
matsk formations in USSR territory. A likely direct ancestor of the genus
Monachus could be some Lower Pliocene form of the genus Pristiphoca
since a member of the recent genus Monachus adjoined the above fossil
genus even in the Middle Pliocene. The geological history of this genus
binds it to the west of the Old World, especially to the western, south-
ern, and southeastern parts of contemporary Europe, to the basin of the
Tertiary Tethys Sea.

Paleontological proof and morphological data of the contemporary
members of the genus Monachus indicate that this genus represents a
phyletic base from which other contemporary genetic branches of the
subfamily originated and evolved. The genus Monachus, more than any
other genus of Monachinae, reveals many features relating it to the sub-
family Phocinae (especially the petrosal bone projecting into the foramen
lacerum, margin of mastoid visible when seen upward, structure of the
zygomatic bones, etc.). It may be regarded as a primary element of the
eight-incisored phyletic branch originating from the primary Phocinae
and serving in turn as a stage for the evolution of its derivative, i.e., the
primary ancestral form of subfamily Cystophorinae.

The genus consists of three species, or about 43% of the species
of the subfamily: (1) Mediterranean monk seal, M. monachus Hermann,

‘IPUDISUINDYIS “PFW чепемеН— SNYIDUOLU W uesuelss}Ipsf[—Z
‘syooidoa "И чеэаанеэ—1 ‘(irysdeyo МУ) 5пузриои ‘seas Уцош jo snuas 94} JO UOTNQIISIP PsJONISUODIY ‘005 “B14 9/5

aaa
03000$ 020+ OCDE 0505 GO O 0501

502

1779; (2) Caribbean monk seal, М. tropicalis Gray, 1850; and (3) Hawai-
ian monk seal, M. schauinslandi Matschie, 1905. Though all of these are
allopatric, the species characteristics are independent. They differ mor-
phologically in the structure and position of the teeth in the row, in the
form of nasal and zygomatic bones, and in other respects. Thus there are
two Atlantic and one Pacific species. At present, practically speaking,
there are two species (both the species are protected in most places);
the West Indian species is apparently wholly extinct.

The fauna of the USSR contains one species, M. monachus Her-
mann, 1779, or 33% of the species of the genus and roughly 0.3% of the
species in the fauna of the USSR.

This species was almost extinct in the territory of the USSR even by
the second half of the last century. (K.Ch.)

MONK SEAL

Monachus monachus (Hermann, 1779)

1779. Phoca monachus. Hermann. Beschaf. Berlin. Ges. Naturf. Freunde,
4, p. 501, pl. 12, 13. Lake and coast of Dalmatia; Adriatic Sea.

1785. Phoca albiventer. Boddaert. Elench. Anim. 1, p. 170. Adriatic Sea.

1800. Phoca bicolor. Shaw. General Zoology, 1, p. 254. Adriatic Sea.

1816. Phoca leucogaster. Peron and Lesuer. Voyage aux terres Austr. 2,
р. 47. Nimes, southern France, Mediterranean Sea.

1828. Phoca hermanni. Lesson. Dict. Class. d’Hist. Nat., 13, p. 416. Adri-
atic Sea.

1838. Monachus mediterraneus. Nilsson. К. Svens. Vet. Ak. Handl., 1837,
p. 238. Adriatic Islands and Greek archipelago.

1848. Phoca crinita. Menis. П Mare Adriatica, р. 153. (V.H.)

Diagnosis

The dimensions are large, evidently larger than many other species of the
genus. The color is darker. The nasal bones are relatively short and form
16-20% of the condylobasal length. Their anterior margin has a median
notch. The upper incisors are disposed in an arc; there is no more than
one additional cusp on the premolars posterior to the main cusp. The
preorbital processes are well developed. The uncinate processes of the
pterygoid bones are not bent angularly outward. (K.Ch.)

Description

In general build, the monk seal does not differ greatly from other seals
378 Of similar size (Fig. 201) but is evidently slightly stockier than many

503

ZZ

2..2) —
EGA LEE
РА РНР ее а.
SY LEE = eas ГАР а
Sate, ЗЕЕ ZS
he, ИР 2277 LLL =i) —
a OIA,

2

, ПОР:
о

—— 4

378 Fig. 201. Monk seal, Monachus monachus (figure by М. М. Kondakov).

species of the true seals (Phocinae) with a heavier and apparently slightly
shortened anterior portion of the body.!””

The head is flattened on top, as for example among the harp seals
(Mohr, 1952). The fore flippers are somewhat shorter than the hind ones.
The claw zone on the fore flippers appears as though incised in a strait
line while the length of the claws decreases quite uniformly from the
first to the fifth digit. The hind flippers have broad terminal lobes due
to an angular folding along the inner edge turned toward the apex of the
median notch, which is deeper than in the true (10-incisored) seals. The
claws on them are highly reduced, very narrow, and barely visible (their
length does not exceed 0.5 cm).

The claws on the fore flippers are not pointed but broad.

The hair coat of adults is rigid, smooth, adheres closely to the body,
and very low. The hairs are very thin; their length on much of the body
is about 0.5 cm (King, 1959*). On the dorsal side of the body of females
from Kilii delta of Dunai River, the longest hairs (5.3 mm) were found
on the nape, their length on the back and in the tail section being 3.3
and 2.5 mm respectively. The hairs are longer on the belly: the longest of
them are in the midportion of the trunk (10.9 mm) and slightly shorter

129 This impression is gained from some published photographs (Mohr, 1952; King, 1956;
Heck, 1966). It is highly possible that this is due to the jugular section of the male actually
appearing in the photographs as massive and comparatively short while the head appears
large with a slightly swollen snout. Confirmation of this is the wide spacing of the zygomatic
bones in the skull of large males exceeding that in females (see p. 506). Quite possibly,
sexual dimorphism does exist among monk seals in the size of the body and the dimensions
of the skull, although no direct evidence is available (see p. 506). However, the suggestion
that the head of this species is unusually large and resembles that of a bear (Sal’nikov, 1967)
is evidently due to a misunderstanding and is explained as due to the poor preparation of
a stuffed animal.

379

504

on the fore portion (10.5 mm). The hairs are the shortest in the tail
section (3.1 mm). The thickness of the hairs is identical in all sections
of the body (0.1 mm) (Sal’nikov, 1959).130

The neonatal pelage is at least three times longer, incomparably
denser, and altogether without nape. It consists of soft, thin hairs.

The labial whiskers are set in five or six rows; their number in a
row on each side of the lip decreases gradually from eight to two as
the number of the row increases. There is one more lone whisker on
top. The total number of whiskers goes up to 70. Their length varies in
relation to the position of the row as also the height of the row and
probably depending on age and even season (assuming that the whiskers
are sloughed during molt). Among the young (less than two years of age)
their length varies from 2 to 15 cm. They are oval in cross section, without
wavy edges and highly tapered toward the apex. There are hairs like
whiskers even in the chin region, their length not exceeding 2 to 3 ст.131

Supraorbital whiskers do exist, contrary to the opinion of some
authors, even among the young animals (not less than three).

The color!*? of the hair coat of the adult animals is dull and quite
often monochromatic. The characteristic predominant shades are gray,
brown, and black, as also contaminated tones; the dark color is intensi-
fied on the dorsal side of the body compared with the ventral. There is
something like a color bloom, creating a faint, bright white or yellowish
hue, due to the alternation of light-colored tips of the hairs with compact
dark hairs.!*

The upper side of the body of adult males is usually a vivid dark
gray or slate-black with an admixture of fairly distinct brown shades. On
the body flanks, the color turns pale and imperceptibly transits into an
even lighter but still vivid gray on the lower side of the body. In this

130 According to some other sources (Schapp, Helwing, and Chizelea, 1962), the length
of the hairs among the much younger animals varies from 4 to 7 mm on the ventral side
of the body.

131 The description was based on the data of King (1956), Sal’nikov (1959), and Schapp,
Helwing, and Chizelea (1962).

132 Description of the color features of the hair coat is very difficult because of the
extremely few specimens. The number of skins in our museums is very small and such
altogether absent for the other species of the genus. Further, due to prolonged storage, the
skins have acquired a fairly distinct yellowish-golden hue.

133 There is one more widely prevalent type of coloration in the form of fine light-colored
bands or variegations, quite abundantly dispersed in large patches at different places on
the body. These are concentrated most often under the chin, along the flanks, and at the
center of the neck, and at various places on the chest and back. Since these variegated
spots are visible (seen through) on the reverse side of the skin (on the flesh side), they may
be regarded as caused by abrasions.

380

505

background, а very large spot stands out in contrast. This spot is tens
of centimeters across, angular, almost white or slightly creamy, rhom-
boid or roughly rectangular, and often resembles a butterfly. This is a
characteristic feature of an adult male. The spot is usually disposed asym-
metrically, most often on the abdomen or on the flanks in the rear half
of the body but closer to the fore flippers. Among adult females (King,
1959*; Sal’nikov, 1959), the dorsal side is dark, blackish or dark gray,
with a slightly silvery or yellowish bloom due to the light-colored tips
of the dark brown hairs. On the flanks, the hair coat gradually turns
lighter and becomes light gray on the entire underside of the body, but
usually without the large whitish spot characteristic of the males. The
yellow bloom often seen on the skins is most likely due to posthumous
color changes caused by oxidation of fat remnants present in the skin.
The color of the tips of hairs among live animals is mostly whitish or
very pale gray, if not white.

The pelage of the newborn is a fairly uniform dark brown or dark cin-
namon on the dorsal surface; it turns lighter on the flanks and apparently
is even lighter in color on the ventral side.

Evidently, depending on whether the animal is a male or female,
on the underside of the body or on the flanks, posterior to the fore
flippers, there is, or is not, a large angular spot of white or almost white
color similar to that described in the case of adult males. As the neonatal
(juvenile) coat is shed, a rigid, very short hair coat closely adhering to the
skin is seen; this coat is an altogether different color and is characterized
by sharp dichromatism: dark gray on the dorsal side and very light or
almost pure white on the ventral side (or ventral side with yellowish
tinge). It may be sharply set off on the flanks, roughly at the level of the
base of the fore flippers from the dark gray with brown bloom of the
dorsal side (Schapp, Helwing, and Chizelea, 1962). Apparently, at this
early age, the color of the first definitive hair coat is identical among
males and females.

The color of the males of transitional age is highly similar to that
of adult animals of the same sex. The former, however, lack the large
light-colored spots of the type described above which are characteristic
of the adult (and possibly the newborn) males (King, 1959*; Kumerlowe,
1966* ).

The skull is somewhat larger than in the other species of the genus
(Fig. 202). Its upper contour when seen in profile usually descends quite
steeply toward the fore end roughly from the apex of the nasal bones.
The narrow and long zygomatic bones have an anteroventral angular
margin distinctly restricted outwardly along the suture. Preorbital pro-
cesses are distinctly manifest in the form of a nipple. The nasal bones

380

381

506

vise oy о и
ий y ИИ MH SN А о
М =

Fig. 202. Skull of monk seal, Monachus monachus (figure Бу М.М. Kondakov).

are anteriorly bifurcated by a median angular notch; the anterior end
of each of them is in the form of an angular or tonguelike prominence.
The frontal-maxillary suture from the nasal bone to the flexure in the
orbit is not shorter than the nasal-maxillary suture and is usually longer.
The bony tympanic bullae are roughly triangular, of irregular form, highly
elongated forward, lean somewhat toward each other, and terminate con-
siderably anterior to the crest of the articular fossa. Their internal margin
is concave with the aboral carotid fossa at their very end. The fontanelles
in the choanal region are very large. The posterior margin of the bony
palate is in the form of an arc with a notch, often asymmetrical.

The condylobasal length of the skull of adult males is 280-302 mm,
of adult females 260-280 mm. The zygomatic width in adult males is
190-215 mm, in adult females 160-180 mm.

The teeth are very massive. All the lower premolars and at least two
upper ones are set aslant relative to the tooth row. Their crowns are
formed mainly with a single massive main cusp at the base of which is a
broad fringe (cingulum) from inside. Anterior or posterior to the base of
the main cusp or on both sides of it, quite often an additional denticle,
barely raised above the cingulum, appears. All the teeth, except the first
premolar, have two roots. The upper incisors are disposed arcuately: the

507

median less large ones are shifted forward slightly compared with the
extreme ones. A semicircular notch and fringe in the form of a cingulum
is seen On the posterior side of all the upper incisors.

The body length of adult males from tip of nose to tip of tail varies
from 210-250 cm; some very large animals could be of much larger
proportions. Nevertheless, the figure of 300 cm and above cited in the
literature (М. Smirnov, 1929; and others) is hardly reliable.!*4

The adult females are probably somewhat smaller although a ges-
tating female caught on the coasts of Corsica measured 278 cm (evidently
up to the tip of the flippers) (Troitzky, 1953). The length of a female
from the Kilii River bed of Dunai (if not sexually mature, quite close to
being so) measured 227 cm (Sal’nikov, 1959). The total body weight of
adult animals goes up to 300 kg and may even exceed it. (K.Ch.)

Taxonomy

The monk seal although undoubtedly representing the ancestor from
which other species of the genus have evolved, is more specialized in
some respects than its descendants. This is manifest in the intensification
of the dental apparatus (premolars and molars more massive with a more
sharply manifest tendency to a transverse setting, i.e., to the displacement
of their anterior edge inward), in the intensification of the maxillary
musculature manifest in a broadening of the coronoid process and the
lower posterior subcondylar portion of the lower jaw. The enlargement
of the zygomatic arches is more significant than among the other species;
the transformation of the bony tympanic bullae has proceeded farther,
more forward advanced, etc.

The phyletic relations of the monk seal with the other species, in a
morphological context agree with the probable direction of the dispersal
of the species of the genus: the Caribbean seal is closest to the European
seal in the structural features of the skull while the Hawaiian seal differs
most. The two former species are related particularly in the steep curva-
ture of the upper line of the profile and generally the greater height of
the skull, the greater mass and tapering disposition of the cheek teeth,
enlarged subcondylar portion of the lower jaw, structure of the anterior
region, general shortening of the zygomatic bones, and structural fea-
tures of the bony tympanic bullae. On the other hand, the Caribbean
seal can be considered closer to the Hawaiian seal in the proportions of
the skull (low index of its width) and in some plastic features (length of

134 Тре body length of adult males as established from the published literature, makes
no claim on adequate accuracy since not a single author has specified the method of
measurement.

382

508

the nasal bones and their contact with the premaxillaries, structure of the
coronoid process, pterygoid bones, contours of the suborbital apertures,
etc.). Some craniological features among the Caribbean seals are of an
intermediate character (reduction of the preorbital processes, form of
the anterior margin of the nasal bones, length of the zygomatic bones,
etc.) (King, 1959*).135

The following general features can be cited:

1. The Mediterranean monk seal, M. monachus, represents a special
species. Some structural features which are of a more primary (primitive)
nature are characteristic of it (arcuate setting of the upper incisors, upper
contour of skull profile steeply dipping forward, presence of sharp pre-
orbital processes, etc.), proximating it to the 10-incisored seals (angular
form of the lower anterior margin of the zygomatic bones).

2. Of the other two species, the Caribbean seal, M. tropicalis, is more
related to it than the other species, while possessing several features
bringing it closer to the Hawaiian М. schauinslandi.

3. М. schauinslandi combines in itself craniological features of a more
primitive character (bony tympanic bullae and some swelling of the pre-
orbital zone of the rostrum) as also features that are more evolved (long,
arcuate zygomatic bones that are concave anteriorly and smoother upper
line of the profile). (K.Ch.)

Geographic Distribution

Coastal waters of the Mediterranean Sea along the continental coasts
of Europe and Africa and also the coastal islands, including those away
from the mainland. Parts of the Atlantic Ocean southwest of Gibraltar
up to the Madeira and Canary islands inclusive, mainland coasts of Africa
in the south up to 20 to 15° М lat. (Cape Cabo Blanco in the Senegal
River estuary), western, Balkan, and southern Asia Minor coastal zones
of the Black Sea (Fig. 203).

Geographic Range in the USSR

Forms a negligibly small portion of the northeastern rim of the range.
The reconstructed boundaries of distribution in the last century or
earlier covered the western strip of the Black Sea adjoining Romania
within the southern part of the coast of former Bessarabia with the region
of Zmeinyi Island (Krotov, 1952), the sea along the western coast of

135 When evaluating these features, it should be remembered that the data for all the
species are extremely scant and the extent of individual variation of several features is not
known. All these could be a source of error.

382

383

509

Fig. 203. Reconstructed distribution of the monk seal, Monachus monachus т
the USSR (К.К. Chapskii).

Crimea, possibly both sides of the Tarkhankut Peninsula (Zernov, 1913),
and farther south and east of the southern coast of Crimea from Cape
Kherson roughly to Gurzuf (Nordman, 1840; Керреп, 1883*; Puzanov,
1929). Some small tongues of distribution intrude from the side of Asia
Minor waters into the extreme southeastern part of the Black Sea toward
Batumi (Sal’nikov, 1959). In the early 1840s, these seals were encoun-
tered quite often on Zmeinyi Island (Nordman, 1840; Puzanov, 1969*).

Reports on the habitats of monk seals in the region of Sevastopol
go back to ancient times although they indicate that seal sightings were
generally rare. At the end of the eighteenth century, it was likewise
reported that seals were “rarely seen on the coasts including Sevastopol
harbor, unlike at other places where they are often sighted” (Gablits!’,
1785*). 38

136 Interestingly, Pallas (1811*), who lived for quite sometime in Crimea at the beginning
of the last century, reported that seals in the Black Sea came only from the Mediterranean
contd...

510

The presence of these seals on the southern coasts of Crimea in
the first half of the nineteenth century was more accurately reported by
Nordman (1840) who indicated sightings in the caves on the rocky south-
western coast. The seals were few, however, and occasionally became tar-
gets for hunters. It is interesting that all the recent published reports on
this animal are usually a repetition of the earlier publications. Reports
of original finds of these seals are extremely rare. Thus, in the 1830s, one
seal was killed between Kuchuk-Lambat and Karabakh (Keppen, 1883*)
and, in 1834, a very large seal was “sighted on a cape at the end of the
land in a botanical garden” (A. Nikol’skii, 1891).

At present, the monk seal lives or reproduces nowhere in the western
part of the Black Sea. The appearance of stray animals in the extreme
northern portions of the Dunai delta represent extremely rare instances
of transgression far beyond the limits of the habitats that are normal for
our time. Such instances were noted at the end of the 1930s and early
1940s at some points in the Kilii delta: on Siberian spits, in the narrow
straits of Prorva where seals were caught in fishing nets, and also at
several other places in the water as well as on sand spits, including the
small islets Limba and Kuril (Sal’nikov, 1959). From 1946 through 1951,
there were five more instances of seals being caught by fishing hooks
(Krotov, 1952), including a female on May 20, 1950. Zmeinyi Island
too falls in the same coastal section of the northwestern Black Sea but
straying of the monk seal onto this island is as rare an event as in the
Dunai delta. There is no positive information whatsoever on finds of
this seal farther north along the western coast of the Black Sea barring a
reference to an “animal like that of a seal” noticed 30 km east of Odessa
in 1950 (Kleinenberg, 1956).

There is no positive information on sightings of the monk seal on the
Crimean coasts in this century and it may be regarded as totally extinct
there (Puzanov, 1929). In Tarkhankut-Bokkal region (a unique place
where it was still considered possible to sight an occasional seal early in
the nineteenth century and early in this century), the seals were obviously
more numerous in the remote past (Zernov, 1913). There is no truth,
however, that seal populations lived and reproduced on the Tarkhankut
Peninsula in the last century. Significantly, there are no other references
in the literature of that time to the existence of this seal. Yet, there are
persistent reports of seal sightings in the region of Medvezhii caves near
Sevastopol beyond the Kherson lighthouse, where these seals were even
hunted at the end of the last century (Zernov, 1913).

(“т Pontum Euxinum adscendit em Mediterraneo’”’). However, he was simply repeating the
widespread belief of the local inhabitants.

384

511

There are three references in the literature pertaining to the south-
ern coast of the Caucasus but they do not inspire confidence. One states
that on the coasts of Adler region a whole herd of animals “with promi-
nent snout and whiskers” was seen once (Poznanskii, 1880); another con-
cerns Batumi region near which some fishermen killed a seal in water
but could not get it out because it drowned (Kleinenberg, 1956). In the
1920s, near the rocks in the region of Cape Zelenyi (close to Batumi), a
few seals settled for several years. Over the course of time their number
dwindled and the last of the animals was killed in 1933 by a dolphin
hunting ship cruising in Chernaya River estuary (Sal’nikov, 1959). The
greater inhabitation of the coasts and the absence of appropriate biotopic
conditions are adequate reasons for disbelieving the appearance of monk
seals on the southern fringes of the Caucasian coast.

Geographic Range outside the USSR

In the past this covered almost the entire coastal strip of the Black Sea
along the Balkan Peninsula (from Dunai to Bosfor) and Asia Minor. In
the Mediterranean Sea basin the distribution encompassed almost the
whole sea coast and the islands except for the long inhabited sections on

_ the southern coast of Europe (most of the French and Italian coasts and

many parts of Spain and Greece) and also some sections on the eastern
coast and African coast. Outside the Mediterranean Sea, the distribution
covered a small section of the Atlantic Ocean along northwestern Africa,
in the south up to Cabo Blanco and even the estuary of the Senegal River
(15° N lat.) and also included the Madeira and Canary islands.

Even at the beginning of the present century, much of the coastal
waters surrounding the southern coasts of Europe fell outside the distri-
bution while the expanses interrupting the distribution along the African
and Asia Minor coasts enlarged. In the 1960s, the distribution com-
prised a few isolated sites of essentially disconnected diminishing pop-
ulations. Some are protected by the government, especially in Bulgaria,
Yugoslavia, and Rio de Oro.

Only stray wandering animals have been sighted, and extremely
rarely, on the Romanian coasts. In the first 50 years of this century, only
nine cases of monk seals caught in fishing nets in Romanian territorial
waters were recorded and the last such catch (in July 1960) after an
interval of 12 years (Schapp, Helwing, and Chizelea, 1962).

In Bulgaria there are two zones of reproduction, both with extremely
small populations, namely Cape Kaliakra and south of Burgas. The rest of
the Black Sea distribution encompasses the coastal belt of Turkey (Ana-
tolia). The boundaries and the nature of distribution of the species there

385

512

are not clearly known but the population can be regarded as extremely
small and widely scattered, mainly in the western regions.

Outside the Black Sea, there are a few places where some small
reproducing colonies still exist and also some regions where these seals
are encountered but their population and reproduction have not been
established despite their presence being known for 20 to 30 years. Repro-
ducing populations of the Mediterranean Sea are encountered on some
islands in the Adriatic Sea along the Dalmatian coast of Yugoslavia,
the Lebanon coast, and possibly here and there the Turkish coast; the
existence of one or some small colonies in Greek waters and on the
Ionian Islands is quite probable; their presence has been established with
certitude on the small islands near Tunisia, in the waters of Morocco,
southern Sardinia, and Corsica. In addition to the aforementioned sites,
some Stray seals are sighted from time to time in the Sea of Marmara, in
the Dardanelles, along the eastern fringe of the Aegean Sea, in Salonik
Bay, on the Rodhos, Kipr, and Kriti islands, in the northeastern corner
of the Mediterranean Sea, and on the coasts of Syria and Israel; the
presence of monk seals is also probable on the Egyptian and Algerian
coasts, on the Baleares, and at other places.

These seals have been reported southwest of Gibraltar in the oceanic
part of their distribution in the region of Medeira (Desert Island) and
especially in the continental coastal sections of northeastern Africa to
the south of Rio de Oro (in the region of Cabo Blanco). The southern-
most point on the coast of northwestern Africa for which information is
available about the presence of this seal is the estuary of Senegal River
(about 15° N lat.). In general, however, the southern boundary of distri-
bution on the Atlantic coast of Africa roughly corresponds to the 20°
winter isotherm of water. (K.Ch.)

Geographic Variation

No information whatsoever is available on the morphological features
of the local populations. (K.Ch.)

Biology!*’

Population. Right now, it is not appropriate to refer even to the very
existence of the monk seal, let alone its population in our ferritorial
waters. In the remote past, however (a century or sO ago), as already

137 Information on biological aspects is based mainly on foreign sources which, however,
are neither abundant nor thorough.

386

513

stated before (р. 510), the seals were regarded as quite common оп
Zmeinyi Island. They were evidently reported in very small numbers on
the southwestern coast of Crimea including Sevastopol region although,
Over a century ago, it was stated that seals in the Black Sea were “few
and only rarely stray animals reach the northern coast” (Kessler, 1861).

The population is not much on the coasts of Bulgaria and Anatolia
of Turkey. In spite of a total ban on hunting and strict conservation
for many decades, the population on the Bulgarian coasts has shown no
upward trend. Just 20-30 animals have been reported in the region of
Cape Kaliakra (Caspers, 1950). The population of the colony south of
Burgas does not even exceed half this number. The Bulgarian population
is essentially the only source for the dispersal of this seal in the northern
part of the western Black Sea including Zmeinyi Island and Crimea. The
rarity of the seal on the Romanian coasts has already been commented
upon.

It is not possible to draw a picture of the present population of
the monk seal on the Turkish coasts of the Black Sea. Some sixty years
ago, 3 or 4 seals were encountered on the Anatolian coasts during a
3-week cruise. This gave grounds for the statement that the sighting
of this seal on the Anatolian coasts “was not so rare” (Zernov, 1913).
Somewhat later, and also during a three-week cruise, “10 to 12 families”
were counted on the west coast of Anatolia, at Zonguldak (Caspers,
1950; Mohr, 1952). There is almost no information from Turkey for the
1960s and that available does not provide a complete picture (Mirsaloglu,
1964 from Kiimerlowe, 1966). It is only known that the seals are sighted
every year not only on the Mediterranean coasts, but also on the Black
Sea coasts and that small colonies probably do exist on the Anatolian
coasts as well (Wijngaarden, 1966* ). Since, however, no seal conservation
measures have been practiced in Turkey in the last half a century, its
population has been considerably depleted and at present is hardly more
than the Bulgarian population. Had the situation been different, solitary
animals would have been sighted in our waters on the Caucasian coast
in the last 30-40 years.

The number of Mediterranean populations, not exceeding 20, is
extremely small. Their existence is reliable only at three places: 1) island
on the Dalmatian coast of Yugoslavia, 2) northern coast of Corsica, and
3) Tunisian islet Galata. Moreover, two populations of indeterminate
strength are encountered on the Ionian Islands and south of the Aegean
Sea. The existence of colonies on Crete, on the southern coast of Turkey,
and also on the coasts of Libya east of Benghazi is doubtful. The largest
population of monk seal is found on the Atlantic coast of Africa, south
of Rio de Oro, directly around Cabo Blanco, where some 200 animals (or

514

even more) have been counted (Kusto and Dyuman, 1953; Wijngaarden,
1966* ).

The total number of all the populations of monk seal equals less
than a thousand. This number is so small that the species is threatened
with extermination.

Habitat. Being an aegialoid [littoral] animal, 1.е., biologically associ-
ated with coasts, the monk seal is confined predominantly to the coastal
zone and is not found deep at sea. Since, however, the coasts within the
distribution zone have been extensively inhabited for a long time now, the
seals have taken to vacant or less populated sites for breeding. These are
mainly rocky islands and generally those sections of the mainland coast
which are not inhabited by man. Thus in Crimea the rockier and more
rugged sections of the southwestern coast are regarded as sites of their
habitation. The seals rarely take to open and low beaches. At least the
seal colonies known at present are not associated with such biotopes.
It is possible, however, that in the past they did live on the vacant and
open beaches of Zmeinyi Island but these were probably only temporary
rookeries not associated with breeding. The shallowness of the coasts
and their low level explain the absence of permanent rookeries on the
vacant and low coasts in the narrow straits of Dunai (Sal’nikov, 1959).

At various sites outside our waters, the seals occupy predominantly
the uninhabited and less inhabited islands, more rarely the mainland
coasts with complex, highly rugged relief, abounding in rock clusters and
various types of rocky recesses. They generally select caves and grot-
toes, especially those with an underwater inlet. In such rock crevices and
caves, protected from the direct action of the surf, the animals eagerly
take to sandy-pebbly beaches. Similar biotopes with caves and grottoes
abound in the Black Sea (on the Bulgarian coasts at Cape Kaliakra and
on the Turkish coasts of Anatolia), in the Mediterranean Sea, and also
on the ocean coast of northwestern Africa (including Cabo Blanco). In
this context, the explanation for the absence of this seal south of Cabo
Blanco, i.e., the coast there is low and sandy, is rather intriguing (since
not wholly correct) (Bétger, 1951*).

Food. Almost nothing is known about the food of these seals in
the Black Sea waters of our country. The stomach of females caught in
Kilii, the narrow strait of Dunai, revealed about 3 kg of semidigested
remnants of flounder (Rhombus maeoticus) (Sal’nikov, 1959). The catch
of seals coincided with the en masse approach of this fish to the coasts
for spawning. Judging from the massive teeth of the monk seal, one
may assume that it specializes in consuming large fish. This has been
confirmed by seal food preferences in zoological gardens and the stom-
ach contents of some specimens caught in the Mediterranean Sea. Thus,

387

SS)

on the coasts of Sardinia the stomach of these seals revealed the rem-
nants of wrasses (Labrus) and sparid fish (Dentex) (Carrucio, 1893; King,
1959*). The seals were noticed playing with large fish measuring about
70 cm in length. They caught them in their mouth after throwing them
up in the air, with the head of the fish turned directly into the mouth
(Caspers, 1950). It is therefore hardly correct to suppose the food of the
monk seal to be large fish, especially such massive and migratory fish
as anchovy and sprat; given the sluggishness of the monk seal, catch-
ing such large fish would be difficult (Sal’nikov, 1959). On the African
coasts, in addition to fish, the seals consume thorny lobsters (Palinurus).
According to fishermen in the region of Cape Kaliakra, a large seal can
consume 15-20 kg, with a preference for sturgeons (Caspers, 1950), but
this estimate is doubtful. The daily ration of the seals in zoological gar-
dens reached 12 kg (Gavar, 1927*) and they consumed all types of fish,
such as whiting, eel, sardine, salmon, etc.

Home range. Under minimal favorable conditions, monk seals are
usually confined year round to the same section of the coast in which they
breed year after year, i.e., the seals lead a settled mode of life. The dimen-
sions of the selected sections evidently vary depending on such factors
as the size of the population, availability of fairly suitable hideouts, food
conditions, etc. The actual population regime on Cape Kaliakra con-
forms to these factors. The boundaries of the occupied coastal sections
could only be tentatively established for each of the given populations.

Daily activity and behavior. Almost no observations on the behavioral
features of the animals transgressing into our waters have been reported.
Data on the behavior of the monk seal outside our country are likewise
scant.

The seals are regularly encountered outside their hideouts in the day
(Caspers, 1950; and others) but may be found sleeping even on the coast
in grottoes when the brightness is not sharp (Troitzky, 1953). Contrarily,
they exhibit the same mobility even in darkness (Kiimerlowe, 1966). It
may be assumed that there is no strict sequence of sleep (night) and
activity (day). They can be regarded as more diurnal than nocturnal ani-
mals. In this respect, observations on the behavior of these seals on Cape
Kaliakra are of interest. With surprising stereotyped regularity, noticed
for hours on August 23, 1941, the seals swimming from the southern
side dove roughly 20 m from a rock. Later, they remained submerged for
brief intervals, obviously in shallow water, often surfacing in between,
at times with the head ducked and only the back showing above the
water. It appeared as though the animals were searching the bottom for
something. This probably was so, considering that the water around Cape

388

516

Kaliakra is exceptionally transparent. Leisurely swimming thus, submerg-
ing and surfacing, the animals turned away for some distance from the
rocks. They continued the reverse journey in the same manner until they
were lost to sight. Nearly eight minutes later they again appeared and
repeated the selfsame behavior. Before submerging, the animal raised its
body quite high above the water and flexed its forepart so as to assume
the required direction at depth. While diving, air was often exhaled even
inside the water, which bubbled up in the form of an arc until the head of
the animal surfaced. From this arc one could judge beforehand the point
of surfacing. On surfacing, the hissing sound of the animal’s expiration
was Often audible (Caspers, 1950).

Monk seals are generally good divers; they usually throw themselves
into water from rocks and are somewhat like sea lions in this respect.
They first assume an appropriate posture by heaving the chest.

It has been assumed that monk seals are not particularly good swim-
mers (Sal’nikov, 1953). Some observations on their swimming capabilities
from the coast and from ships (Caspers, 1950) do not justify changing
this view. Nonetheless, a slowly swimming animal could hardly seize a
large fish. Divers with aqualungs could not approach adult monk seals
to photograph them (Kumerlowe, 1966).

The relations between individual animals are hardly known but
Bulgarian fishermen were witness to scuffles which, in their view, arose
when a new animal entered the colony. The reports of fishermen on
regular and severe scuffles between adult females sound somewhat
strange. Scuffles are accompanied by loud laryngeal cries resembling at
times the bellowing of calves, barking, and even the scream of a beaten
or bitten dog (Wolf, 1818*; Carrucio, 1893; Caspers, 1950; Morales-
Agesino, 1950* ). The voice of young seals is generally similar to that of an
adult but fainter and less “fervent” (Morales-Agesino, 1950*) while some
have a more characteristic voice. A very young specimen caught close
to Livorno, uttered the sound “ovaavaavava” in captivity, sometimes
something like a sneeze was heard interspersed with an intense dull
sound, and at times the animal emitted a bleat or simply a loud scream
(Wolf, 1818; after Mohr, 1952). A newborn seal caught near Split gave
out laryngeal sounds similar to “oa” (Priemel and Mohr, 1952*).

Observations of the behavior of captive animals have revealed well-
developed higher nervous activity. This is manifest in comprehension
and great affinity to man, as noted in its ability to respond to his com-
mands to some extent. A very young animal from around Livorno became
totally domesticated after a few months and would “kiss” its master while
producing a sound similar to belching. On being brought to Nurnberg,
it became totally silent (Wolf, 1818; after Mohr, 1952). Another pup,

389

S17

which had learned to recognize its trainer, would cry out on sighting her
(Ber, 1838*).

Seasonal migrations and transgressions. Being generally confined to
a single section of the habitat, monk seals do not undertake periodic
or distant migrations. At the same time, the transgressions of stray sin-
gle animals to fairly long distances from the breeding site are widely
known although not frequent. Such animals were sighted in almost all
parts of the range and even at its very boundaries. The reasons for these
transgressions are not known. It is difficult to say for certain whether the
seasonal arrival of fish for spawning plays a significant role in this respect
in certain regions. Nevertheless, this phenomenon in particular is used to
explain the appearance of some animals and their being trapped in fishing
nets in Dunai delta (Sal’nikov, 1959). However, transgressions here are
extremely rare and reveal no regularity whatsoever. Were they regular
migrations, in spite of the extremely small population, the migrating ani-
mals would first have been encountered in somewhat larger numbers and
more often, and secondly their departure from Kaliakri would have been
perceivable. But, far beyond the boundaries of the colony, on Cape Kali-
akri, single animals and almost invariably young ones are encountered
(Calinescu, 1936). These phenomena can more naturally be explained as
cases of transgression.

Reproduction. Very little is known. The period of mating follows the
cessation of lactation although it can commence even somewhat earlier
and is evidently protracted. Its calendar periods vary for different animals.
Most animals evidently mate in the autumn or at the end of summer.
The total duration of the embryonic period (from the moment of mating
of the female to parturition) has been put at 10 (Caspers, 1950) or 11
(Troitzky, 1953) months. The whelping period falls at the end of summer-
autumn. It is July-August for the Bulgarian populations (Caspers, 1950).
Much earlier periods have been reported for the Anatolian coast pop-
ulations of Turkey: a female caught near Zolgundak delivered a pup in
a zoological garden at Ankara on May 5, 1962 (Mursaloglu, 1964 from
Kumerlowe, 1966). According to other indirect data (Zernov, 1913), the
period of whelping on the coasts of Anatolia is roughly in the second
half/end of July. In the Mediterranean Sea the whelping period extends
from mid-August through September and even October (Carrucio, 1893;
Dathe, 1934; Troitzky, 1953) and on Rio de Oro, even later, possibly
around mid-November (Agasino, 1950 from King, 1956). The reference
to spring whelping (Postel, 1950*) is evidently not wholly correct.

It is impossible to establish the dimensions of the newborn because
of the absence of information on the method of measurement and on
age. Pups with a fresh umbilical cord, found in Dalmatia on September

518

19, 1933, measured roughly 90 cm and weighed 26 kg (Рае, 1934).
A fully mature fetus was removed from the womb of a female killed
on September 27, 1947 on the coast of Corsica; it measured 120 cm in
length and weighed 17 kg (Troitzky, 1953).

Lactation extends for 6-7 or 7-8 weeks (Caspers, 1950; Troitzky,
1953). The total duration of the reproductive cycle according to some
authors (Caspers, 1950) is 12 months and according to others (Troitzky,
1953), about 13 months and hence births are not an annual phenomenon.
It is difficult to say which view is more correct but nevertheless it may
be assumed that births are an annual feature in most cases.

Growth, development, and molt. Accurate data are not available on
postnatal growth and development. Since the length of the smallest pup
at the end of December in Rio de Oro measured 150 cm (Morales-
Agasiono, 1950*), it may be assumed that the pups add 25 -30 cm during
lactation. The period of molt of the neonatal hair coat is not known. It
is also not known whether the pup remains on the coast throughout lac-
tation or goes into the water. Some young ones with the neonatal coat
fully preserved were caught in the sea. Thus, the Russian hydrobiological
expedition in mid-August of 1912 caught on the Anatolian coast a pup
that was still completely covered in a firm neonatal hair coat. It molted
only after a few weeks (Zernov, 1913). In the second case, a newborn
pup with a fleshy umbilical cord was caught on the coasts of Dalmatia.
Both these cases point to the ability of even extremely young animals
to enter water without molting the primary, neonatal coat. It has been
pointed out in the literature, however (Troitzky, 1953), that the pup goes
into the water for the first time at the end of lactation and possibly molt.
The embryonic hair coat is thus sported for not less than 1.5 months.

A newborn in captivity, weighing 26 kg, was bottle-fed for one week
with a mixture of half oatmeal and half cow’s milk with a small addition
of cod liver oil. This mixture was given six or seven times a day. The pup
died due to injuries sustained during transport (Dathe, 1934).

The total growth duration before the onset of sexual maturity has
not been established. It has been assumed that the monk seal reproduces
for the first time at about four years of age (Troitzky, 1953).

Enemies, diseases, parasites, mortality, and population dynamics. Six
species of helminths parasitizing the gastrointestinal tract were detected
among monk seals (Delyamure, 1955): cestodes Diphyllobothrium hians,
р. latum, О. lanceolatum, and Diplogonoporus tetrapterus; and nematodes
Contracaecum osculatum and Terranova decipiens.

These seals face no competition with other pinnipeds. Besides man,
sharks pose a threat to the juveniles at some places.

390

59

Field characteristics. This is a large seal of deep dark monochromatic
coloration on the dorsal side and a different coloration on the flanks and
belly depending on age and sex. A large light-colored spot in the pos-
terior half of the body is a characteristic feature among pups and adult
males. This is the only species of seal in the Mediterranean and Black
seas and also on the Atlantic coast of northwestern Africa. (K.Ch.)

Economic Importance

This seal has no economic importance because of its small popula-
tion. In some countries (Bulgaria, Yugoslavia, and in Rio de Oro), this
animal is protected. It needs to be protected throughout the range of its
distribution. (K.Ch.)

Subfamily of Hooded Seals and Elephant Seals,
or 6-incisored Seals

Subfamily CYSTOPHORINAE Gray, 1866138

These are seals of large and extremely large dimensions. The body length
with tail along the dorsal surface (Lc) of adults varies from 180 to 500 cm
or more, the males being slightly larger and heavier than the females.

The hind flippers are slightly longer than the fore flippers. The first
two digits of the fore flippers are longer than the third and subsequent
ones. The claws are quite massive. By the time of maturity, the males
grow a dermo-muscular, sac- or proboscislike process on the anterior
upper portion of the snout joined to the nasal cavity. This is capable of
considerable enlargement on being filled with air. There is one pair of
teats.

The skull is massive, broad, with a roomy cranium and zygomatic
arches markedly protruding laterally; the zygomatic width considerably
exceeds the width of the skull between the mastoid processes. The nasal
processes of the premaxillae fall far short of reaching the nasal bones.
The nasal opening is wide open and its upper and lateral margins extend
far backward. The preorbital processes are well developed.

138 The independence of the subfamily Cystophorinae has come under dispute quite
recently (King, 1966). Considering the controversial nature of this new interpretation of the
family Phocidae, however, its existing structure has been retained. The subfamily is repre-
sented in our waters by a single species, which is rather a random find. It nonetheless plays
a role in our sea hunting activity in the North Atlantic and hence has been described fairly
fully here. The species is also of interest because its biology and distribution were studied
quite completely only in the last decade; yet although our scientists put considerable effort
into its study, still little is known about it. (У.Н.)

392

520

The incisors in the upper jaw number two and, in the lower jaw,
one on each side. The extreme upper lateral incisors are similar to the
canines and slightly longer and thicker than the medial ones. At least the
first two premolars have a single root and an almost wholly undivided
crown. The cheek teeth are relatively small.

Biologically, the species of the subfamily are extremely divergent.
Some of them are pagophilic with no association with the land (hooded
seal) while others have almost no association with ice floes and reproduce
on land (elephant seals). All of them undertake extensive migrations.
In the period of reproduction, males are confined mainly to one and
the same rookery along with the lactating females and offspring. The
pups at birth are not covered with a luxuriant embryonic hair coat of
nearly white color, characteristic of the pups of most of the 10-incisored
seals (Phocinae), but rather with a short firm hair coat. Some mem-
bers of the subfamily (hooded seals) live in pairs at the time of mating
while others (elephant seals) form harems. The former mate in water and
the latter on land. Fairly large cephalopods and fish play an important
role in their nutrition. Hooded seals molt far away from their breeding
sites. Elephant seals, however, molt in the very same rookeries and with
a far more intense scaling of the epidermis, which comes off in large
patches.

The distribution is interrupted and bipolar (borealnatal). Some
members of the subfamily are distributed in the arctic and subarctic
Atlantic (hooded seals) and others (elephant seals) mainly in the
Southern hemisphere, predominantly along the edge of Antarctica, and
enter only insignificantly into the Northern hemisphere on the western
American coasts (Fig. 204).

The origin of the hooded seal dates back to the Miocene. The seals
of this branch originated somewhere in the west of the Old World; the
earliest known finds of Cystophorinae have been dated to the lowermost
Middle Miocene strata of France (Burgundy). The center of origin of the
elephant seals is not known with certainty but possibly falls in the north.
But there is a view that they evolved from the Antarctic 8-incisored seals
(King, 1966) somewhere in the Southern hemisphere. The occurrence
of elephant seals north of the equator in the Pacific Ocean, especially
along the Californian coast is quite a recent phenomenon, evidently in
the most recent stages of the Quaternary period. The route to this place
ran apparently from the subantarctic regions along the Chilean coasts
and was facilitated by the cold Peruvian currents.

The system of the subfamily in the conventionally accepted form
and as adopted here, is simple. It consists of two genera—hooded seals,
Cystophora Nilsson with one species (see ahead), and elephant seals,

5211

Mirounga Gray with two species: M. leonine Linnaeus, 1758 and M. angu-
stirostris Gill, 1866. Some suggested radical changes in the structure of
the family, concerning also the subfamily Cystophorinae (King, 1966), as
mentioned above, cannot yet be accepted.

One genus, Cystophora, is distributed in the northern part of the
Atlantic Ocean; one species of the second genus, Mirounga, in the
Antarctic and the cold temperate waters of the Southern hemisphere
(M. leonina), and the third on the western coasts of North America,
somewhat north and south of 30° N lat. (transgressions into the north
up to Vancouver). The ranges of the two species of the genus Mirounga
are distinctly separate.

Both genera are presently of economic importance but more so the
hooded seal, which is a preferential target of sea hunters. Many rookeries
of elephant seals destroyed in the last century are now being successfully
restored by practicing conservation; at places, as in South Georgia, these
are being exploited strictly along economic lines.

In the USSR fauna the subfamily is represented by a single genus,
the hooded seal, Cystophora Nilsson, 1820. But its presence in our waters
is not a regular phenomenon (see below).

From the middle of the 1950s, the hooded seal began to play a
positive role in the economy of our country, as Soviet hunting ships
were able to negotiate the ice floes in the Greenland Sea. (K. Ch.)

Genus of Hooded Seals
Genus Cystophora Nilsson, 1820

1820. Cystophora. Nilsson. Skand. Fauna. Dagg. Djur., I, p. 382.
Cystophora borealis Nilsson = Phoca cristata Erxleben.

1911. Cystophoca. Brass. Aus dem Reiche der Pelze, 668. Substituted for
Cystophora Nilsson. (У. H.)

These are seals of large dimensions.

The skull of the adults is massive, with thick bones, but relatively
short and broad. The cranium is greatly enlarged but nonetheless stunted.
In spite of the large volume of the orbits, the interorbital space is quite
broaa, slightly more than the maximum diameter of the alveolus of the
canine.

The features of the bony tympanic bullae viewed from below resem-
ble a trapezium somewhat; the posterior carotid aperture is disposed on
their posterior side along the inner margin. The very large and open nasal
aperture is highly enlarged in the posterior upper portion and extends
posteriorly beyond the anterior margin of the orbits. The entire bony
palate is very long and covers more than one-half the total length of the

522

‘puluoa] Dsunop ‘[еЭ$ Зие44э]э WIDYyINOS э41 JO UOTINGLNSIP

Jo souepunog aieunxoidde эцз 1214эр soul] usyo1g ‘эеи1л0ц401$

АЭ ‘s]eas pososioul-9 Jo Апитезап$ ay) Jo чоппа1“а “p07 ‘314

See
Wy 000$ 0009 0006 0005 COCO! 0 001

165

393

573

skull. The posterior margin of the bony palate is fringed with a wavy
line for the most part, like two symmetrical gentle notches converging
medially and forming a projection set backward. This projection often
has an additional narrower notch/bifurcation in the middle. The palatine
bones are elongated and form a very broad bony palate. The choanae
are broad and high; their longitudinal bony septum reaches or falls just
slightly short of the posterior margin of the bony palate. The length of
the zygomatic bones (without processes) is usually not more than double
the smallest width. The lower (or posterior) processes are slightly longer
than the upper. The maxillary bones are contiguous with the nasals in a
small section; their length is usually about one-third that of the nasals
and usually less than their width.

The jugular (paroccipital) processes are well developed and bent
backward. The nasal bones are considerably advanced forward and are
very prominently suspended above the nasal opening. Their anterior
portion is fairly enlarged and usually terminates in an obtusely angled
median projection (see Fig. 206).

The rostral portion of the skull has no bulges anterior to the orbits
and has a concave and not convex (as in 10-incisored Phocinae) profile
when viewed from above. The angular process of the lower jaw is weakly
manifest.

Cranial sutures not persisting until old age, in addition to occipital
(including the lambdoidal) and basal sutures and, in rare cases, even the
anterior palatal suture, are characteristic. The dental formula is:

2 1 4 1
La) Ст» Pig Mi.

The canines are very massive; the lateral incisors in the upper jaw
are slightly larger than the inner ones and similar to the canines; the
molars and premolars are relatively small, with low crowns, slightly flat-
tened from the sides and almost not differentiated. As long as they are
not worn out, only a shallow indentation is visible on some as though
separating the rear section of the crown in the form of a denticle which,
however, may not be present at all. Sometimes, a similar additional den-
ticle is seen in the front. The true molars of the upper and lower jaws
have two independent roots. The rear premolars to quite often have two
independent or partially used roots.

Adult males are perceptibly larger than females. The claws are well
developed on the fore as well as hind flippers. The color of the adults is
bright and spotted. The whiskers are horn or brownish in color, flattened,
and with wavy edges. On the upper side of the snout from the nostrils
to the forehead, the adult males have a hollow dermo-muscular saclike

394

524

process formed by an overgrowth of the nasal cavity. At rest, it is barely
visible and hangs limply with its fore edge above the tip of the snout. It
can be highly inflated during excitation.

These seals are biologically associated with drifting ice floes. They
do not form harems. They feed on fish and cephalopod mollusks.

Distributed in the subarctic and arctic Atlantic from Canada to the
western parts of the Barents Sea, insome years up to the White Sea inclusive.

These seals undertake distant migrations from the wintering and
breeding sites to the regions for molt and feeding. Their main biotope
is the marginal strip of drifting sparse pack ice. In the period of feeding
they cover even much farther boreal regions to the south.

The origin of the genetic branch has not been traced conclusively
with the finds of teeth reported thus far dated to the Middle Miocene
formations of France and the Pliocene formations of Belgium. Mesotaria
ambigua, described from these finds, is regarded as a possible predecessor
of the contemporary genus Cystophora.

An immediate problem for taxonomists is to provide a more
thorough explanation of the morphological differences and similarities
between the hooded seal on the one hand, and species of the subfamily
of true seals (Phocinae) and the subfamily of monk seals, Monochinae,
on the other.

The genus consists of a single species, the hooded seal, Cystophora
cristata Erxleben, 1777.

This seal is a random find in the USSR where it penetrates into the
northern part of the White Sea (its inlet and neck sections).

As one of the most important targets of sea hunting, this seal is caught
predominantly in the international waters near Jan Mayen, Iceland, Green-
land, and Canada for its skin, fat, and other byproducts. (K. Ch.)

HOODED SEAL!??
Cystophora cristata (Erxleben, 1777)

1777. Phoca cristata. Erxleben. Syst. Regn. Anim., I: 590. Southern
- Greenland and Newfoundland.
1785. Phoca cucullata. Boddaert. Elen. Anim., p. 107.
1820. Cystophora borealis. Nilsson. Scand. Faun. Г: 383. Southern
Greenland and Newfoundland.

139 This seal is sometimes referred to as the Atlantic gray seal (“tevyak’”’), which is not
correct as this name has been assigned to another species (Halichoerus grypus). Hunters
sometimes call it the “klappmyss,” a word borrowed from the Norwegians.

525

1824. Phoca leucopla.'*° Thienemann. Reise im Norden von Europa,
р. 102. Grimsey Island, north of Iceland.
1825. Phoca mitrata. G. Cuvier. Oss. foss., 5, p. 210.

1843. Phoca isidorei. Lesson. Rev. Zool., p. 256. d’Oleron Island, France.
(V.H.)

Diagnosis
Only species of the genus.
Description

In general appearance, the hooded seal is similar to the large species of
the subfamily of true seals, 1.е., common or harp seal, except that it is
larger (Fig. 205).

The body is elongated and the fore flippers have a significantly trun-
cated posterior upper margin. The hind flippers do not form a deep
recess ON maximum expansion.

Пре
WZ =

Е SE: FP PLCS: р
I a CE, ae =F
= и ae 2 _

=

и Fr
р
Wye 7

395 Fig. 205. Hooded seal, Cystophora cristata, mature male and female (figure by
М.М. Kondakoy).

140 Ellerman and Morrison-Scott (1961*) assign this name to an individual variant of the
harp seal. (V. H.)

395

526

The length of the anterior portion of the body from tip of nose to
base of fore flippers is about 31% in females and 34% in males; length
of fore flipper in both males and females about 14%, and maximum body
girth about 65% of body length (L. Popov, 1960).

Adult males differ sharply from all the other seals of the North
Atlantic in having a special saclike formation on the anterior upper part
of the head. It represents a spacious paired enlargement of the nasal
cavity between the skin cover and skull and nasal cartilage longitudi-
nally divided for the most part by an elastic mucous septum. In the
normal (collapsed) state, it is comparatively less recognizable because it
is externally bound on the nose bridge by a transverse fold and anteri-
orly [it] forms a pouch of the rostrum hanging above the mouth opening.
When the animal is in a state of excitation, this saclike cavity becomes
highly inflated under the pressure of air generated by the closure of the
nostrils and stands out prominently above the anterior portion of the
snout in the form of a high and thick semicircular balloon with a length
up to 30 cm and height up to 25 cm (Fig. 207). In addition to inflating
this subcutaneous sac, the males are also capable of blowing a typical
balloon out through either nostril. Such a balloon is formed by inflating
with exhaled air the elastic longitudinal mucous membrane separating
the saclike cavity.!*!

The hair coat of adults is distinctly dichromatic. Dark spots of
extremely variable size and shape are scattered quite haphazardly on a
light gray background (sometimes darker on the dorsal than ventral side).
These spots are slightly angular and quite often of fanciful shapes that
frequently merge when the contours are particularly ornate. The spots in
the midportion of the body are considerably larger, with more complex
outlines, but more sparsely scattered than in the anterior portion, where
they become very small close to the head and look like dabs. The
spottiness on the dorsal surface is somewhat denser than on the ventral
side. The spots are dark brown, often brownish-black or almost black.
The anterior portion of the head, hind flippers, and tail are more deeply
pigmented and often monochromatically dark. The speckles on the fore
flippers are usually quite distinct.

Among males with this type of coloration, much darker animals are
sometimes encountered. The spots in them are so large and numerous,
especially on the upper part of the body, that there is virtually no
space left for the main background; hence such animals appear to be

141 Brondsted, 1931*; Olds, 1950; Trenze, 1950*; Berland, 1958; Mohr, 1958*; Sierts,
1958*; L. Popov, 1960; К.К. Chapskii.

396

397

527

Fig. 206. Skull of the hooded seal, Cystophora cristata (figure by N.N. Kondakov).

Fig. 207. Schematic structure of the nasal cavity of the male hooded seal,

Cystophora cristata, at rest (A) and in a state of excitation, filled with air in one

of the paired chambers (B), causing extension and inflation of the elastic nasal

septum (С). 1—posterior portion of the nasal cavity; 2—eye; 3—cartilaginous

section of the nasal septum; 4—elastic nasal septum; 5—anterior portion of nasal
cavity; 6—nostrils; 7—inflated balloon (after Berland, 1966).

a monochromatic grayish-black from a distance. The females are practi-
cally indistinguishable from males in coloration though there are refer-
ences that the main background among females is lighter and the dark
spots more contrasting (Collett, 1911-1912).

The age-related color changes of the skin are significant. The first
coat with which the pup is born is characterized by the total absence

396

397

528

of spots and sharp color differences between the ventral and dorsal
surfaces. The upper part of the body from the tip of the nose to the
tail is a monochromatic bluish-gray with a bluish-olive tinge. This col-
oration changes sharply on the flanks to almost total white on the lower
side of the body. The color of the inner and lower sides of the flip-
pers is similar. The boundary between the dark and light colors runs
from the lower margin of the nostrils 1.5-2 cm below the eyes almost
through the ear opening or slightly above it, diverging onto the sides of
the [shoulder] blades, and forms a long narrow projection in the mid-
portion of the outer surface of the fore flippers. Posterior to them, the
boundary extends along the flanks of the body (above the axillae) and
reaches, almost without flexure, the hind flippers, terminating in their
upper part, and onto the tail.

Such an unusual coloration of pups is seen in the last stages of
embryonic growth, a few weeks before birth (Reinhardt, 1865*).

Another characteristic feature of the hair coat of the newborn is that
it is not typical of seal pups born on ice floes, 1.е., soft, wavy, and very
dense; instead the embryonic coat is short, rather dense, and somewhat
stiff. This type of coat is seen among other pagophilic seals after the
embryonic coat has been shed, 1.е., the coat of the newborn corresponds
to the second coat of other seals. The embryonic coat is seen in the later
stages of fetal growth but is shed in the womb itself sometime before
birth. The discarded embryonic hair, in the form of dense matted feit
and round disklike clumps, is often detected in the amniotic fluid (Olds,
1950; L. Popov, 1959; Yakovenko, 1959; V.K. Shepeleva). The mech-
anism of matting of. the embryonic hair is not wholly clear but this is
hardly due to fetal movements. It is more probable that the sloughed hair
is swallowed by the fetus and becomes rolled into clumps during passage
through the intestine under the influence of peristalsis and during excre-
tion (Mansfield, 1963). The hair clumps consist of fairly homogeneous
light gray hairs, thus revealing the color of the embryonic coat.

The hooded seal is among the largest of the true seals. The body
length along the dorsal surface (Lc) of adult males from the Greenland
Sea measures 210-250 cm, the largest reaching almost 280 cm; the body
length of most adult females is 180-200 cm but some measure almost
227 cm. The average length of males is 235.7 cm, of females almost
195 cm (L.A. Popov, 1959, 1961).!42

12 According to Degerbol and Freuchen (1935), the largest males reach a length of
350 - 400 cm, which is rather doubtful (the method of measuring the length was not specified;
it was probably with the hind flippers). According to Mansfield (1963), the average length
of males from the western Atlantic is 233.7 cm and that of females 203.2 cm.

398

529

The weight of adult males (length along the dorsal surface Lc
225 - 240 cm) is 260-300 kg; adult females (200-215 cm long), however,
weight 145-160 kg (L. Popov, 1959, 1961). The average weight of
Canadian-Newfoundland male hooded seals is considered as 317.5 kg
(Mansfield, 1963). A large male measuring (Lc) 275 cm long caught on
October 31, 1954 on Faeroe Islands weighed 375 kg (Ehlers, Sierts, and
Mohr, 1958).

The condylobasal length of the skull in adult males (Fig. 206) varies
from 241.5 to 293, 0, average 271.5 mm; in females 218.0 to 238.0, average
225.3 mm; zygomatic width in adult males 158.0-223.0 mm, females
155.0-176.0 mm; mastoid width in males 151.0-171.0 mm, females
144.0- 156.0 mm; length of upper tooth row in males 58.0-85.0 mm,
females 50.0-65.0 mm (L. Popov, 1959, 1961).

The os penis in adult males is 20.5-21.0 cm long, 2.5 -2.6 cm wide,
2.0 cm thick, and weighs 32-38 в (Mohr, 1958*). (К. Ch.)

Taxonomy
Only species of the genus.
Geographic Distribution

Predominantly the pelagic regions of cold streams along the southern
fringe of the arctic zone and its adjoining northern sections of the boreal
strip of the Atlantic.

Geographic Range in the USSR

Only the extreme northeastern branch of distribution extends to the ter-
ritorial waters of the USSR, not beyond the northern sections of the
White Sea covering its inlet and neck sections. Stray hooded seals are
sometimes encountered here; family groups are rarer. These are more in
the nature of stray transgressions of mothers with pups than regular vis-
its. The hooded seals transgress into these regions possibly together with
herds of harp seals migrating to breeding sites. Such instances, though
not arising every year, are well known to the White Sea hunters who
sometimes sight and catch a few animals while hunting for harp seals.

Geographic Range outside the USSR

The southern distribution of the range passes slightly south of the winter
boundary of drifting ice floes from the coastal zone of America in the
region of the Gulf of St. Lawrence and Newfoundland Island, encom-
passing the extensive pelagic expanse to the south and east of the latter
and extends (with some bend toward Davis Strait) toward the waters

398

530

of southern Greenland. Encircling its southern extremity, it runs farther
northeast, covering Denmark Strait, and extends north and northeast and
especially east of Iceland through the Greenland Sea and northern part
of the Norwegian Sea, then descends close to the Scandinavian Penin-
sula considerably below the boundary of winter drifting ice floes. Along
their fringes and receding from the peninsula to the south, evidently up
to the coastal belt, the boundary extends to the southwestern portion of
the Barents Sea to the threshold of the White Sea into its northern part.
The northern boundary of distribution is the fringe of dense accumu-
lations of perennial arctic pack ice between Spitsbergen and Greenland
and evidently the same latitude in the straits between western Greenland
and the eastern fringe of the Canadian archipelago (Fig. 208).

Within these limits of the main distribution, the populations of the
hooded seal are unevenly distributed. Associated in distribution with the
fringes of fairly sparse drifting ice floes, the hooded seals in the course
of their annual cycle drift alternately north and south, partly due to the
seasonal variations in the position of the ice floe fringes.

The southern boundary of distribution in the winter extends from
the southwest, roughly from 45° N lat. at Newfoundland, to the north-
east up to the Polar Circle (perhaps slightly south along the Norwegian
coasts). In summer, on the contrary, it shifts into a more arctic position,

1000 2000 3000 4000 5000 km
| | jee ie

Fig. 208. Distribution of the hooded seal. 1—breeding zone; 2—molting

zone; 3—probable courses of migration; 4—including with pups (after

Rasmussen, 1960, with additional information); 5—distribution boundaries

including probable boundaries (broken line). Dots in the White Sea represent
sites of transgression into the USSR waters (К.К. СварзКи).

399

aye)

toward 60° in southern Greenland and Labrador to the Polar Circle in
the northeastern regions of Iceland and in the much higher latitudes
toward Spitsbergen and the western part of the Barents Sea. Some pop-
ulations migrate in the course of the year from some sections of the
distribution to others in search of food or sites for breeding, molting,
and wintering (see “Seasonal Migrations and Transgressions”’). The pas-
sive migrations caused by these factors and the active migrations in a
relatively short period sharply modify the boundaries of distribution. This
reduces the overall zone of distribution into a tentative concept, defined
as the maximum distribution range. Thus the distribution is extremely
dynamic.

The distribution can be described in greater detail as follows: its
western boundary could be drawn along the Atlantic coast of Canada
to north of Nova Scotia, encompassing the Gulf of St. Lawrence and
the extensive zone of open waters of the Atlantic around Newfoundland,
along the coast of Labrador to Hudson Bay, which is intersected by it
in the westernmost section, and farther north along the entire coast of
Baffin Land with almost the whole Strait of Lancaster and the Admi-
ralty Strait opening into it included. Even more northward, the distribu-
tion includes the entire eastern rim of Baffin Land, Smith Strait, Kane
Basin, and Robson Bay. This represents the northern limit of distribution
between Canada and Greenland. Subsequently, following the coastline
of western Greenland, the boundary descends to its southern extremity.
Thus Baffin Bay and Davis Strait fall wholly in the distribution. Along
the eastern coast of Greenland, the boundary runs from Cape Farewell
roughly up to 75° М lat., is separated there from the land usually by
broken ice floes, tends northeast up to Spitsbergen, descends along its
western side and runs east into the Barents Sea reaching 30° (Nansen,
1924) and even 50° E long. (L.A. Popov, 1960). From this extreme east-
ern limit, the distribution turns southwest (White Sea) and later, entering
the southern boundary traces back in the direction described above.

Since breeding among hooded seals is largely localized, some exten-
sion of their spatial division is possible. Apparently, there are three fairly
isolated geographic groups, which vary in population and importance.
Two of these groups inhabit the western part of the range from the Gulf
of St. Lawrence and Labrador to the northern region of Davis Strait
and southern Greenland in the east to Denmark Strait. In the breeding
period the hooded seals of these groups are concentrated in two regions:
on both sides of Newfoundland Island, mostly north and northeast of it,

43 п compiling the geographic distribution, the data of @ritsland (1959), Rasmussen
(1960), Mansfield (1963), and @ynes (1964) were used.

400

532

and to а lesser extent westward in the Gulf of St. Lawrence. The third
population is distributed in the northeastern part of the range on the ice
floe edges between Denmark Strait and Spitsbergen. Jan Mayen serves
as the region of whelping for this group (for more details, see “Seasonal
Migrations and Transgressions’’). (K.Ch.)

Geographic Variation

Not studied.
Biology

Population. The population of the hooded seal has yet to be precisely
determined. At the end of the 1950s, the world reserves were put at
300,000 - 500,000 (Scheffer, 1958). Aerial surveys were made in 1959 and
1960 in Denmark Strait. The total population of the species was put at
500,000 (Gritsland, 1960). Some higher estimates, also assumptions, put
it at 500,000 - 700,000 (Rasmussen, 1960).

The long-term steady hunting in Denmark Strait could serve as a
rough index of seal reserves. On average, 62,500 hooded seals were
caught there annually from the 1870s. Only a very large herd of hun-
dreds of thousands could have sustained such large catches. Considering
this phenomenon, a figure of 500,000 seals was cited even in later years
(Chapskii, 1966).

Habitat. Mostly similar to that of the harp seal with which the hooded
seal shares much in ecology. Both species are confined to drifting ice
floes, mainly to their fringe zones. But while the herp seal prefers com-
paratively young and low ice floes, the hooded seal selects contrarily
very stable perennial pack ice for reproduction. The sites of the hooded
seal invariably have frequent waterways since this seal does not make air
holes in the ice floe (Nansen, 1924; L. Popov, 1960). As it avoids dense
accumulations of ice floes, the hooded seal is not encountered on shore
ice. It usually does not enter the deeply protruding bays and straits or
narrow straits between islands packed with ice floes. Ц does not enter
deep into the Canadian archipelago, transgressing only into the more
southern broad straits opening into Davis Strait and Baffin Bay. For the
same reason, it does not approach coasts densely packed with ice floes,
remaining confined to their outer fringes.

During the breeding period hooded seals are usually encountered
inside the ice massif (which the hunters’ schooners cannot always pene-
trate), on stable arctic pack ice, and around very young ice floes alternat-
ing with open water pools. The accumulation of ice floes with open water

401

583

pools is particularly characteristic of regions with a slow, often circular
drift at the confluence of opposing currents. Jan Mayen constitutes such
a region of relatively stable, coarsely and finely broken ice floes. Here, as
a result of the interactions between the cold eastern Greenland streams
with the northwestern branch of the Gulf Stream, favorable conditions
arise for feeding as well as for prolonged residence on the drifting ice
floes. The hooded seals find very similar conditions in Denmark Strait,
which represents their molting region.

To a much greater extent than the harp seal, the hooded seal is a
pelagic inhabitant of open expanses and is seen close to coastlines only
in the period of migrations; at this time it prefers the coastal strip of
southern, mainly southwestern Greenland. In winter and early spring it
is dispersed sparsely in pelagic regions, far from ice fringes, especially
in the Norwegian Sea (see “Seasonal Migrations and Transgressions’’).
In this very period of winter fattening it is also confined to the western
Atlantic in the vicinity of the Newfoundland coasts and evidently close
to Iceland (Wynes, 1964).

Like other pagophilic seals, the hooded seal requires ice floes for
reproduction and molt. This need is also felt when the animals are
not fat enough and hence suffer excessive heat loss in water. Ice floes
play some role also as a substratum for resting and provide a means
of escape from aquatic enemies. The affinity for a strip of ice floes is
evidently caused by food conditions, promoted by the polar front zone
in general, and the available strip of spring-summer ice floe fringes in
particular.

Food. This aspect has not been studied. Only very general refer-
ences are available regarding specialization of the dental apparatus for
grasping and tearing large adible objects including fish (Nansen, 1924;
L.A. Popov, 1961). These are essentially the sea perch, flounder, cod,
and cephalopod mollusks; the seasonal changes in the distribution of
the hooded seal are probably related to the migration of cod. In the
period of suckling pups and even during molt, adults and others in the

. transitional age group do not obviously cease to feed and go out to sea

quite often. Their stomachs revealed food remnants, in particular squids
(L.A. Popov, 1959; М.Уа. Yakovenko). Nevertheless, feeding in these
periods is very poor and highly irregular. During lactation and molt the
animals become quite emaciated (Rasmussen, 1960); intensive feeding
occurs after mating, before congregation in the molting region, and also
at the end of molt.

On transition to independent feeding, the young seals initially feed
on amphipods, small squids, and other edible objects. An adult male in a

534

zoological garden was fed on herring, whiting (hake), cod, coalfish [pol-
lack], flounder, witch flounder, turbot, and sea perch; this seal refused
to eat the latter, however, when a choice was available. Initially, when
live fish were fed, it consumed eel, carp, and tench. There was a distinct
preference for herring, with up to a hundred consumed daily (Ehlers,
Sierts, and Mohr, 1958).

Home range. It is practically impossible to establish the home range
of such a widely migrating and wandering animal as the hooded seal. The
area of drifting ice floes which the seals select for reproduction is also
not amenable to precise calculations, albeit the animals are confined
to the same icy regions at this time. Much depends on the actual ice
conditions of a given year, type of drifting ice floes, features of approach,
and the initial disposition of the animals. Moreover, the density of animal
disposition varies even during the short duration of lactation under the
influence of ice floe movement.

The animals live in “families” in the regions selected for reproduc-
tion. The whelped female is confined to a definite ice floe with her pup;
the adult male also stays closeby on the same ice floe, forming a mating
pair with the chosen female. Such pairs with suckling pups are disposed
at different distances from each other. Generally, each pair occupies a
separate ice floe. But when the floe is very large, other pairs occupy it
but at distances of tens of meters or sometimes far less. Thus occasion-
ally the hooded seals are so densely disposed at sites of reproduction
that the latter resemble the nursieries of harp seals (Rasmussen, 1954*;
A.V. Potelov).

In general, however, very uneven distribution of the seals is a char-
acteristic feature during the nursery period. In the large concentration
north and northwest of Jan Mayen Island on March 13, 1962, on aver-
age five mating pairs with pups were encountered per mile of traverse.
On March 24 of the same year, slightly eastward, up to 15 “families”
were counted on some ice floes measuring roughly 100 x 100 m; 15
to 25 animals were sighted from a ship on March 26 on the spits of
a coarsely broken ice floe (Khuzin and Potelov, 1963). In general, in
the winter-spring season some mating pairs occupy from 670 m? to
roughly 0.6 km? at reproduction sites. Sometimes such mating pairs
with pups are typically interspersed among masses of harp seals
(R.Sh. Khuzin).

During molt on the ice floes in Denmark Strait the disposition of
hooded seals is denser and more stable. The distance between animals
is just a few tens and sometimes even a few meters. During local move-
ments and more so during migrations, the concept of “home range” is

402

535

no longer applicable due to the wide scattering of the animals and their
uninterrupted movements.

Hideouts and shelters. The hooded seal does not seek hideouts and
shelters on the ice floe or in the snow. As a rule, it does not make air
holes in the ice floe but utilizes the natural openings and open water
pools for emerging onto and exiting from the ice. When, however, the
open water pools have a thin ice cover, the seal can pierce it for res-
piration. In 1956, a hooded seal under observation pierced a young ice
cover 4.5 cm thick with its head, thrust up out of the water, surveyed
the surroundings, filled its lungs, and submerged never to be seen again.
Perhaps the seal managed to breathe at other places in a similar manner
as Open water was nowhere visible, the entire surface was frozen and
calm, and frost prevailed (M.Ya. Yakovenko).

Daily activity and behavior. These aspects have not been studied.
During the molting season, the animals rest long periods on the ice
floes interspersed with brief spells of submergence in water. Males and
females enter water even more often during the period of suckling the
pups. The animals spend time predominantly or exclusively in water for
the purpose of feeding and moving about.

Seasonal migrations and transgressions. Hooded seals are among the
widely migrating seals. Although not much is known about their migra-
tions, their general characteristics have been established. During the
breeding season the productive males and mothers are grouped in two
widely separated regions. One group concentrates in the southwestern
part of the range, on the southern extremity of Labrador opposite the
Strait of Belle Isle and the northernmost extremity of Newfoundland. A
small population locates in the Gulf of St. Lawrence. The other region
occurs on the eastern boundary of the range, northwest of Jan Mayen
Island.

The hooded seals reproducing in the westernmost section of the dis-
tribution, having completed this biological cycle by early April, abandon
the ice floes for sites abounding in food. They first travel along the edges
of Labrador north toward Davis Strait and later, turning east, approach
the western Greenland coasts rich in fish and feed for sometime right
at the banks as also in the coastal regions of southwestern Greenland.
Further, some groups of seals move higher and approach Greenland
at latitudes 65 to 66° and others (the majority apparently) remain in
the feeding grounds, not moving north of 60 to 61° N lat. and are seen
roughly at the same time in May on the southern coasts of Greenland.

Having recovered their fat reserves to some extent by the end of
May to early June, the hooded seals reach the southernmost tip of
Greenland, move round Cape Farewell, and continue their course along

403

536

eastern Greenland (instances are known of hooded seals marked near
Newfoundland being recovered at Cape Farewell).

Later, the seals migrate farther northeast along the ice edges to
Denmark Strait, the main (though not the only) site where hooded seals
gather in large numbers and remain for molting in June and July.

In its first year, the juvenile does not undertake this migration east-
ward; it migrates evidently farther north into Baffin Bay and remains
there until August.

Having completed molt, the adult hooded seals begin to leave
Denmark Strait in the first half of July and scattering widely, return by
the same route to the southern tip of Greenland, and passing it repeat
their journey in a reverse direction. Roughly from mid-July through mid-
August, hooded seals emaciated during molt are caught on the coasts of
southwestern Greenland. Returning to their birth place, the Canadian-
Newfoundland population again visits the western Greenland fish banks
and feeds well. These seals then pass through Davis Strait and are seen
on the Labrador coasts in early autumn, proceeding south into the region
of the Great Newfoundland banks for feeding. They return north in
February to the whelping zone and concentrate again on ice floes before
commencing a new annual cycle.!*4

The populations reproducing in the region of Jan Mayen Island,
after whelping and mating also scatter quite extensively but their
migration does not attain the scale that is characteristic of the
Canadian-Newfoundland populations. The pups abandoned by their
mothers migrate initially passively with the moving ice fringes and are
encountered at April end to early May as highly scattered solitary animals
in the fringe zones from 69 to 76° N lat. (L.A. Popov, 1960; and others).

From April to June the adults are scattered even more widely among
thin ice floes and beyond the fringe zones. They are then capable of dis-
tant migrations in search of food. These migrations sometimes extend
right up to the White Sea. A large collection of hooded seals, some-
what like a nursery in the Greenland Sea (on Jan Mayen Island), was
encountered around May 10, 1962 in the middle of the Barents Sea
(73° 38’ -74°00'N lat. and 24°28’ -35° 20/E long.) (V.A. Potelov). Much
of the population nevertheless tends southwest into Denmark Strait
where it molts in June. The rest is scattered on the fringes of the Green-
land Sea to north of Jan Mayen roughly up to 75° N lat. and molts there
on pack ice (Wolleback, 1907; L. Popov, 1959; R.Sh. Khuzin).

144 According to Allen (1880), Nansen (1924, 1939), Bartlett (1928*, 1929), @ritsland
(1959), Rasmussen (1960), Mansfield (1963), and others.

404

557

At the end of the molting period, the Jan Mayen hooded seals are
again scattered over an extremely wide expanse whose boundaries were
only recently deciphered. It was assumed that they were confined to the
high latitudes and appeared near the mainland only very rarely. There
were frequent reports from fishermen using nylon nets to catch cod and
halibut in the Norwegian Sea, of hooded seals being caught in them at
Olesund, northern Tronnelag, Vesteralen region, and other places along
the Norwegian coast. Hooded seals were seen there even in winter and
were very Common in spring. Such instances were particularly numerous
in the spring of 1959 (@ynes, 1964). These facts compel us to recognize
that, in winter and even in spring, the seals of this species stay regularly
at relatively low latitudes not only in pelagic zones, but also close to the
coasts, entering even into the Norwegian fjords.

Instances have been recorded of scattered solitary animals at differ-
ent points on the coasts of Scotland and England; one find has been
reported even from Biscay coast of France. In most cases these were
young animals, some of which had only just been born. Along the Amer-
ican coast of the Atlantic Ocean south of Nova Scotia, instances have
been reported even recently of the appearance of solitary animals in the
region of New Jersey and on the coast of Massachusetts; a dead female
was found right on the coast of Florida in the winter of 1916 (Miller,
LSI):

The transgression of a hooded seal into the eastern zone of the
Yenisey estuary is wholly exceptional in the matter of distance involved
and is inexplicable (N. Smirnov, 1908, 1929).

Reproduction. The period of mating sets in immediately after the
completion of lactation and extends roughly for two weeks. Even by
around April 10, ovulation has occurred among almost all the mature
females; the bulk of the animals mate in the first week of this month
(L.A. Popov, 1959; Oritsland, 1964).

Precise information is not available about the copulation process
but animals were observed mating in water. The pair slowly floated close
to the surface, periodically thrusting out for respiration. The extremely
large size of the males, larger than the females, suggests that they woo
the females anew each year. Traces of fights in the period of heat have
been detected on the skin of males (Yakovenko, 1959).

The lag in implantation of the blastocyst has been established as 3 to
4.5 months from the time of mating, i.e., up to July end to early August
(Oritsland, 1959, 1964).

The hooded seals whelp in different parts of their range almost
simultaneously, in most cases in mid-March; pups appear only very rarely
at a much later period.

404

538

Fig. 209. Pup of hooded seal with mother. Jan Mayen region (photograph by
A.V. Yablokov).

Twins have not been reported among the females studied but some-
times two pups have been seen near the same female.

Unlike other seals, sexual maturity among hooded seals sets in rel-
atively early; in rare instances, the first ovulation occurs at the age of
two years but in most cases does not result in gestation (Oritsland, 1964;
Yakovenko and Khuzin, 1965*). Even at the age of three years, only
half of the females are pregnant. Only 80% of the 4-year-old females
whelp while 10% of the five-year-olds still remain nonproducers (@rits-
land, 1964). Subsequently (most often from seven to eight years), females
whelp every year (number of barren animals is not over 5%). The ability
to become fertilized and whelp is maintained right up to the end of their
lives, which can be taken as 35-36 years.

The onset of sexual maturity among males occurs somewhat later.
Until three years of age, they are mostly immature.'® At four years
some males evidently become mature but number about 30%. This rises
to 80% among five-year-olds (L. Popov, 1960). Commencing from six
years of age, all males are mature (Khuzin and Potelov, 1963) though,

145 According to Yakovenko (1959), 3-year-old males (with four layers on the cross section
of the canine) are already mature and evidently come into heat since the skin of one such
animal revealed wounds inflicted in a fight.

405

539

according to other authors (L.A. Popov, 1960), immature animals are
still encountered at this age, their proportion reaching 13%.

The sex ratio among the newborn is 1:1 as observed from the numer-
ical ratio between the males and females in the molting rookeries. Of
1,125 pups examined, 49.2% were males (@ritsland, 1964).

Growth, development, and molt. The length of the newborn along
the dorsal surface, judging from the size of the fetuses measured in
the period of whelping (about mid-March and even in the first few
days of April), varies from 79 to 112 cm, on average 100 cm (Collett,
1911-1912; Nansen, 1924146 L.A. Popov, 1959; Potelov and Yablokov,
1967*; V.A. Potelov).

The newborn weighs 12-27 kg, on average 19.3 kg (V.A. Potelov).
Other figures too have been cited (23 - 30 kg) but the pups were probably
already full grown and better fed.!*’

After 10-15 days of lactation, pups (Fig. 210) grow to 117 cm (Lc)
on average while the weight increases up to 37.5 kg on average, the
range being 23-55 kg (V.A. Potelov). The fat content of the milk has
been reported as 43% (Khuzin and Yablokov, 1963) and on average 60%
(with a protein content of 4-9%) (V.A. Potelov).

Toward the end of the period of lactation, the thickness of the fat
layer in the skin increases to 4-5 cm from 1-1.5 cm at birth (L. Popov,
1961; Shepeleva, 1971); of this, the thickness of the fatty tissue is
3.5-4.0 mm.

The subsequent overall growth of the animal proceeds with the same
intensity: the body length of yearlings measured along the dorsal sur-
face (Lc) increases to 135-150 cm, up to 160-170 cm at two years,
and 170-185 cm at three years of age (Yakovenko, 1959). According to
others (Rasmussen, 1960), the body length measured along a straight line
(Lcv) among yearlings of males and females is identical at an average of
133 cm!48; sex begins to exert a pull in the two-year-olds; the average
length of males then is 155 cm, of females 152 cm. At three years of
age, the males measure on average up to 172 cm, females up to 164 cm.
The same intensity of growth continues up to 10 years of age (when the
males grow to an average of up to 218 cm and females up to 193 cm).

146 Nansen and Collett indicate the length of the newborn as 90 cm. The measurement
was evidently along a straight line, i.e., Lev.

147 The much older data of 3-5 kg (Ognev, 1935) or 6-8 kg (М.Р. Vinogradov, 1949)
are undoubtedly erroneous.

148Треге is a view that sex-related differences in body length exist even at the time of
birth and increase slightly during lactation (Potelov and Yablokov, 1967*) but the data in
support of this view are not very convincing.

406

540

Fig. 210. Pup of hooded seal. Greenland sea (photograph by V.K. Shepeleva).

Growth continues, although at a slower rate, up to at least 20 years of
age.

The age-related changes in color of the hair coat reveal some char-
acteristic features. The original, neonatal color remains unchanged for a
year. After the first molt (i.e., at the beginning of the second year), the
first spots, not yet vividly pigmented, appear on the anterior portion of
the back and partly on the neck (Nansen, 1924; Yakovenko, 1959). In the
course of a year their number increases and these spots extend over much
of the back and are also seen on the chest but not in the belly region. In
the first few months of the fourth year (after the third molt), the main
background on the upper side of the body turns olive-gray and is cov-
ered with the innumerable dark brown spots characteristic of the color of
adult animals. The underside of the body also acquires a brownish tinge
and is speckled with spots scattered haphazardly. The snout becomes
dark. On attaining the age of seven years, the main background turns
lighter in color while the spots become darker (Yakovenko, 1959). This
skin pattern evidently remains unchanged until the death of the animal.
According to some authors, the immature animals of both sexes are dis-
tinguished from the adults only in the number of spots, which increase
far more with age; the main background, however, remains uniformly
light in color (L. Popov, 1959b).

406

407

541

The nasal skin зас is not manifest at all among the newborn males in
the first year. In the second year it becomes prominent as a “tiny cushion”
slightly enlarging when the animal is excited (L. Popov, 1959b). The sac
as such is manifest in the fourth year (Yakovenko, 1959). A real “hood”
was seen in a five-year-old male (age established based on marking)
(Rasmussen, 1957).

The hooded seal begins to molt three months after the end of the
whelping period. Denmark Strait is the main site of congregation of
hooded seals of all ages (except under-yearlings) for molting; they gather
there from around June 10 and leave toward July 20. Thus the molting
period extends for 1.5 months. The molting rookeries of the hooded seal
are less dense than those of the harp seal but are far denser than the
nurseries of the former. There are no distinct age- or sex-related differ-
ences whatsoever among molting hooded seals: the animals are disposed
alternately in close proximity. Outside Denmark Strait, no other such
region of en masse collection of hooded seals in the molting period is
known, but small numbers of molting hooded seals are seen in Jan Mayen
and Spitsbergen regions and generally almost all along the fringes of ice
floes in the Greenland Sea (Qritsland, 1964; R.Sh. Khuzin).

Enemies, diseases, parasites, mortality, competitors, and population
dynamics. The young seal spends its early life on ice floes in an almost
sterile icy environment under the protection of adults without exposure
to any factor of natural mortality to which this population is sensitive.
Later, when the juvenile enters the water and fends for itself on the
fringe of ice floes, it may fall prey to the Greenland shark or killer
whale. The latter possibly attacks rather frequently even the adult seals.
Among other enemies is the polar bear but the hooded seal is much less
threatened by this animal than the other arctic seals; furthermore, the
population of this predator in the Jan Mayen region is not much. In July,
1956, the stomach of a polar bear killed in the Greenland Sea revealed
the remains of a hooded seal pup (M.Ya. Yakovenko).

This seal is parasitized by 15 species and two larval forms of
helminths.!*? From among the trematodes, Metorchis albidus infects
the gall bladder and bile ducts of the liver, Opisthorchis tenuicollis
and Pseudamphistomum truncatum the bile ducts of the liver; from
among the cestodes, Anophryocephalus anophrys and Diplogonoporus

49 The review of helminths was compiled by scientists of the Department of Zoology of
Crimean State University based on the following data: Rudolphi (1819), Siebold (1848*),
Brown (1893*), Price (1932), Berland (1963), Skryabin (1950*), Delyamure (1955),
Delyamure, A. Skryabin, and Treshchev (1965*), Delyamure, Skryabin, and Alekseev
(1964*), Treshchev, Zavaleeva, and Potelov (1967), and others.

407

542

tetrapterus infect the duodenum and the small intestine while
Diphyllobothrium latum, D. pterocephalum, Pyramicocephalus phocarum,
and Diphyllobothriidae g. sp. infect only the small intestine; from
among the nematodes, Contracaecum osculatum, Terranova decipiens,
Phocascaris phocae, Ph. cystophorae, and Anisakidae g. sp. parasitize
the stomach and small intestine while Skrjabinaria spirocauda parasitizes
the heart, lungs, and blood vessels; from among the acanthocephalans,
Corynosoma strumosum and C. semerme infect only the intestine.

The dissection of 293 hooded seals (Treshchev et al, 1967)
established a high rate of infectivity; helminths were detected in 61.4%
animals. If, however, the newborn (96 pups) are excluded from the
total number of dissected animals and only the adults (197 seals) are
taken into consideration, the percentage of infectivity rises to 91.4.
The most severely infected seals were aged 1 to 12 years. The most
frequently infected organ was the small intestine (in 58.02% animals);
less frequently infected were the stomach (29.35%), duodenum (27.9%),
and even less the heart (3.4%), lungs (1.02%), and liver (0.68%).

Parasites sometimes cause severe diseases. The nematode Ph.
cystophorae is capable of perforating the intestinal wall of the host. 5.

Fig. 211. Head of an adult female hooded seal (photograph by A.V. Yablokov).

408

543

spirocauda, a parasite of the heart, probably kills the hooded seals at six
to seven years of age (Delyamure and Treshchev, 1966).

The seal louse Echinophthirius horridus parasitizes the skin and the
seal mite Halarachne halichoeri the nasal cavity.

Competition in the feeding grounds can arise only from the harp seal.
In practice, however, there is no such competition as the food reserves
suffice to meet the needs of both species; furthermore, the hooded seal,
more than the harp seal, lives on deep-water nektonic life. Had there
been an acute competition, the two species would not probably have
formed nurseries in the same regions. Most of the feeding grounds of
these two species are nonetheless distinct.

Some idea of the population reduction can be gleaned from hunt-
ing statistics, especially a perceptible increase in younger animals in the
herd, i.e., a relative reduction in the number of mature animals and
an increased percentage of young ones. In 1956, the young constituted
27.5%, which rose to 34% in 1957 and to 49.5% in 1958. It was consid-
ered that the killing was exceptionally high in the 1950s and the state
of the herd at the beginning of the 1960s caused anxiety (Rasmussen,
1960), necessitating a ban on hunting in Denmark Strait. In turn, during
the years of World Wars I and П the forced cessation of hunting using
ships promoted the restoration of reserves. The reduction at present is,
however, quite perceptible. Hunting is the main factor responsible for
the population dynamics of the hooded seal.

Field characteristics. The under-yearlings are easily distinguished by
the two-toned coloration of the skin, 1.е., bluish-gray on top and white
below. The adult animals are recognized by their large size and con-
trasting spottiness. The hooded seal can readily be distinguished from
the harp seal with transitional (gray spotted) coloration by the former’s
fairly large blackish-brown and dark brown spots often fusing into fanci-
ful patterns on a gray background with densely scattered spots and dabs
in the anterior portion of the body. A positive feature of recognition
is the saclike process on the head of adult males, which is quite often
inflated through the nostrils (Fig. 212, A, B). Distribution in separate
“families” on ice floes during the nursery period is also a useful species
characteristic. (K.Ch.)

Economic Importance

The economic importance of the hooded seal in the world hunting indus-
try is quite high. As an object of hunting using ships in the North
Atlantic, it occupies second place after the harp seal. The main region of
hunting now falls in the international waters near Jan Mayen. Sections

544

1 i

Try yyy

МЫ

РР!
>

409 Fig. 212. Head of an adult male hooded seal. A—with a “cap” slightly inflated

by the nasal cavity (after Mohr, 1963); B—schematic depiction of fully inflated
nasal cavity (“сар”) (figure by К.К. Chapskii).

409

410

409

545

of the “polar front” to the north of Newfoundland are of much less
importance. In both these regions the hooded seal is caught together
with the harp seal. Until recently, hunting was practiced in Denmark
Strait and half a million hooded seals were caught there in the 1860s. In
good hunting seasons at the end of the last and early in this century, 15

to 20 hunting ships caught 40,000 to 57,000 seals. In the region of Jan

Mayen, however, even in the first two decades of this century relatively
few hooded seals were caught. Large congregations were not found in
this area. The catch rose markedly from the 1920s. In 1926, Norwegians
caught 53,000 hooded seals in the expanse from Spitsbergen to Denmark
Strait (Sivertsen, 1941).

In the postwar years (from 1956 through 1958), hunting notably
decreased in Denmark Strait and was totally banned in 1961. In the
region of Jan Mayen, however, it continued at a high level. From 1946
to 1958, it averaged at different times 35,000 - 59,000 - 44,000 seals per
annum. The catch of hooded seals in the Newfoundland area (in which
Norwegians and Canadians hunt) represented about 10% of the hunt in
the Jan Mayen region. In these years the catch averaged 5,700 animals,
mainly pups.

The Soviets took to hunting the hooded seal in 1955 and continued
to do so up to 1967. The ice floes of the Greenland Sea in the region of
Jan Mayen constituted the hunting region. Here the harp seal was the
main target.

The extent of hunting is depicted in Table 29.

Relative to the harp seal, also hunted in this region, the catch of the
hooded seal varied from 10-15% of the total catch in the early years of
hunting to almost 70% in some subsequent years.

There is nothing specifically different in the hunting of hooded seals
using ships compared to harp seals except that the ships have to operate
under more severe icy conditions, often approaching each “family” group
separately. The hunting operations for some reason or the other involve

Table 29. Catch of hooded seal by Soviet hunting ships in the Greenland Sea

(R.Sh. Khuzin)
Year Total catch Pups in total catch
1960 4,575 1,472
1961 11,781 7.101
1962 9,562 3,874
1963 7,303 3,470
1964 8,861 4,706

1965 8,380 5,326

546

great difficulties as well as much time. Hunting itself is not а very com-
plicated affair due to the slower self-preservation reflexes of the animals ©
in the period of reproduction.

The main problem is locating the rookeries since aerial assistance
is not used. Although the hooded seal represents an enticing target (an
adult animal yields more fat than a harp seal, while the skin of a juvenile
hooded seal is highly valued as a commercial fur), it was not specially
sought in the early years of hunting and only whatever came in the way
of a ship was taken. Later, however, hunters laid greater emphasis on
the hooded seal. After World War II, the Norwegian hunters invariably
sought the juveniles so that the catch of pups exceeded that of adults,
amounting to 55.5-86% of the total catch. On average, in the first 13
postwar years the Norwegian hunters killed 70% of the juvenile hooded
seals available in the Jan Mayen region. The fashion for seal skin over-
coats played no mean role in this regard. The hair coat of the pups
is distinguished by denseness, moderate luxuriance, excellent color and
luster, while the skin is quite durable.

The skin with the subcutaneous fat layer when removed by the
hunters (without the flippers) weighs 5 to 11.5 kg (average 9.1) in the
case of a newborn but averages 20 kg at the end of lactation. The weight
of an adult skin with the fat in early spring (during the breeding season)
in the case of males (length along the dorsal surface, Lc, 225-260 cm)
varies from 45 to 60 kg. The weight of the blubber in males constitutes
roughly 60% of the total weight and in females impoverished during lac-
tation 30-40% (L. Popov, 1961). The meat is used as feed in fur animal
farms.

To implement rational exploitation of the resources of the Jan
Mayen hunting region and to ensure conservation at a stable level for a
long time, agreements are imperative between the countries involved in
hunting the hooded seal.

Efforts in the field of research should be directed primarily toward
an accurate census of the pups of the hooded seal; a comprehensive
study of its ecology, including intensity of reproduction; migration char-
acteristics; degree of isolation of the Jan Mayen herd from the western
Atlantic (Canadian) populations; etc. The major task is to establish ratio-
nal norms of permissible kill to prevent the exhaustion of hooded seal
resources.

Some measures restricting hunting have already been implemented
in Norway and the USSR. These measures provide: (1) restricted hunt-
ing in the breeding season in the Jan Mayen region (commencement
of hunting not earlier than March 20 and its cessation not later than

547

May 5); (2) banning the reentry of a ship for hunting in the same sea-
son; (3) banning hunting in Denmark Strait; and (4) banning hunting in
summer оп drifting ice floes north of Jan Mayen Island. (K.Ch.)

411 ORDER OF WHALES

Order CETACEA Brisson, 1762

413

COHORT OF WHALES
COHORT Mutica

ORDER OF WHALES
Order CETACEA Brisson, 1762

These are highly specialized mammals, totally adapted to aquatic life.

Dimensions vary from moderate to large and these animals are the
largest among mammals. Body length and weight range from 1.1 m and
30 to 300 kg in some large dolphins to 33 m and 100 to 120 tons in blue
whales (Fig. 213).

The body is streamlined, spindle-shaped or teardrop-shaped, elon-
gated or somewhat shortened: Such a shape offers the least resistance
to swimming in water (Fig. 214). In common dolphins the body is better
streamlined than in such fast-swimming fish as gray mullets and boni-
tos (Shuleikin, Luk’yanova, and Stas’, 1937). Resistance to swimming in
water is also reduced by the reduction of ear pinnae and the placement
of mammary glands and teats in special pouches in females and the penis
in a special slit in males.

The forelimbs are modified in the form of fin-shaped flippers, which
essentially serve as rudders. The hind limbs are totally reduced. At the
end of the laterally flattened caudal crest there are paired horizontal
caudal flukes without a skeleton. The caudal section of the trunk serves
as the main locomotory organ. In most species an unpaired dorsal fin
without a skeleton performs the role of a stabilizer while swimming. The
head is often massive, sometimes about one-fifth of the body length, and
is usually fairly elongated. It terminates obtusely or acutely, or has an
extended rostrum, i.e., the so-called “beak”. The head in almost all these
animals merges without a visible neck into the trunk, which gradually
narrows with no sharp boundaries into the caudal crest.

Whales do not have a compact hair coat. Baleen whales have some
stray bristles on the snout, typically structured like the whiskers of land
mammals. These hairs evidently play a specific role in searching for

414 Fig. 213. Comparative size of whales (figure by М.М. Kondakov). A—toothed
whales: 1—common dolphin, 2—pilot whale, 3—killer whale, 4—Baird’s beaked
whale, 5—female sperm whale, 6—male sperm whale; B—baleen whales: 7—blue
whale, 8—fin whale, 9—sei whale, 10—humpback whale, 11—Minke whale,

12—gray whale, 13—bowhead whale.

massive plankton collections (up to 400 nerve tips present in the follicle
of one such hair). In the adult state, among the toothed whales, only the
river dolphins possess whiskers: the Ganges River dolphin—Platanista
gangetica and the Amazon dolphin—Jnia geoffrensis living in muddy
river water. In most of the other toothed whales (except for the white
whale and narwhal), whiskers are present only in the embryo.

414 Cutaneous glands are lacking. The skin consists of an epidermis, der-
mis, and a very thick layer of subcutaneous fatty cellular tissue [blubber]
with no distinct demarcation between the last two layers (Fig. 215). The

553

415 Fig. 214. Characteristic body form of whales (figure by М.М. Kondakov). 1—right
whale; 2 and 3—rorquals; 4 and 5—dolphins; 6—white whale; 7 and 8—sperm
whales.

554

)

ме

eR
S =

SN

416 Fig. 215. Structure of the skin of whales according to У.Е. Sokolov.
1—epidermis; 2—dermal papillae; 3—dermis; 4—subcutaneous fatty cellular
tissue; S-—subcutaneous musculature; 6—bundles of collagen fibers; 7—fat cells.

epidermis in the body sections exposed to the maximum resistance of
water while swimming (anterior margins of the fins and anterior part of
the head) attains maximum thickness. The inner surface of the epidermis
has numerous cells in which dermal outgrowths (papillae) grow vertically
upward toward the kin surface. The pigmentation of the epidermis deter-
mines the body color of whales. At the boundary of the epidermis and
dermis, pigment cells—melanophores!—are visible. The color of some

‘Numerous white or weakly pigmented spots (scars) of varying size are usually found
on the skin of whales. These are scars from bites during fights, attacks by sea lamprey,
parasitization by some Protozoa, crustaceans, etc.

555

cetaceans has been correlated with age-related variations (for example,
among white whales and narwhals). The dermis consists of a thin layer of

416 dense intertwined bundles of collagenous fibers and papillae rising from
this layer into the epidermal cells. In the subcutaneous fatty cellular tis-
sue, Straight fascicles of collagenous fibers are considerably larger than
in the dermis and widely separated from each other, and the entire space
between them is filled with accumulations of fat cells. The thickness of
the subcutaneous cellular tissue varies in different seasons: for example,
among female fin whales, it is 7.8 cm in December and grows to 12.7 cm
by March (Slijper, 1936).

The peculiar structure of the skin of whales evidently promotes
development of a laminar flow of water around the swimming animal,
with a minimum expenditure of energy when developing high speed.

The caudal flukes (Fig. 216) under the skin cover consist of three lay-
ers: one deep and two superficial. The superficial layers are formed of the
tendons of m. longissimus and m. hypaxialis lateralis? (Narkhov, 1937)
and spread fanlike into the flukes (Felts, 1966). The deep layer consists

С р Е

417 Fig. 216. Figure showing structure of the caudal flukes in various species of
whales (figure by N.N. Kondakov). A—Atlantic right whale; B—gray whale;
C—fin whale; D—common dolphin; E—sperm whale.

2 According to A.V. Yablokov (1959), the tendons of these muscles do not reach the
caudal flukes.

417

418

556

of a fairly dense bunch of large bundles of collagen fibers between which
groups of fat cells occur. The caudal flukes are capable of minor indepen-
dent movements (Zenkovich, 1952). The internal structure of the dorsal
fin is similar to that of the deep layer of the caudal flukes. In the fins,
along with normal arteries and veins, complex vessels exclusively typical
of whales are seen. Each of these vessels consists of a large artery with a
very thick middle layer (tunica media) surrounded by 10 to 13 very fine
thin-walled veins. The base of the dorsal fin has more than 20 such ves-
sels, which are fewer in number in the caudal flukes and flippers (Tomilin,
1957). These typical vessels are seen in the body parts but differ in struc-
ture; their formation is evidently determined by functional requirements.

The elasticity of the caudal flukes can evidently vary during the move-
ment of the animal due to blood pressure controlled by complex vessels
and the general distribution system present in the principal supply vessel
(Pershin et al., 1970). It is possible that the effect of hydroelasticity in
the caudal flukes has a bearing on the swimming of whales.

Skeletal bones essentially have a typical spongy structure and con-
tain a large amount of fat. Clavicles are lacking (Fig. 217). The sternum
is highly variable in size, shape, and articulation, depending on the num-
ber of ribs attached. The scapula is broad, flabellate [fan-shaped], and
has a poorly developed crest. The humerus is extremely reduced. The
radius and ulna are also highly reduced, flattened, and greatly broadened
(Fig. 218). Flattening of the limb bones is evidently associated with the
limb performing a typically new function similar to that of the aileron
of an airplane wing. In view of the fact that the load on the limb is per-
pendicular to it, the limb is flattened in the same direction (Druzhinin,
1924; Yablokov, 1959). All the bones of the free limb are firmly inter-
connected and sometimes even fused, and with a common integument.
Only the shoulder joint is movably articulate. Nevertheless, antebrachial
muscles are preserved in the majority of whales. More often than among

v 3 BAB Е 56
чопрдурорие оч“ * ^

Fig. 217. Skeleton and body contour of the common dolphin, Delphinus delphis
(figure by N.N. Kondakov).

418

557

Fig. 218. Skeleton of the limbs of whales (schematic). A—bowhead whale; B—fin
whale; C—blue whale; D—common dolphin (figure by N.N. Kondakov).

other mammals, the fusion of the carpals is noticed. The number of digits
is four or five. The number of phalanges in the middle digits is more and
in the outer ones less. The number of phalanges in the embryo is more
compared to the adult, 1.е., they reduce in number during the course of
ontogenesis (Kiikenthal, 1889; Yablokov, 1959).

The hind limbs form in a fairly early embryonic stage but soon dis-
appear. Their rudiments are very rarely preserved. Among adult whales,
only the rudiments of the pelvis arranged in the musculature, in the
form of two bony shafts (fused ilium, ischium, and pubis), are preserved

- of the hind limbs. Their articulation with the vertebral column has dis-

appeared but the muscles of the urogenital system are attached to them
and, obviously for this reason, their total reduction has not taken place
(Yablokov, 1959).

The vertebral column consists of only four sections: cervical, tho-
racic, lumbar, and caudal. Sacral vertebrae are lacking due to the disap-
pearance of the hind limbs. The cervical section is represented by seven
highly reduced vertebrae which may be fused into a single unit or sev-
eral groups. In the rest of the sections, the number of vertebrae varies.
Transverse and spinous processes attain large proportions. Hemal arches,
similar to those in the lower vertebrates, appear in the caudal section.

Е
ИЕ = = —
И: 2727
ПРЕ. 2 ge
i PN

419

418

419

S\N
\

\

He
WN

м

PO ро
М
Ny
Wi

у

Bai
№
oN
-\

а
N
4

Sins
а

\) RTS

Sas

т.е

Fa

\
\
К

Fig. 219. Rudiments of the hind limbs of the sperm whale. A—outer view, В
and C—X-ray pictures (figure by М.М. Kondakov, after А.А. Berzin, 1970).

The number of ribs varies from 10 to 17 pairs. The anterior two to
eight pairs are normally joined to the vertebrae; the posterior ribs have
neither capitulum nor collum and are articulated only to transverse verte-
bral processes. The anterior eight pairs of ribs are joined to the sternum.
The thoracic cage, unlike in most land mammals, is not laterally com-
pressed but round in cross section or slightly flattened dorsoventrally.

The bones in the skull of whales are arranged symmetrically or asym-
metrically. In the latter case, the bones on the right side are broad-
ened and shifted leftward while the left ones are thickened. Some bones
may overlap others. The cranial section is shortened and rounded. The
supraoccipital bone is highly developed and joined with the frontal bone
anteriorly while the parietal bones are shifted laterally. The nasal bones
are small and the outer bony apertures are shifted far backward and lie
directly anterior to the neurocranium. The nasal passages are short and
run fairly vertically. Maxillary and premaxillary bones and the vomer are
highly extended and form a rostrum. The bony palate is elongated due
to the large pterygoid bones, as a result of which the choanae appear
displaced backward. The mandibles are long and have a rudimentary
coronoid process, which is sometimes lacking.

In the adult state, only the members of the suborder of toothed
whales possess teeth, which are homodont and serve only to hold and kill
the prey. In the embryo of baleen whales, teeth begin to form but soon

420

959

cease to grow and are resorbed. Among adult baleen whales, the teeth
are functionally replaced by special, characteristic horny formations of
the palatal epithelium, the so-called “whalebone” [baleen] which serves
to strain the tiny planktonic organisms on which these whales feed. The
triangular plates of the whalebone are fixed along the edge of the upper
jaw parallel to each other in such a way that the smaller side of the
triangle falls toward the upper jaw while the larger side is turned out-
ward and the hypotenuse is inside the oral cavity. The inner part of the
plate of the whalebone is split into innumerable fine and long keratinous
filaments. The baleen plates are slightly separated from each other and
the filaments of the neighboring plates are intermeshed to form a gauze-
like structure. When the whale swims with its mouth open, water passes
through this structure continuously while the planktonic organisms are
retained on its surface. By closing its mouth, the whale pushes out the
water from the oral cavity with its tongue, which also helps to push the
food particles into the esophagus: whales do not masticate food. The
tongue is well developed and soft lips are absent. Salivary glands are
either altogether lacking or are rudimentary.

The stomach is complex and consists of three to fourteen sections
(Fig. 220). This complexity is explained by the absence of food mastica-
tion. The length of the intestine varies: it may be five or six times the
body length as, for example, among baleen whales (Mysticeti) or among
species of the family of beaked whales (Ziphiidae); 15 to 16 times among
sperm whales and bottlenose dolphins; and up to 32 times among La
Plata dolphins (Pontoporia blainivillei). The liver is relatively small and
a gall bladder is absent. The pancreas is elongated and faintly lobular,
far more rarely disjunct; its ducts open into the bile duct.

External nares (blowhole) are paired among baleen whales and
unpaired among toothed whales, shifted to the top of the head, and
have valves that close them during diving. Among the toothed whales,
the nasal passage above the skull has special air sacs. Elongated laryngeal
cartilages of toothed whales enter the choanae and thus maintain a
constant contact between the larynx and the air sacs located above
without interruption even during swallowing. The trachea and bronchi
are shortened. The short passages transporting air accelerate respiration.
The lungs are single-lobed with highly developed smooth muscles. The
bronchioles have a series of smooth muscle sphincters (among toothed
whales) or contain a large number of muscle fibers in the upper portion
of the alveolar septa (among baleen whales). The connective tissue of
the lungs is highly elastic. Lymphoid tissue is almost totally absent. The
number of alveoli is relatively large and their dimensions somewhat
larger than among land mammals. The epithelium of the bronchi and

560

420 Fig. 220. Internal organs of whales (figure by М.М. Kondakov). 1—lungs; 2—heart;
3—diaphragm; 4—stomach; 5—Пуег; 6—intestine; 7—kidney; 8—urinary bladder.

421

561

glands is almost devoid of mucous cells, which is evidently associated
with the absence of dust in the air intake (ЗШрет, 1958a). Alveolar septa
contain two layers of capillaries and a very large amount of clastic fibers.
All the bronchi have a ciliate epithelium and are surrounded externally by
cartilage of irregular shape. These cartilages are also seen in the terminal
bronchioles (not found among beaked whales, Berardius).

Whales can survive without breathing for quite sometime and remain
under water from 2-10 to 30-40 min (up to two hours, according to
some authors). They are insensitive to large amounts of carbon dioxide
in the blood; its accumulation does not regulate their respiration, as in
land mammals, but an oxygen deficit in the blood does.

The long duration of diving is ensured by the large capacity of the
lungs, insensitivity of the respiratory center of the brain to the accumu-
lation of carbon dioxide in the body, high content of myoglobin, and
slightly higher oxygen-holding capacity of the blood due to its higher
hemoglobin content and its high concentration of erythrocytes (Irving,
1938; Taarra, 1950; Korzhuev and Bulatova, 1952; Kleinenberg, 1956a,
1957).

Hemoglobin content in 100 ml of blood, g .

Sei whale 15.6
Sperm whale 15.8
Cow 12.4
Man 13.0

Respiration in whales can usually be divided into several stages:
(1).exhalation after prolonged diving; (2) intermediate brief inhalation
and exhalation; and (3) deep inhalation before prolonged diving. Dur-
ing intermediate inhalation and exhalation, the whale does not dive very
deep, surfaces almost always vertically, and breathes at regular intervals.
The number of intermediate inhalations/exhalations varies among differ-
ent species; the longer the animal remains under water during diving,
the greater the number. There is a particular reflex movement during
inhalation: the animal strikes the water with its caudal fin in an up and
down movement which invariably keeps the top of the head with the
blowhole afloat (Tomilin, 1957). Exhalation can commence even under
water, as a result of which the air expired under pressure forms a water
jet that is generally characteristic in shape and size for various species.?

3 Apparently, the jet or fountain may also be formed as a result of the condensed
vapor present in the warm air expired (in high latitudes) and also as a result of holding
and spraying of the water which remains, as the whale surfaces, in the cavity in which the
blowhole is located.

422

562

Some whales are capable of diving to great depths (Юг example,
sperm whales can sink deeper than 1,000 meters). The quick surfacing
of the animal from a great depth should liberate the gaseous nitrogen
dissolved in the blood under the high pressure generated during diving.
Otherwise the blood vessels would become clogged with nitrogen bub-
bles, leading to caisson disease and even to the death of the animal. But
whales do not suffer from this disease. This is probably explained by the
fact that the amount of nitrogen present in the lungs slightly exceeds
the nitrogen-holding capacity of the tissues while no more air enters the
lungs during diving (unlike what happens in divers).

The brain in most respects is highly differentiated but in many other
respects has preserved extremely primitive features that are not found
among the members of other orders of mammals (Fig. 221). These differ-
ences are mainly: significantly narrow cortex in cross section while layers
Гапа V-VI, the oldest in evolution, are predominantly developed; overall
high cellular content of the cortical plate; and developmental features
and ratios of the zones of old and intermediate cortex (Filimonov, 1949).
The weight of the brain in absolute terms is maximum in mammals but
is insignificant in relation to the body weight (especially among large
whales). The brain of white whales weighs 2,180 to 2,340 g (Kleinenberg
et al., 1964), fin whales 6.5 to 7.2 kg, and sperm whales 7 to 8 kg (Kojima,
1951; Janes, 1952). Large convolutions and longitudinal fissures are seen
in the short rounded form of the brain. Intense development of the cortex
of the hemispheres of the forebrain is a characteristic feature. Without
exception, the cortex houses the paleocortical, archicortical, and inter-
stitial zones (Filimonov, 1949). The cerebellum attains large dimensions
(up to one-fifth of the weight of the entire brain). The hypophysis is
highly developed.

Fig. 221. Brain of the common dolphin, Delphinus delphis (figure by N.N. Kondakov).

422

563

The structural peculiarities of the cetacean brain are associated with
the high degree of development of the higher nervous activity of these
animals. Many species of whales are easily tamed and trained. With
respect to higher nervous activity, some whales (for example, the bot-
tlenose dolphin) may be placed among the higher mammals although
lower than the anthropoid apes (Voronin, 1970).

Reduction of the rhinencephalon is a characteristic feature. The bul-
bus olfactorius is lacking in most whales (Kiikenthal and Ziehen, 1893)4
and the lobus hippocampi are poorly developed. The V and VIII pairs
of cranial nerves associated with the sensitivity of the facial region and
auditory organs are highly developed. Among bottlenose dolphins (Tur-
slops truncatus), the following features of the brain have been noted
(Kruger, 1959): a prominent flexure between the mesencephalon and
the diencephalon; reduction of tactile thalamic region, evidently asso-
ciated with the absence of hairs in adult animals (this is true of most
toothed whales); intense development of n. ventralis medialis, definitely
suggesting that whales have a sense of taste (although many investiga-
tors refute this); and so forth. Amony some toothed whales (narwhal,
pilot whales, white whales, and common propoises), a small number of
typical pits with minute processes at the bottom have been noticed on
the surface of the root of the tongue. They evidently function as organs
of chemical perception (Yablokov, 1957, 1961a); taste buds are found
in them (Yablokov et al., 1972) as well as many efferent ducts of the
albumen and mucous glands.

The eyes are small (Fig. 222). The cornea and sclera attain consider-
able thickness and the optic muscles are well developed. The crystalline
lens has a characteristic spherical form. Eyelids are not developed. Vision
is evidently monocular and myopia is typical (Kellogg, 1928; and others).
The lachrymal glands are reduced and the nasolachrymal passage is lack-
ing. An oily secretion from Harder’s gland protects the eye from the
mechanical and chemical action of water. Conjunctival glands, not found
among other mammals, are present (Weber, 1886). The olfactory organs
and Jacobson’s organ are reduced.

Sense organs among baleen whales and some toothed whales (for
example, the Amazon dolphins, Га) are represented by vibrissae dis-
posed on the snout and sometimes all over the trunk (Fig. 223). Numer-
ous nerve ends approach the roots of the vibrissae surrounded by blood
lacunae (Japha, 1910). Among all whales, the sensory nerve endings rise

4 The bulbus olfactorius has been described (Flatau and Jacobson, 1899; Filimonov,
1949) in the bottlenose whale, Hyperoodon rostratus, the blue whale, Balaenoptera musculus,
and the fin whale, B. physalus.

422

423

564

Fig. 222. Eye of the fin whale, Balaenoptera physalus (figure Бу М.М. Kondakov).

along the dermal papillae almost directly at the outer surface of the skin
so that the entire surface of the body appears well innervated.

The auditory organs are highly modified (Fig. 224). The concha is
reduced. The external auditory meatus opens outward behind the eye as a
tiny aperture; its lumen is invariably covered. It is possible that the rudi-
mentary auditory meatus may serve as an independent organ perceiving
pressure changes (Yamada, 1953). The tympanic membrane bends out-
ward (among baleen whales) or inward (among toothed whales), and is
externally covered among baleen whales by a typical ear plug of ker-
atinous epithelium and сегатеп.> In view of the fact that the petrous
tympanicum is not only isolated from the other skull bones, but also sur-
rounded by special air cavities, it is possible that whales perceive sound
signals through the auditory meatus. It has been suggested that sound is
perceived through the body surface (especially the head), transmitted to
the auditory zone, and that the petrous tympanicum, which is capable of
vibrations, acts as a resonator. It has also been suggested that the lower
jaw is also capable of perceiving sound from the external environment
and transmitting it to the auditory organs.

Whales are capable of receiving a wide range of sound waves:
from 150 to 120-140 thousand Hertz (Kellog, 1953; Slijper, 1955, 1960;
Bullock et al., 1968), i.e., even ultrasonic vibrations. The high degree of
development of the auditory section of the brain of toothed whales points
to their particularly acute sense of hearing, which is almost unique among
mammals. Among baleen whales, hearing is inferior to that among
toothed whales but better than that among land mammals (Ogawa and

> The age of whales can be determined by counting the layers in a slice of this plug
(Purves, 1955).

565

423 Fig. 223. Vibrissae of whales. Top to bottom: Atlantic right whale, fin whale, and
gray whale (figure by N.N. Kondakov).

Arifuku, 1948). Whales, like bats, are capable of echolocation (Kellogg,
1958). Since vocal cords are lacking, whales cannot produce sound by
the methods ordinarily used among mammals. However, that they do
produce diverse sounds has long been known. Evidently they result from
vibrations of the lower portion of the nasal septum (Tomilin, 1957) or
vibrations of the folds of the outer valve due to passage of air from the
dorsal nasal sacs (Yablokov, 1961b). It should be noted that dolphins
have at least two systems of sound production: they are capable of
simultaneous production of location signals and whistles. It has been
424 suggested that whistles are produced by means of the larynx; impulse

424

566

Fig. 224. Structure of the auditory organs in the fin whale, Balaenoptera

physalus. Top—outer view; bottom—section (figure Бу М.М. Kondakov). 1—outer

opening; 2—membrane; 3—tympanic membrane; 4—ligament between tympanic
membrane and malleus; 5—ear plug; 6—auditory ossicles.

signals are apparently produced by the air sacs (Evans and Prescott,
1962; Norris, 1964). The typical structure of the frontal section of the
skull and the disposition of the fatty “lens” anterior to it enable the
animal to judge the direction of sound waves (Romanenko et al., 1965).

The blood circulatory system of whales has several characteristic
features, some of which are also encountered in other aquatic and ter-
restrial mammals (Slijper, 1959): (1) arterial rete mirabile is present on
the inner side of the vertebral column and particularly well developed
in the regions of the neck, thorax, between the ribs, at the base of the
brain, and around the spinal cord. It is formed of a large number of sinu-
ous arteries of the muscular type interconnected by several anastomoses.
The histological structure of the vessels points to their capacity to modify
considerably the volume of blood flowing in them; (2) the venous rate
mirabile is present at the base of the skull and is particularly massive in
the abdominal zone; (3) an expansion occurs in the hepatic vein (lack-
ing in baleen whales); (4) two vertebral veins running along the ventral
side of the vertebral column are almost identical in diameter throughout
their length and lack valves, indicating that blood can flow in them in
both directions: through the major уу. costocervicales they run into the

425

567

anterior vena cava and through the veins between the ribs into the pos-
terior vena cava. Since all these features of the blood circulatory system
attain maximum development among smaller whales, they are evidently
associated not with deep and prolonged diving of the animals, but with
frequency of diving and respiration. A reference has already been made
to the veinous vessels of fins typical of whales (arteries surrounded by
small veins). In addition to the very high (compared to terrestrial mam-
mals) oxygen capacity of the blood, the considerable concentration of
sugar and sodium chloride in it is a characteristic feature of whales.
This is evidently associated with the deposition of additional reserves
of food required by the animal during prolonged underwater residence.
The diameter of erythrocytes in whales varies from 6.6 to 10.0 microns
(Lenfant, 1969).

The body temperature of whales is similar to that of land mam-
mals and varies from 35 to 40°C (the upper limit was recorded among
wounded whales or dolphins caught after chasing). The maintenance of a
high body temperature in water, which conducts heat many times better
than air, is facilitated by the thick layer of subcutaneous adipose tissue.
Killed whales 60 h after the moment of death and in cold water recorded
a temperature close to 30°C®, 1.е., only 6-7°C loss of heat during this
time (Zenkovich, 1938). The temperature of the outer layers of the skin
among whales is close to the temperature of the surrounding water. This
is achieved by minimal heat dissipation by the body as required for nor-
mal activity in such a highly heat-conducting medium as water. In the case
of over-heating of the whale body (for example, on rapid and prolonged
swimming), dissipation of heat occurs from the entire body surface but
more intensely from the surface of the fins.

Compared to land mammals, the musculature of the trunk and limbs
has undergone the most intense modification. As a result of losing
the locomotory function, the following muscles in the forelimbs are
very weak: m. romboideus, m. serratus, m. levator scapulae, and m.
infraspinatus; m. trapezius and m. teres minor are totally reduced; and
m. deltoideus, m. subscapularis, and m. paniculus carnosus are highly
developed. In the trunk, m. latissimus dorsi and m. ileo-costalis passing
through the dorsal side of the vertebral column as also m. hypaxialis
lateralis and m. hypaxialis medialis passing along its ventral side are
particularly well developed. The former are attached anteriorly to the
skull and the latter to the lateral processes of the posterior thoracic

6 The high body temperature of the killed whales may also have been partly main-
tained by the process of putrefaction; nevertheless, this fact points to the considerable
heat-insulating capacity of the subcutaneous adipose tissue.

427

568

vertebrae and to the last of the ribs; posteriorly, these muscles are
attached to the caudal vertebrae and (Narkhov, 1937) partly transit into
massive tendons running into the flukes and diverging fanlike there.

The kidney of whales is multilobulate and relatively much larger
than in land mammals (Slijper, 1958a). The urinary bladder is small;
the absence of a sphincter in its neck facilitates frequent urination in
small quantities which possibly may serve as a signal for other whales
(Yablokov, 1961a). Whales evidently do not swallow sea water and the
required moisture is provided by the food intake in addition to the water
of metabolism. The absorption capacity of the kidneys of whales is not
so high as to extract fresh water from the swallowed sea water. The
concentration of sea water was found to be more than that of the urine
(if whales swallowed sea water, this ratio would be reversed) (Table 30).

The testes are located in the abdominal cavity. In blue whales, the
length of the testes may reach 45 cm and the weight 45 kg (Slijper, 1966).
Of the other glands, only the prostate is present. The penis is long and
narrows gradually toward the tip; at rest, it lies in a special sheath. At its
base, the penis bends in an S-shape. An os penis is lacking except in the
right whale, Balaena glacialis. The cavernous tissue is poorly developed
in the penis. The crus of the crura penis are attached to the rudimen-
tary pelvic girdle; here, mm. bulbo- and ischiocavernosi are separated.
Among adult females, the ovaries are partly or wholly surrounded by
an extensive infundibulum of the oviduct. The oviducts extend into the
bicornuate uterus, which has longitudinal folds on the inner surface. The
portio vaginalis enters the vagina. The outer slitlike genital opening is
surrounded by small liplike bulges between which the boneless clitoris is
located. Ureters open into the vulvar region; there is no sinus urogeni-
talis. The vagina contains several ringlike folds which probably perform
the role of valves, protecting the genital system from water entering into
it during copulation and whelping. In female baleen whales, the genital
Opening and the anus are separated from each other by a considerable
distance while in toothed whales, these are located in a common Sac and
are surrounded by a common sphincter (Fig. 225). The males are capable
of fertilizing throughout or for much of the year.

Ovulation among whales is perhaps stimulated by copulation
(Sokolov, 1950, 1954; Sleptsov, 1952; Tomilin, 1957; Yablokov, 1959) but
the recovery of spermatozoa in the vagina of female common dolphins
having mature but not opened-up follicles in the ovaries casts doubts on
this assumption (Sokolov, 1961). On fertilization, the corpus luteum of
pregnancy develops in the ovary and, after resorption, traces of it are
preserved for a long time (presumably throughout life) in the form of
a corpus albicans. The age of the female can possibly be determined by

569

—

КИГИ /
“LD, 77772 22222772 2 ly
РРР Е: EE LLL РРР
ИРИ 77 РРСРР. й
РР РРР й
ат - 4% Z Os
SEES рр. Я. - 27
jp
eee a
——<— Segoe eon ee oe te IP ie
CLE
1 2 A
“
7)
РРР Я
ПАР Ре РОРЕРРРРРРР ,
ИР 2777; GLEE Ze 22222 i
ne. We ИИ SOLIS РРР Ce; 72 АА: РР = й 4 di
a VE
a LEED J
РЕ i
<< cere lw =
=== He НИЕ Е Liza >
Zig — =
<->» oe = Е
пе 22 В
427 Fig. 225. Zone of disposition of the urogenital organs in the fin whale, Balaenoptera

physalus. A—male; B—female (figure by N.N. Kondakov). 1—genital
opening; 2—anal opening; 3—sac for mammary teats.

counting these bodies. Among pregnant females, two or three fetuses can
be found in the uterus at the commencement of pregnancy but shortly
thereafter only one is found. The placenta is diffuse, of the epithelio-
choreal type (Zhemkova, 1965), and may partly extend into the second
horn of the uterus. In the late stages of pregnancy, the embryo is usually
situated with the tail toward the womb exit. The lobes of the flukes are
convoluted, the dorsal fin (when present) bent down towards the back,
and the flippers turned towards the tail. The sex ratio of embryos is
roughly 1:1 (Table 31).

Birth takes place under water. The calf is born fully developed and
capable of independent movement; its body proportions are highly sim-
ilar to those of adults (Zemskii, 1958a) but its length up to 1/4 to 1/2
that of the mother. Some female whales can be fertilized soon after par-
turition, during the lactation period. The calves are suckled under water,
the duration of each suckling being short (a few seconds). The calf holds
the teat of the mother (Fig. 226) between its tongue and upper palate.

428 An exception is the sperm whale calf, which holds the teat in the corner
of its mouth (Slijper, 1966). Milk is sprayed into the oral cavity of the
calf by the contraction of special muscles in the female. Newborn calves
feed very frequently; for example, calves of bottlenose dolphins suckle

570

one
TTS 6 LY Bes (On) СИТ STS pss a4 Crs ССР OLY STS a3e]U9019g

$цэци9э@$
179 ILS 887 Lvz LI 61 79 05 6 18 LSI PLT Jo лэашам

o[RUIS.J SIN этешэ ae a[ewi3y ae эгешэН эти эешэ Te o]ewsy aye
[е1оТ, ЭТЕЧАА 11э4$ эгеЧчм yoeqduinyy ЭГеЧм эп[а ЭТЕЧА 19S ЭГеЧм и злетоплтеа

(ISs6T ‘озелай рце эп21 {0661 ‘одип[ рие anziy) зоАлашо э[еЧм Зиоше OBI хэб “TE э[ает,
= LE Ors bs от 8'6 06? = ЕСС тотем BIS
bE 6'5 6$ 1Z 81 = == OLS ETE if STEM SITY A
= = ObZ-OLT от OL = = 078-08? — 15-855 с urydjoq
= = O8T = = = == 08$ 10°Z т этечм зопа
19 = Ove = = = == OZ 095 I yey onig
1-64 0-1 065-055 9'9 ЕЕ [9 0/5 057-085 = t эГечл uly
ТУТ = 0/5-09%% 85-6966 6531 CL-9S 055-056 05%-055 == с ayeym uly
Iy-9T 6-95 OLE-O81T TPr-9T Z-€0 58-9 06Z-OFZ 055-088 9r'7 Ф ayeym 125
LCZEG= ВСЕ Т 10/9095 = oe == = 059-05 = ф этеЧл 125
1Z 8$ OLE = 6 ps 05$ 00$ 67< I этецм 19g
’Od 705 5) ЗИ i) Я ем 6211] Do suoneuIwajap oTeUM
Jo} [/WIu ‘и0ел11э940) jo Joquinyy

(6€61 ‘1эЦэуэЯ) sofeyA Jo autin ay} Jo ион 15040) ‘0$ тает,

9th

9¢L

427

429

ЙЕ

Fig. 226. Teat of a lactating female fin whale, Balaenoptera physalus (figure by
N.N. Kondakov).

roughly once every 26 min day and night (McBride and Kritzler, 1951).
The mammary glands of the female lie along the sides of the genital
opening. Two teats (one on each side) lie in slitlike folds and project
outward only during lactation. Female whales produce a large amount
of milk daily: from 200 to 1200 g among dolphins, up to 90 to 150 liters
among fin whales, and 200 liters among blue whales (Sleptsov, 1955).
The milk is thick and usually cream-colored. Its surface tension is 30
times more than that of water, which is particularly important consider-
ing that suckling takes place under water (the milk spray does not spread
on water). The nutritive value of whale milk is very high (Tables 32, 33,
and 34).

A different composition of the milk of fin whales has been given by
Ota et al. (1955) (Table 33).

During the period of suckling the growth of calves proceeds very
rapidly: for example, by seven months of age the calf of a blue whale
had grown to 16 m from 7 м at birth, 1.е., the average daily increment
of length was 4.5 cm (Slijper, 1960).

Sexual dimorphism among whales is manifest mainly in the differ-
ences of body length between males and females. Among baleen whales,
the females are larger than males while the converse is true in most
toothed whales. Whales are mainly herding animals and live in schools
of a few animals to hundreds and thousands. They are encountered

428

428

428

572

Table 32. Composition of the milk of large whales (2, averages) (Zenkovich, 1952)
Species Fat Dry residue Water
Fin whale 43.57 13.50 42.93
Blue whale 40.25 13.70 46.05
Humpback whale 39.93 14.02 46.05
Gray whale 53.04 6.38 40.58
Sperm whale 37.30 8.00 54.70
Cow 3 РА TUS)

Table 33
Fat Protein Water Lactose Inorganic
matter
33.0 13.3 53.4 0.3 1.0
31.8 12.3 55.0 0.2 0.7
32.5 10.5 54.1 1.4 1.4
Table 34. Composition of the milk of small whales (2) (Ural’skaya, 1957)
Species Fat Protein Sugar Dry Ash Water
residue

Common dolphin 43.71 5.62 1.45 7.53 0.45 48.76
Common dolphin 41.56 4.88 1.49 6.82 0.45 51.62
Common porpoise 45.80 11.19 1633 13.09 0.57 41.11
Common porpoise 33.90 5.22 1.28 7.10 0.60 59.00

Bottlenose dolphin 46.10 11:55 1.5 13.50; 0.38 40.50

along coasts as well as in the open sea. Some are capable of ascend-
ing along major rivers opening into the sea while some species are
regularly confined to rivers. The food of many is specialized; plank-
tophagous, teuthophagous, ichthyophagous, and sarcophagous animals
are known (Tomilin, 1957). Whales include fast swimmers (for example,
killer whales and many dolphins) and relatively slow-moving animals
(gray whales). Most whales regularly remain on the surface while some
(for example, sperm whale) can dive to great depths.

The population of the various species of whales is not identical.

Some whales are most numerous and can be found in thousands of
schools (common dolphin) while others are very rare and only a few
sightings have been reported (some members of the genus Mesoplodon

573

(beaked whales) dwarf sperm whale, and false killer whale). Natural fluc-
tuations in population have not been studied but can hardly be signifi-
cant. As a result of senseless hunting, the population of many species of
whales has sharply decreased. Thus, right whales and gray whales have
come close to total extinction. International conventions have imposed
a total ban on the hunting of these whales and their population is now
slowly recovering. The hunting of even sperm whales and fin whales,
numerous in the recent past, has to be totally banned to maintain their
population at the present level. A sharp reduction in whale hunting or
its total ban for several years, followed by rational and strictly controlled
hunting, should be considered.

Periodic migrations are characteristic of many whales. Among some
species, the distance covered during migration is relatively small (Azov-
Black Sea common porpoise migrates from the Azov Sea into the Black
Sea and back) while migrations among others cover long distances: some
large whales travel from tropical waters to high latitudes.

Most whales are monogamous. Periods of mating and whelping are
usually prolonged. They give birth to one (rarely two) calves. The mate-
rial instinct is strongly developed among females.

Apart from man and the killer whale, there are no enemies of
cetaceans.

Cetaceans are universally infected with endo- and ectoparasites
(crustaceans). The latter are particularly typical of large whales.

Whales are found in all the oceans, in most of the world seas, and in
some rivers and lakes. Two factors determine the distribution of whales:
food and water temperature. Some species are widely distributed and are
encountered in warm as well as cold seas (some species of the dolphin
family), others have a small range (gray whales inhabit the subtropical
moderate and cold waters of the North Pacific Ocean and the Chukchi
Sea), still others are even more restricted (narwhals do not leave the
Arctic waters), and finally, the ranges of river, lake, and estuary forms
are altogether insignificant.

At present, relatively few fossil remains of whales are known.
Remains of the more primitive of the known whales have been found in
the Middle Eocene of North Africa. There is no unanimity of opinion
regarding the ancestors of whales. Some scientists support the origin of
whales from ungulates, with which they share many common features,
such as diffuse placenta, bicornuate uterus, and complex stomach. The
number of chromosomes and the reaction of the precipitation of serum
proteins of whales confirm this hypothesis (Makino, 1948; Boyden and
Gemery, 1950). Based on the structural similarity of the skeleton and the
dental system of extinct whales with the primitive carnivores (creodonts),

430

574

some scientists assume that the whales originated from the latter. А view
has been expressed that the ancestors of whales were much older than
creodonts, i.e., cretaceous insectivorous forms (Slijper, 1958a). Some
authors (Beddard, 1900; Kukenthal, 1900; Hosokawa, 1950; Kleinenberg,
1958; Yablokov, 1964; Anderson and Jones, 1967) suggest a diphyletic
origin of the contemporary order of cetaceans, i.e., that the baleen
and toothed whales evolved from different ancestors and that their
evolution proceeded by convergence and not divergence; hence it would
be more correct to regard these suborders as orders. In confirmation of
this hypothesis, numerous morphological differences between the baleen
and toothed whales are cited but such evidence cannot be regarded as
sufficiently convincing.

The ancient whales, Archaeoceti, are sometimes regarded as conver-
gent with other whales although they have nothing common in phylogeny
and hence should not be included in the order of cetaceans (Yablokov,
1964). But even this assumption is not supported by adequately convinc-
ing data (Mchedlidze, 1970).

The systematics of Cetacea has not been properly worked out. As
mentioned earlier, even the content of this order has been disputed by
some. The composition of some families and subfamilies has also not
been conclusively established. Usually (Simpson, 1945), three suborders
are distinguished in the order Cetacea: modern toothed whales (Odon-
toceti Flower), baleen whales (Mysticeti Flower), and extinct whales
(Archaeoceti Flower). Sixteen families (nine are extinct) and 173 genera
(of which 137 are fossil forms) are usually recognized in the order. The
total number of contemporary species of the order is 81 (Hershkovitz,
1966). The USSR is host to 6 contemporary families, 19 genera (roughly
52% of the world fauna), and 23 species (roughly 28% of the world fauna)
(Tomilin, 1957). The suborder Archaeoceti comprises three families:
Protccetidae Stromer (Middle Miocene), Doridontidae Miller (Eocene
and Miocene), and Basilosauridae Cope (Eocene and Oligocene), and 14
genera (Simpson, 1945). The family Protocetidae includes short-bodied
forms such as Protocetus with a length of not more than 2.5 m and also
long-bodied animals (in the available long bodied form of Eocetus, the
skull alone measures 120 cm in length). Evidently, Eocetus was the source
of the family Basilosauridae while Protocetus and Pappocetus gave rise
to the members of Dorudontidae (Deshaseux, 1961).

The family Dorudontidae comprises animals 8 to 9 m in length with
an elongated and flexible cervical section and serrate cheek teeth while
the family Basilosauridae includes animals of 8 to 22 m in length with
some skeletal characteristics attaining a high degree of development
within the suborder (Romer, 1966).

431

432

55

The economic importance of whales is quite significant although уаг-
105 representatives are not of equal importance in this respect. Some
whales are caught in large numbers every year (fin whales and sperm
whales) while others are hunted only at random. Valuable edible and
commercial products are obtained from almost all organs of the whale.
Oil is rendered from the skin and bones (commercial and edible). The
meat contains 20 to 26% protein and can be a source of diverse pro-
tein concentrates (frothing and emulsifying agents, substitutes for egg
protein and deficient peptone for preparing nutrient media in microbi-
ology) (Faingersh et al., 1953). The pancreas of whales can be used to
produce insulin (1 kg of pancreas contains from 1,000 IU in the blue
whale to 3,000 in the sperm whale; Egorova, 1953) and commercial pan-
creatin (skin-softening agent; Bodrov et al., 1958); the liver can be used
to produce vitamin A (1 g of liver contains from 400 IU in the hump-
back whale to 5,800 in the sperm whale; Mrochkov, 1953), vitamin В,
and campolon (Bodrov et al., 1958). The brain of whales can serve as
raw material for producing cholesterol (frozen brains of fin whale, blue
whale, and humpback whale contain roughly 2.5% or about 11% of it in
terms of dry matter; Egorova and Lebedeva, 1953).

The skin of whales, apart from providing fat, is useful as leather. The
upper denser sections of the sperm whale are used to produce sole leather
while the lower more porous ones are used as soft leathers (Zaikin,
1953). (V.S.)

Key to Suborders of Cetacea

1 (2). Teeth absent. Numerous horny plates on upper jaw form straining
apparatus. External nares bipartite. Skull symmetrical. Middle sec-
tions of movable halves of lower jaw flexed outward. Only first pair
of ribs joined to sternum. ... Suborder of baleen whales, Mysticeti.

2 (1). Teeth present. Straining apparatus absent. External nares not
bipartite. Skull asymmetrical in frontal section. Middle sections
of immovable halves of lower jaw straight or flexed inward. Not
less than three pairs of ribs joined to sternum..................
уе ие 8 Suborder of toothed whales, Odontoceti (See below).

SUBORDER OF TOOTHED WHALES

Suborder ODONTOCETI Flower, 1867

Size small, medium, and large (body length up to 2 m, up to 4 to 6 m,
and up to 21 m). The body is torpedo- or teardrop-shaped.

432

433

576

Fig. 227. Skull of the common dolphin, Delphinus delphis (figure by М. М. Kondakov).

In the skin, the dermal layer and, in most cases, its fat-free portion
are well distinguished. The network of elastin fibers is relatively poorly
developed (except in small dolphins). A hair coat is lacking in most
species. The body color is monochromatic, sometimes with typical white
spots and bands.

The skull bones (Fig. 227) are sharply asymmetric although, in the
early stages of embryogeny, the skull is characterized by all features char-
acteristic of the skull of land mammals (Sleptsov, 1940). But this similar-
ity disappears very soon as a result of the intense growth of the supraoc-
cipital and premaxillary bones and also the supraorbital processes of the
frontal bones, and the skull of the embryo then resembles that of an
adult. Uneven development of the skull bones is also seen in the late
stages of embryonic growth and extends into the postembryonic stage
(Sleptsov, 1940c). The reasons for the genesis of asymmetry have not
been accurately ascertained thus far: some researchers suggest that the
unequal pressure of water on different sections of the skull while swim-
ming is probably responsible while others regard this asymmetry as a
consequence of the reduction of olfactory nerves. The nasal bones are
poorly developed and do not cover the posterior part of the bony nares.
The nares are displaced leftward and open into a common chamber. Max-
illae, premaxillae, and nasal bones are notably shifted onto the frontals
and almost wholly cover them. Primitive features of structure are seen in
the skull: presence of os parasphenoideum, tabularia, and postfrontalia,;
the number of bony components in the skull of embryos is more than in
juvenile and adult animals (Sleptsov, 1949c). Cervical vertebrae may be

433

577

fused. Among the fast-swimming species, the number of caudal vertebrae
is more. Among the deep-diving animals, because of the reduced num-
ber of ribs (in the sperm whale, their number decreases to three), the
latter are joined to the sternum by flexible joints or attached by cartilage
(beaked whales).

The dental formula varies from о or : to roughly 2. The teeth are

homodont and monophyodont. Three types of teeth are distinguished
(Fig. 228) (Yablokov, 1958a):

i

Simple peg-shaped teeth with highly developed pulp cavity and thin
cement and enamel layers among adult animals. This type of dental sys-
tem is found in the common dolphin, common porpoise, pilot whale,
etc. Except for the pilot whale, all these animals have a large number
of teeth uniformly distributed in the jaws.

. Teeth with highly developed layer of cement; enamel lacking on the

teeth of adult animals; the tooth crown in the young animals has a
thin layer of enamel (Lenberg, 1911). Teeth are simple, peg-shaped,
larger than in the first type, and their number goes up to 30-50. The
pulp cavity is well developed or absent. This tooth type is seen among
sperm whales, white whale, apparently in dwarf sperm whale, Risso’s
dolphin, dwarf killer whale, Irrawaddy River dolphin, and false killer
whale.

. Teeth are flat, wedge-shaped, with highly developed enamel layer and

cement filling the pulp cavity and thus covering the entire tooth with
the exception of the crown; further, the cement layer typically adheres
to the enamel layer in the midportion of the tooth. The number
of teeth is small and they occur only in the lower jaw. This type is
characteristic of Baird’s beaked whale, beaked whales, and apparently
the rest of the members of this family except for the Tasman beaked

Fig. 228. Teeth of whales. A—common dolphin, Delphinus delphis; B—white
whale, Delphinapterus leucas; C—Baird’s beaked whale, Berardius bairdi (figure
by N.N. Kondakov).

434

578

whale. The teeth of the killer whale occupy an intermediate position
between the first and second types. They are similar to the teeth of the
first type but differ from them in their large dimensions, comparatively
small pulp cavity, and large amount of dentine. In these respects, they
resemble the teeth of the second type but, in contrast to them, have a
poorly developed cement layer.

In most of the toothed whales, the number of teeth is subject to indi-
vidual variation. Among some species, teeth are fewer in the upper than
in the lower jaw (Baird’s beaked whale, sperm whale, Risso’s dolphin,
etc.); among others, on the contrary, the number of teeth in the upper
and lower jaws is identical (narwhal, white whale, common porpoise, and
black finless porpoise).

Annual layers are deposited on the teeth and the age of toothed
whales can be determined by counting these layers in a section of a
tooth. A unique feature of some cetaceans is that the left (rarely the
right) upper tooth has transformed into a tusk, as seen in the narwhal
males (less commonly, in females also), reaching a length of 2.7 m.

The digestive system among all the species is totally separated from the
respiratory tract and has several characteristic features (Yablokov, 1958b).
The tongue, unlike in land mammals, has a different structure and function.
It is highly mobile and is covered with a thick sheath consisting of many
layers of horny epithelium and a massive layer of connective tissue. The
tongue positions the captured quarry in the oral cavity, pushes it into the
gullet, and prevents water from entering. The soft palate is absent.

The initial sections of the digestive tract are covered inside with
many layers of horny epithelium, which evidently protects the tract from
damage by hard portions of food swallowed whole. The multichambered
muscular stomach (Fig. 229) among species feeding predominantly on
cephalopods (sperm whale and Baird’s beaked whale) has a lining of
glandular epithelium in the first section (Sleptsov, 1955). The subsequent
sections of the stomach apparently take part in the process of absorbing
the food; their structure is similar to that of the intestine. A cecum is
lacking (seen in Platanista; Weber, 1927) and the intestinal sections are
not distinctly separated from each other.

The relative length of the intestine varies markedly from species to
species.

Ratio of length of intestine to body length
(Yablokov, 1958b)

Bottlenose whale 1:6.0
Sperm whale 1:15.0-17.5

435

433

579

Common dolphin 110451225
Common porpoise 1:12.2 13.8
Bottlenose dolphin 1:14.4- 15.8
Chinese finless porpoise 1:6.5
White whale 1:61 5951

The respiratory system too reveals several characteristic features.
The external nares are not bipartite (Fig. 230) and are supported by
dense dermomuscular folds, such that the projections of one fall into
the depression of the other. Several special paired air sacs lie above the
skull: the upper ones dorsally, the lower ones proximally, the rear ones
nasofrontally with additional cavities, and the lateral ones enterolaterally
(Kleinenberg and Yablokov, 1958). Their function consists in supporting
the nasal passage while diving; the greater the pressure of water, the more
intense their action. Moreover, they produce sounds.

The larynx projects into the internal nares in the form of a tube
formed by elongated arytenoid cartilages and the epiglottis (Fig. 231).
This tube is covered by a special muscular sphincter. Bronchioles contain
annular sphincters of smooth muscles (lacking in bottlenose whales).
These are best developed in smaller species which dive often and thus
breathe often and have a relatively large lung capacity.

Apparently, the system of sphincters enables adaptation to pres-
sure changes in the lungs during frequent dives and powerful inhala-
tion/exhalation (Slijper, 1958a).

2 Lk 4 >
А №
Ху In? \
ie EMSS NAS
\ Sy h АИ
Ne ies
SY ASA 1
S NSE yy
SS <<
RSI 3
WS
WEY
REY

Fig. 229. Structure of the stomach of toothed whales (figure by N.N. Kondakov).

1—esophagus inlet; 2—first section of stomach; 3—second section of

stomach; 4—third section of stomach; S—sphincter; 6—duodenal ampule;
7—commencement of duodenum.

580

The brain has well-developed ventral nuclei in the auditory nerve,
anterior Olives, etc. (Ogawa and Arifuku, 1948). The auditory organ dif-
fers significantly from that of baleen whales in the shape of the tympanic
bulla and the mode of its attachment to the skull (Yamada, 1953).

Sexual dimorphism is manifest among many species and is most dis-
tinctly seen in the dimensions of the animals: the males of most toothed
whales are larger than the females. There are other differences too in
some species. Thus, the dorsal fin among male killer whales is much
higher than in females while a tusk is a feature of male narwhals (mainly).

In the course of postnatal development, changes occur in body
dimensions and its sections. Some species undergo color changes too
(for example, white whales which are dark colored at birth turn lighter
and become white with advancing age).

>

<
<

<
3 ~

СХ

А В

Fig. 230. Unpaired blowhole among toothed whales, viewed from above (figure
by N.N. Kondakov). A—bottlenose whale; B—sperm whale.

Fig. 231. Structure of the upper respiratory passages (figure by N.N. Kondakov).

436

581

Almost all toothed whales are fast swimmers, which has а bearing
on their food characteristics. Some are capable of deep and prolonged
submergence. They live on the coasts as well as in the open sea.
Some are found in rivers and species of the family of river dolphins,
Susuidae, regularly inhabit large rivers (Ganges and the Amazon). All
the toothed whales, unlike baleen whales, swallow prey entire. Depending
on the predominant food, several adaptive groups can be distinguished
(Tomilin, 1957).

1. Ichthyophagous whales, feeding mainly on pelagic schools of fish.
Their rostrum is long and narrow and the teeth numerous and covered
with enamel. The common dolphin and dolphins of the genus Stenella
belong to this group.

2. Bentho-ichthyophagous whales consuming mainly bottom-
dwelling fish and invertebrates. The rostrum is considerably shortened
and the number of teeth less. Common porpoise, white dolphin,
bottlenose dolphin, etc. belong to this group.

3. Teuthophagous whales, feeding mainly on cephalopods. These
have a small number of teeth but none at all on the upper jaw. Sperm
whale, beaked whales, bottlenose whale, etc. belong to this group.

4. Teutho-ichthyophagous whales (pilot whale and false killer whale),
consuming cephalopods and fish. They occupy an intermediate position
between the second and third groups. The rostrum is reduced in length,
but broadened because of the premaxillae. Although the number of teeth
is less, they are well developed.

5. Sarcophagous whales (killer whales), along with fish also feed on
large marine mammals. They have short but massive jaws with few but
very strong teeth.

Toothed whales are distributed in all the oceans, in almost all the
open seas of the world, and in some rivers and lakes.

Among the primitive Odontoceti of family Agrophiidae Abel, 1913
(Upper Eocene), the early stage of bone extension in the skull has
been observed: the maxillae began extending posteriorly and covering
the frontal bones while the nostrils occupied a position above the orbit.
The members of Agrophiidae apparently represent the ancestors of
Squalodontidae Brandt, 1873 (Oligocene-Miocene) which in the Lower
Miocene gave rise to contemporary dolphins (Delphinidae) and sperm
whales (Physeteridae) and during the Miocene to beaked dolphins
(Ziphiidae).

The systematics of the suborder of toothed whales has not been
adequately studied, not only with respect to morphology and diagnosis
of the species, but even the number and composition of the families.

582

Most scientists have по doubts about the correctness of recognizing
river dolphins, beaked whales, and sperm whales as individual families
(Kellogg, 1928; Nishiwaki, 1972 placed the dwarf sperm whale in the fam-
ily Kogiidae). Views differ regarding the white whale and the narwhal,
dolphins per se, and the common porpoise (for more details, see p. 586).
Combining all the three above-mentioned groups of whales into two
families is evidently more correct (Hershkovitz, 1966; Yablokov et al,
1972). Thus the suborder of toothed whales includes three contemporary
superfamilies and four families: river dolphins—Platanistoidea Simp-
son with one family of river dolphins, Platanistidae Gray (= Susuidae);
dolphins—Delphinoidea Flower with two contemporary families: dol-
phins, Delphinidae Gray and narwhals, Monodontidae Gray; and sperm
whales—Physeteroidea Gill with two families: sperm whales, Physeteri-
dae Gray and beaked dolphins, Ziphiidae Gray (= Hyperoodontidae).
The suborder comprises 31 genera and 71 species (Hershkovitz, 1966).
In the USSR, the members of two superfamilies and three families (not
river dolphins), 15 genera (roughly 48% of the world fauna) and 17
species (roughly 23%) are found (Tomilin, 1962).

The suborder includes five extinct families: Agrophiidae Abel
(Upper Eocene of North America); Squalodontidae Brandt (Upper
Oligocene-Upper Miocene of Europe, Lower Miocene of South America,
Australia, and New Zealand, and Middle and Upper Miocene of North
America); Eurhinodelphidae Abel (Lower Miocene of South America,
Lower and Upper Miocene of Europe, Middle Miocene of North
America, and Upper Miocene of Japan); Hemisyntrachelidae Slijper
(Middle Miocene of North America and Lower Pliocene of Europe);
and Acrodelphidae Abel (Lower and Upper Miocene of Europe and
Middle Miocene and Lower Pliocene of North America).

Some toothed whales have acquired economic importance in the
recent past. Sperm whales are even now caught in large numbers. At the
same time, many of the scarce and less numerous species are practically
of no importance.

Not all the species of toothed whales which could serve as valuable
targets of hunting are caught in sufficient numbers in the waters of the
USSR (for example, killer whales) and some are not caught at all (for
example, dolphins in the waters of the Far East). (V.S.)

Key to Species of the Suborder of Toothed Whales Inhabiting and
Probably Found in USSR Waters

1( 6). Anterior part of snout strongly projecting beyond tip of lower
jaw. Functional teeth absent or not more than three pairs in

2768},

3) (12).

4 (5).

5 ( 4).

6( 1).

7 (16).

Sec):

9 (10).

10 ( 9).

583

upper jaw and not less than seven pairs in lower jaw. Symphysis
of lower jaw halves covers more than one-third their length.
Body dimensions large (3-5 m in newborns and up to
15-21 m in adults). Head accounts for one-third body length.
Blowhole at end of snout. Dorsal fin low, hump-shaped. Lower
jaw with 18-28 pairs of massive teeth. First cervical vertebra
freenMemaAINGeMUSEG HET. ом аа able t os eae:
MIDE TOR fade ne 2 Sperm whale, Physeter catodon (p. 801).
Body dimensions small (not more than 4 m in adults). Head
accounts for roughly one-fifth body length. Blowhole at center
of head. Dorsal fin concave along posterior margin. Lower jaw
with 9-16 pairs of small curved teeth. All cervical vertebrae
fused.

Body length 2.7-3.4 m. Lower jaw with 12-16 (less often, 10
Onli) spairssOf teeth eR VERT ORs PN TIA rte. 2 on S30
PEMA, ДЕН Pygmy sperm whale, Kogia breviceps (p. 841).
Body length 2.1-2.7 m. Lower jaw with 8-11 (less often, 13)
pairs of teeth. ... Dwarf sperm whale, Kogia simus (p. 845).
Anterior part of snout slightly projecting or not projecting
beyond tip of lower jaw. Functional teeth present in both upper
and lower jaws. If absent in upper jaw, their number in lower
jaw does not exceed seven pairs. Symphysis of lower jaw halves
does not exceed one-third their length.

Teeth absent in upper jaw, not more than two pairs in lower
jaw. One or two pairs of furrows on neck converging anteriorly.
Notch not present on posterior edge of caudal flukes. Dorsal
fin closer to flukes than to flippers.

Width of midportion of straight, long, thin rostrum not less
than one-seventh its length. Only one pair of teeth in lower jaw,
located far behind anterior end of jaw. Teeth highly flattened
laterally (their smaller diameter not less than one-half larger
diameter). Fully developed tooth laterally trapezoid.

In skull, antorbital notches relatively poorly developed.
Aperture of pair V of nerves in maxilla anterior to aperture in
premaxilla or at the same level. Palatine bones not contiguous.
Ratio of length of tooth crown to width more than 6:1 ......
.... Stejneger’s beaked whale, Mesoplodon stejnegeri (р. 866).
In skull, antorbital notches relatively well developed. Aper-
ture of pair V of nerves in maxilla posterior to aperture in
premaxilla. Palatine bones contiguous. Ratio of length of tooth
crownto width notwmore Вай 4 ei. lees teresa iaielerst

438.

584

Te:

12 (13).

13 (12).

14 (15).

15 (14).

16, (7):

17 (26).

18 (23).

19 (22).

20 (21).

21 (20).

22 (19).

23 (18).
24 (25).

Width of midportion of rostrum not more than one-sixth its
length. Teeth located on anterior end of lower jaw and conical
(larger diameter exceeds smaller by not more than 1.5 times).
Length of head about 20% of body length. Massive crests
running along maxillae reaching level of skull apex. Maximum
diameter of anterior tooth usually not more ап 2 ст ......
Northern bottlenose whale, Hyperoodon ampullatus (p. 885).
Length of head about 13-17% of body length. Low crests run-
ning along maxillae not reaching level of skull apex. Maximum
diameter of anterior tooth does not exceed 2 cm.

Two pairs of teeth present. “Beak” flattened dorsoventrally.
Rostrum from above not triangular ......................4.
te eer Baird’s beaked whale, Berardius bairdii (p. 850).
One pair of teeth present. “Beak” not flattened dorsoventrally.
Rostrum) from, abovetniangular.., Aisa) ана в oe eee
eRe TING ET. Cuvier’s beaked whale, Ziphius cavirostris (p. 876).
In most species, teeth present in upper and lower jaws. Teeth
absent in upper jaw only in Risso’s dolphin, and in lower jaw
in narwhals. In latter case, however, width of midportion of
rostrum 2/5 - 1/3 its length. Furrows not visible on neck. Notch
present between caudal flukes.

Dorsal fin lacking.

Head rounded, without “‘beak”’.

Number of teeth not more than 44. Neck distinctly visible.
Horny protuberances absent on dorsum.

Body monochromatic (white or yellow in adults). Up to 44 teeth
in upper and lower jawsiit 32 при Benue Ot tec A ees
SLUNG, EELS. ee White whale, Delphinapterus leucas (p. 757).
Body of adult animals spotted (monochromatic, dark, in juve-
niles). Teeth lacking or not more than one pair оп upper jaw.
In males and sometimes in females, left upper tooth in form of
Spirallyatwistedy tusk ооо sR di. лены:
Dy AEROS ее. Narwhal, Monodon monoceros (p. 791).
Number of teeth up to 100. Neck lacking. Horny protuberances
presenton dorsum 2. ive: ase oie 12 Ae Rees ome
.. Black finless porpoise, Neophocaena phocaenoides (p. 750).
“Beak” well developed. Number of teeth exceeds 160.

Entire body black, with narrow light-colored band only along
abdomen from neck to tail; band forms rhomboidal patch on

439

25 (24).
26 (17).

27 (42).
28 (29).

29 (28).

30 (31).

31 (30).
32 (37).
33 (34).
34 (33).
35 (36).
36 (35).

37 (32).
38 (39).

39 (38).
40 (41).

breads tae. обои rors sR ее О ЗН о.
.. Northern right whale dolphin Lissodelphis borealis (р. 646).
Body black except for white lower sections and abdominal flanks
.. Southern right whale dolphin, Lissodelphis peroni (р. 651).
Dorsal fin present.
“Beak” absent or barely perceptible.
Body length exceeds 4 m. Flippers broad (length not more
than twice width) and rounded. Teeth massive (diameter more
than 2.5-2.8 cm). Body dark in color. White band on abdomen
extending as tongue on flanks in region of anal opening, its tip
directed anteriorly. Dorsal fin high, especially in males (up to
Пети ай ее Bee aid Mid fo ke Killer whale, Orcinus orca (p. 680).
Body length less than 4 m; if more, flippers narrow (length
exceeds width by more than 2.5 times) and pointed. Tooth diam-
eter not more than 2.5-2.8 cm. Body color different. Dorsal fin
smaller.
Teeth absent in upper jaw (sometimes one or two pairs seen);
two to seven pairs present in lower jaw ....................
sys APCs es EEE Risso’s dolphin, Grampus griseus (p. 696).
More than six pairs of teeth in upper jaw and more than seven
pairs in lower jaw.
Teeth in upper and lower jaws not more than 8- 13 pairs each.
Low dorsal fin in anterior half of body. Head anteriorly obtuse
with high forehead. Teeth only in anterior half of upper jaw
овал AGEL Pilot whale, Globicephala melaena (p. 702).
Relatively high dorsal fin roughly midbody. Head anteriorly
rounded. Teeth in upper jaw not exclusively in anterior half.
Entire body black. Roughly 50-51 vetebrae .................
а False killer whale, Pseudorca crassidens (р. 674).
Body dark gray, edge of jaws and region around anal opening
white: Roughly/67 -7Ojvertebrae! 335. ae. 5 ek. ВВ
LS nS. Se AEP Dwarf killer whale, Feresa attenuata (p. 719).
Teeth in upper and lower jaws more than 15 pairs each.
Body size of adult animals exceeds 2m .....................
ан Broadsnout dolphin (melon-headed whale),
Lagenorhynchus electra (p. 672).
Body size of adult animals exceeds 2 m.
Dorsal side of caudal crest with high keel. White field on
body flanks sharply set off from surrounding dark background.

586

41 (40).
42 (27).

43 (44).

44 (43).

45 (52).

46 (47).

47 (46).

48 (49).

49 (48).

50 (51).

440 51 (50).

Number of teeth in upper jaw (15-24 pairs) usually less than in
lower(2 2228 pairs). ОЕ О sty ots eae
нана BA ROS Dall porpoise, Phocoenoides dalli (p. 738).
High keel on dorsal side of caudal crest lacking. Body color dif-
ferent. Number of teeth in upper jaw (16-30 pairs) usually more
than. in lower, jaw:(16 227 (pairs) $0.20 JA. hee: 2 he
В ORE Common porpoise, Phocoena phocoena (p. 722).
“Beak” distinctly perceptible.
Symphysis of lower jaw more than one-fourth its length. Tooth
CLOWNS! BTOOVEG NEAL AE ARNT NO ЗОО ран...
И ALLL Rough-toothed dolphin, Steno bredanensis (p. 591).
Symphysis of lower jaw less than one-fourth its length. Tooth
crowns smooth.
“Beak” of moderate length or short. Body length exceeds “beak”
length (from tip of snout to beginning of corpus adiposum) by
more than 25 times. Diameter of teeth in middle of tooth row
exceeds 3 mm. :
Teeth not more than 20 pairs in upper jaw and 26 pairs
in lower. Diameter of teeth in middle of tooth row exceeds
6 mm.“Beak” dark in color. Keel absent on dorsal side of caudal
Стел nuns Bottlenose dolphin, Tursiops truncatus (p. 632).
More than 26 pairs of teeth each in upper and lower jaws. Diam-
eter of teeth in middle of tooth row less than 6 mm. If number
of teeth less (up to 22 pairs), their diameter more (up to 7 mm).
Beak white. Keel present on dorsal side of caudal crest.
Tip of “beak” white. Trunk dark-colored above right up to joint
with flippers. Number of vertebrae exceeds 84. Number of teeth
in upper and lower jaws less than 28 pairs. Pterygoid bones
separated by gap, broadened anteriorly .....................
.. White-beaked dolphin, Lagenorhynchus albirostris (р. 660).
Tip of “beak” dark. Dark color of trunk does not descend to
joint with flippers. Number of vertebrae less than 84. Number
of teeth in upper and lower jaws more than 28 pairs. Pterygoid
bones adjacent or separated by gap, not broadened anteriorly.
Number of vertebrae 74-78; ribs 12-14 pairs; teeth in upper
jaw usually 30 -32 pairs. Narrow dark band extending along body
flanks from base of flippers to lower caudal crest ...........
ELE ie WOON OLR Rn ten Da Pacific white-sided dolphin,
Lagenorhynchus obliquidens (p. 664).
Number of vertebrae 77-82. Ribs 14-15 pairs. Teeth in upper
jaw usually 35 -38 pairs. Dark band along body flanks from base

of flippers to lower caudal crest absent ...... тан Se
Atlantic white-sided dolphin, Lagenorhynchus acutus (p. 656).

52 (45). “Beak” long. Body length exceeds “beak” length (from tip of
snout to beginning of corpus adiposum) by not more than 20
times. Diameter of teeth in middle of tooth row not more than
3 mm.

53 (54). Dark band extending from flippers to chin. Palate with two
longitudinalfgroOvesime HI BAO AB, RR OSL MIS
ОЛ. Common dolphin, Delphinus delphis (р. 607).

54 (53). Dark band extending from flippers to chin absent. Palate with-
out grooves.

55 (56). Body dark gray above, white below. Length of rostrum of skull
twice length of cerebral section. Number of teeth 148258 ......
.. Long snout [spinner] dolphin, Stenella longirostris (p. 604).

56 (55). Body black or blackish above, white, whitish, or ash-gray below.
Length of rostrum less than twice length of cerebral section.
Number of teeth 32-50.

57 (58). Body black above, white below. Number of teeth 4-2? .......

.. Blue-white [striped] dolphin, Stenella coeruleoalba (р. 594).

58 (57). Body blackish above, whitish or ash-gray below. Number of
teeth м.

59 (60). Body ash-gray below. Maxillary teeth small, less than 3 mm in

diameter, Number of vertebrae 79 8.

Malay [Pan-Tropical spotted] dolphin, Stenella dubia [= айе-

пиащ] (р. 601).

60 (59). Body whitish below. Maxillary teeth large, more than 5 mm in

_ diameter. Number of vertebrae not more than 70 ...........

а: Bridled dolphin [Atlantic spotted dolphin],

Stenella frontalis (p. 602). (V.S.)

SUPERFAMILY OF DOLPHINS
Superfamily DELPHINOIDEA Flower, 1864

Cetaceans of small and medium dimensions.

The body of most of the dolphins is well proportioned. The caudal fin
bears a deep notch between the flukes. Many species have a large dorsal
fin (sometimes lacking) in the midbody. Furrows not seen on the neck.
Rostral part of head stretched into a well-distinguishable “beak” which,
however, may be poorly developed or altogether absent. The “beak” may

441

588

be sharply demarcated from the frontal portion of the head or may pass
smoothly into it. The body is monochromatic (dark or light in color)
or dark above and light-colored below, or dark with light-colored bands
that vary in disposition, shape, and number.

The zygomatic and temporal bones are poorly developed in the skull.
The supraoccipital and frontal bones overhang a small temporal fossa.
Crests are lacking on the maxillae. The pterygoid and nasal bones are
relatively small. The palatine bones are joined along the midline of the
palate and are not separated from the vomer. The zygomatic process of
the squamosal bone is highly reduced.

The symphysis of the lower jaw is not more than one-third the jaw
length. The width (spacing) of the lower and upper jaws is nearly similar.
The number of teeth varies from © to 5.

These are migratory animals living in schools. Their food is diverse:
fish (ichthyophagous whales), cephalopod mollusks (teuthophagous),
cephalopods and fish (teutho-ichthyophagous), and warm-blooded
vertebrates (sarcophagous).

The species of the superfamily live in almost all the seas of the
world right up to the icy seas in the high Arctic. Some are very widely
distributed, almost throughout the world (common dolphin and killer
whale) while others are more localized. Some species (genus Sotalia)
live in rivers (South America and southern Asia).

Members of the family are known from the Lower Miocene, having
separated from the ancient Squalodontidae. The oldest members (Argy-
rocetus, Schizodelphis, and Delphinavus) had a long and narrow rostrum
and a large number of teeth. The contemporary form of the dolphin can
be traced to the end of the Miocene-Early Pliocene.

The systematics of dolphins has been differently interpreted by var-
ious scientists.

In the superfamily of dolphins, Delphinoidea, one family—Delphi-
nidae (Tomilin, 1957), two families—Delphinidae and Monodontidae
(Hershkovitz, 1966; Yablokov её al., 1972), or three families—Monodon-
tidae, Delphinidae, and Phocoenidae (Simpson, 1945) have been
distinguished. In more detailed systems, the number of families has
risen to four—Monodontidae, Stenidae, Phocoenidae, and Delphinidae
(Andersen and Jones, 1967), or even six—Monodontidae, Delphinidae,
Grampidae, Globicephalidae, Orcaelidae, and Phocoenidae (Nishiwaki,
1966; Ridgway, 1972).

It would appear to be more correct to combine all these groups into
two families: dolphins (Delphinidae) and narwhals (Monodontidae).

442

589

Representatives of both the families are found in the USSR fauna.
The extinct forms are represented in the superfamily of dolphins by
the family Eurinodelphinidae (three genera), family Hemisyntrachelidae
(two genera), and family Acrodelphidae (six genera). Moreover, 24 gen-
era belong to the family Delphinidae (Simpson, 1945).

Some representatives of the family are of economic importance
and are hunted in large numbers. Others are almost not caught
because of their small number or for other reasons and have no
economic importance. (V.S)

Family of Dolphins
Family DELPHINIDAE Gray, 1821

Animals of small, medium, and large dimensions.

The dorsal fin in most cases is well developed. There is no neck
between the head and the trunk.

The rostrum of the skull is usually longer than the cranium. The
petrous temporal bone does not grow toward the skull but is attached
to it by ligaments. The cervical vertebrae may be fused (atlas and axis
more often fused). The anterior ribs have a double articulation with the
vertebrae. The ulnar process is well developed in the forelimb.

Open seas, coastal zone, and rivers are inhabited.

The geographic distribution of the members of the family covers the
range of the superfamily; only the northern limit falls more southward.

The family comprises 18 genera,! of which 14 have been reported (or
may still be found) in USSR waters: rough-toothed dolphins, Steno Gray;
spotted dolphins, Stenella Gray; common dolphins, Delphinus Linnaeus;
bottlenose dolphins, Tursiops Gervais; right whale dolphins, Lissodelphis
Gloger; shorthead dolphins, Lagenorhynchus Gray; false killer whales,
Pseudorca Reinhardt; killer whales, Orcinus Fitzinger; Risso’s dolphins,
Grampus Gray; pilot whales, Globicephala Lesson; dwarf killer whales,
Feresa Gray; common porpoises, Phocoana G. Cuvier; Dall porpoises,
Phocoenoides Andrews; and black finless porpoises, Neophocaena Palmer.
The remaining four genera inhabit waters outside the USSR: Sotalia Gray,
Lagenodelphis Fraser, Cephalorhynchus Gray, and Orcaella Gray.

The species of some genera are of economic importance. (V.S.)

1 According to P. Hershkovitz (1966), the placement of Lagenorhynchus electra Gray in
an independent genus, Peponocephala Nishiwaki and Norris (Electra auct—nom. praeocc.),
requires further justification.

444

442

590

Genus of Rough-toothed Dolphins
Genus Steno Gray, 1846

1846. Steno. Gray. Zoology Voyage Erebus and Terror. I, p. 30. Delphinus
rostratus Cuvier = Steno bredanensis Lesson, 1828. (V.H.)

Small dolphins reaching up to 2.5 m in length (Fig. 232).

The “beak” on the head passes smoothly into the slope of the fore-
head. The dorsal fin is triangular, high, with a notch along the posterior
edge [falcate].

The body is dark gray on the dorsum, light gray with yellowish-white
spots on the flanks, and white on the abdomen.

The rostrum is long and narrow (Fig. 233), its length more than three
times its width. The symphysis of the lower jaw constitutes over 30% of
its length. The pterygoid bones are adjacent. The postorbital processes
of the frontal bone are small. The teeth are large, with grooved crowns.
Teeth 20=27 (usually 3).

Vertebrae 65 - 66. The first and second vertebrae are fused.

The biology of the dolphins of this genus is almost unstudied. Their
population is insignificant. They live in groups and feed on squids and
fish.

The geographic range covers the temperate and warm waters of the
Atlantic and Pacific oceans and also the Indian Ocean. They have not
been reported in the waters of the USSR. However, they might be found
in the region of Kuril Islands and the Sea of Japan.

The genus comprises one species: the rough-toothed dolphin, S.
bredanensis Lesson, 1828.

Sometimes the rough-toothed dolphin, together with the genera
Sotalia Gray and Sousa Gray, is placed in a separate family, Stenidae
Fraser and Purves, 1960 (Anderson and Jones, 1967). (V.S.)

Fig. 232. Rough-toothed dolphin, Steno bredanensis (figure by N.N. Kondakov).

591

442 Fig. 233. Skull of rough-toothed dolphin, Steno bredanensis (figure Бу М.М. Kondakov).

ROUGH-TOOTHED DOLPHIN?

Steno bredanensis Lesson, 1828

1817. Delphinus rostratus. Desmarest. Nouv. Dict. Hist. Nat., 9, p. 160.
Bretagne, France. Nom. praeocc.

1823. Delphinus frontatus. G. Cuvier. Rech. ossemens foss., 5, p. 278. Por-
tugal. Description does not conform to the rules of nomenclature.

1828. Delphinus bredanensis. Lesson. Hist. Nat. mamm. oiseaux decon-
verbés depuis 1788. Substituted for Delphinus rostratus Desmarest,
1817. (У.Н.).

Single species of the genus. The body is well proportioned. The “beak”

is somewhat compressed laterally. The dorsal fin lies roughly midbody.

The flippers are broad at the base.

* Russian name corresponds to the English name. (V.H.)

446

592

A sharp demarcation of different colors is a characteristic feature of
body coloration. The snout is sometimes white.

The adult male measures 2.23 m in length and weighs 102 kg. (У.5.)

Transgressions and habitation in our waters of the Sea of Japan and
near the Kuril Islands are possible (Fig. 234).

Outside the USSR, this dolphin is found in the Pacific Ocean south
of the strip Japan—Hawaiian Islands—California; in the Atlantic Ocean
from the southern part of the North Sea (Holland) and waters of France
in the east and Virginia in the west to roughly 40° $ lat. (Tristan da
Cunha, Argentina); and in the Indian Ocean from the northern coasts
and Java to the Cape of Good Hope.

Geographic variability has not been established.

Biology not known. Lives in small schools. (V.H.)

Genus of Spotted Dolphins
Genus Stenella Gray, 1866

1864. Clymene. Gray. Proc. Zool. Soc. London, p. 237. Delphinus
euphrosyne Gray, 1846. Nom. Ргаеосс.

1866. Stenella. Gray. Proc. Zool. Soc. London, p. 214. Steno attenuatus
Gray, 1846.

1866. Euphrosyne. Gray. Ibid., p. 214. Clymene euphrosyne Gray. Nom.
ргаеосс.

1880. Prodelphinus. Gervais. In: Van Bénéden et Gervais. Ostéographie
des Cétacées, p. 604. Substituted for Clymenia Gray, 1864. (V.H.)

Small dolphins; body length up to 2.7 m.

The body is well proportioned. The dorsal fin is located midbody and
its apex curved backward. The flippers are crescent-shaped. The “beak”
is long and narrow. The lower jaw is slightly longer than the upper.

The body coloration differs markedly from species to species and,
further, is subject to considerable individual variation. The dorsum and
upper sections of the body flanks are usually dark; the lower portion of
the body flanks and abdomen are light in color.

The premaxillae are curved in the rostral part of the skull. The ptery-
goid bones are adjacent. Longitudinal grooves are absent or very small
on the palate. The symphysis of the lower jaw constitutes less than one-
fifth the length of the jaw itself. The teeth are small, sharp, and 34-65
in each half of the jaw.

The cerebral section of the skull is broad in the occipital region
and flat in the frontal region. The structural features of the skull of the
spotted dolphins on the one hand are similar to the common dolphins,
Delphinus, and on the other to the genus of bottlenose dolphins, Tursiops.

ul uy
yy et
ЕЕ В

Tas:
Ant
Peay

int]! Ih
ПИН
ОКА ОА

il

| Lent
1

ий |

|

|
|.
||

Л

1
|

1000 0 1000 2000 3000 4000 5000 km

О
НИТИ :
НИ

НН

Fig. 234. Range of rough-toothed dolphin, Steno bredanensis (У.А. Arsen’ev).

443

594

The common features of spotted dolphins and common dolphins are the
long rostrum, slightly exceeding the cerebral section of the skull, the
large number of very small teeth, and premaxillae usually confluent with
the rostral part of the skull. However, as in bottlenose dolphins, spotted
dolphins do not have longitudinal grooves on the bony palate.

Long, thin, spinous and transverse processes are characteristic of the
vertebrae. The cervical vertebrae may be fused. The sternum is T-shaped
and consists of four sections. Ribs 15 - 16 pairs. In the flippers, the second
and third digits are the longest (2nd one longer).

Biology almost not known.

Its distribution (Fig. 235) ranges from the cold to the tropical waters
of the Pacific and Atlantic oceans (including the Mediterranean Sea),
from the Bering Sea and Greenland to the Cape of Good Hope, Cape
Horn, and Australia, and the Indian Ocean. Not caught in our waters.

The composition and systematics of species of the genus have not
been adequately studied. Ten species are usually recognized but their
number seems to have decreased: (1) S. asthenops Cope, (2) S. clymene
Gray, (3) S. coeruleoalba Meyen, (4) S. crotaphiscus Cope, (5) S. dubia
G. Cuvier, (6) S. frontalis G. Cuvier, (7) S. graffmani Lonnberg, (8) S.
longirostris Gray, (9) S. malayana Lesson, and (10) 5. pernettyi Blainville.*

Only the blue-white dolphin, S. coeruleoalba, is encountered in the
waters of the USSR while the appearance or residence of S. dubia, S.
frontalis, and S. longirostris is possible. (V.S.)

BLUE-WHITE [STRIPED] DOLPHIN
Stenella coeruleoalba Meyen, 1833

1833. Delphinus coeruleo-albus. Meyen. Nova. Acta Leop.-Carol., 16, 2,
р. 609. Near La Plata estuary.

1846. Delphinus styx. Gray. Zoology Voyage Erebus and Terror, 1, p. 39.
Atlantic Ocean around South Africa.

1846. Delphinus euphrosyne. Gray. Ibid., p. 40. North Atlantic, waters of
England.

1848. Delphinus lateralis. Peale. U. S. Explor. Exped., Mammalia, p. 34.
Pacific Ocean at 13°58’ N lat. and 161°22’ W long. (V.H.)

Diagnosis

Body length up to 260 cm. The dorsal surface is black and the ventral sur-
face white. A narrow black band runs from the eyes to the anal opening

3 In alphabetical order, after Hershkovitz (1966).

a т, TT | ТИ.

ih i es И
мн a el i О

ИИ р.
И, |

i i!

yt i!
1 И Nh!

.

м

iN И
it

0

1000

1000 2000 3000 4000 5000 Кт

Fig. 235. Range of spotted dolphins, Stenella (V.A. Arsen’ev).

445

447

447

596

and from the eyes to the base of the flippers. The flippers are small.
The length of the rostrum is less than double the length of the cerebral
section of the skull. Teeth 4-28. Vertebrae about 76-79. (V.S.)

Description

The main body measurements of female and male blue-white dolphins
(Fig. 236) caught in the waters of Japan (Tomilin, 1957) are respectively
(in cm): body length 200 and 196 (may go up to 2.7 m); distance from
tip of snout to base of flippers 48 and 50, up to blowhole 34 and 35, up

to anterior edge of dorsal fin 93 and 94, and up to anal opening 137 and

138; height of dorsal fin 17 and 15, length of the base of dorsal fin 31
and 28; width of caudal flukes 67 and 44; and length of flippers 27.

Of the more than 3,000 blue-white dolphins caught at the end of
November, 1967 at village Kawana (Honshu Island), a random sample
of 75 adults (body length 170 to 256 cm) comprised 59 females and
16 males; 71 newborn calves (body length 97 to 144 cm) comprised 36
females and 35 males (Sokolov and Kasupa, 1969). Among the females
studied, six with a body length ranging from 184 to 217 cm (184, 199,
206, 208, 213, and 217) were immature; two with a body length of 139
[sic.] and 213 cm had not given birth before but were ready for fertiliza-
tion (with large follicles in the ovaries); and nine had calved before (they
bore traces of the corpus luteum of pregnancy in the ovaries) but were
now barren or had lost their calves. The body length of these last nine
females was: 217 cm (two), 224, 226 (two), 229, 231, 232, and 239 cm.
The ovaries of two of them (body length 232 and 239 cm) contained
large follicles. Of the remainder, 40 females were lactating (the status
of two females could not be ascertained) and the ovaries of three con-
tained mature follicles. The body length of these 40 females was (in cm):
209, 216 (two), 218 (two), 219, 221 (three), 222, 223 (four), 224 (four),

Fig. 236. Blue-white dolpin, Stenella coeruleoalba (figure by М.М. Kondakov).

597

225, 226 (four), 227 (three), 228 (two), 229 (three), 230, 231 (three), 232

(two), 233, and 234 (two).
The body length of immature males varied from 170 to 214 cm (170,

176, 184, 191, 195, and 214 cm) and of mature males from 224 to 256 cm:

224, 228, 232 (two), 233, 235, 237, 239, 241, and 256.
The condylobasal length of the skull (Fig. 237) of these dolphins

448 from La Mancha was (in cm): 47, length of rostrum 29, width of rostrum
at base 12, length of lower jaw 41, and length of symphysis of lower jaw
4 (Tomilin, 1957). Phalangeal formula: Ц, Пу-1о, Ша, [V4, and У, _,

(V.S.)

4
ра
of
РА
Я
Я
7
4

ERAS
А
х

447 Fig. 237. Skull of the blue-white dolphin, Stenella coeruleoalba (figure Бу М.М.
Kondakov).

598

a
Be Aue
‘oe

{| и

250 0 250 500 750 1000 km
ЕЕ

BN]

150 160 170

130 140

448 Fig. 238. Range of the blue-white dolpin Stenella coeruleoalba in the USSR (V.A.
Arsen’ev).
Geographic Distribution

This dolphin inhabits the warm and temperate belts of the Atlantic and
Pacific oceans. Studies of its distribution are highly schematic.

Geographic Range in the USSR

Waters of the Pacific Ocean around the Kuril Islands and possibly the
waters of the Bering Sea (Fig. 238).

Geographic Range outside the USSR (Fig. 239)

Strays in the Atlantic Ocean northward up to the coasts of Great Britain,
Shetland and Orkney Islands, and southern Greenland; known in the
waters of Quebec and Nova Scotia. Not reported in the Northern, Baltic,

1000 0 1000 2000 3000 4000 5000 km le
\ `

150

Fig. 239. Species range of the blue-white dolphin, Stenella coeruleoalba (V.A. Arsen’ev).

449

450

600

or Barents seas.* Distributed southward up to Rio de La Plata along the
South American coast and up to the southern tip of Africa. Lives in
the Mediterranean Sea. In the Pacific Ocean, known from the coasts
of Japan and the Kuril Islands in the west and from the Bering Sea,
British Columbia, states of Oregon and Washington in the east up to
New Zealand. [51с.] (V.A.)

Geographic Variation

Our knowledge of the geographic variation of the blue-white dolphin
is extremely scant and this aspect has generally not been discussed in
the world literature. In our literature on the fauna of the USSR, the
existence of two subspecies is sometimes acknowledged (Tomilin, 1957,
1962). These are: the nominal form in the seas of the Far East and the
Atlantic form, S. c. euphrosyne Gray, 1946, for “the Baltic Sea, possibly
the Barents 5еа”. The main difference between them, in addition to some
skull features, lies in coloration (band from eyes to base of flippers single
in the former and double in the latter). Sometimes these two forms are
treated as independent species. Their nomenclature differs according to
different authors.

As pointed out above (see “Geographic Distribution”), the existence
of the Atlantic form (euphrosyne) in our waters is just not possible.
According to the convention prevailing in our literature, the name S.
c. coeruleoalbus Meyen, 1833 may be retained only tentatively for the
blue-white dolphin found in our Far Eastern waters. However, it would
be better to use the binary name. It is quite possible that the use of the
name styx Gray, 1846 might be more correct.

The systematics and nomenclature of the species and forms of the
genus Stenella are complex and in a high state of confusion (compare,
for example: Ellerman and Morrison-Scott, 1951, 1962; Tomilin, 1957,
1962; Hall and Kelson, 1959; Hershkovitz, 1966; and others). A radical
review of the subject, based on substantial evidence, is required. (V.H.)

Biology

The biology of the blue-white dolphin has almost not been studied and
practically no observations have been made on the behaviour of these
animals at sea.

‘Information and assumptions on habitation in these seas (Tomilin, 1957, 1962) have
not been confirmed in the latest compilations (Ellerman and Morison-Scott, 1951, 1966;
von den Brink, 1958; Koval’skii, 1964; Bobrinskii et al., 1965;- Hershkovitz, 1966; Siivonen,
1967). (V.H.)

451

601

The largest of the measured embryos was 103 cm long. All the young
dolphins 120-150 cm long were comparatively recently born since their
stomach contained exclusively milk or milk together with small quanti-
ties of the remnants of squids. The smallest female with a corpus luteum
in the ovary measured 215 cm in length; apparently females of this size
attain maturity. Among animals of this length (or slightly larger), the
ovaries and testes begin to lengthen rapidly. Among dolphins caught in
November, 1956, 12 were immature females, 72 gestating, and 27 simul-
taneously gestating and lactating. The oldest of these dolphins was 18
years of age (Nishiwaki and Yagi, 1953).

Sufficient numbers of the blue-white dolphin are caught in the
coastal waters of some regions of Japan and China. Hunting is done
in small crafts and with small-bore harpoon guns. The kill is sold in fish
markets. Economic importance is very negligible. (V.A.)

MALAY [PAN-TROPICAL SPOTTED] DOLPHIN
Stenella dubia [= attenuata] С. Cuvier, 1812

1812. Delphinus dubius. G. Cuvier. Ann. Mus. Hist. Nat., Paris, 19, p. 9.
French waters. [nomen dubium]

1826. Delphinus malayanus. Lesson. Voyage autour du monde...,
Zoologie, I, p. 184. South China Sea, strait. between Java and
Kalimantan (Borneo). [nomen dubium]

1846. Steno attenuatus. Gray. Zoology Voyage Erebus and Terror, Г,
p. 44. Type locality not established. (V.H.)

Body length up to 183 cm. The color is blackish above and ash-gray

below (including the caudal flukes). The flippers are large. The length

of the rostrum is less than double the length of the cerebral section of

the skull. Teeth 3-44. Vertebrae 79-81.

The body is well proportioned, resembling the common dolphin

(Fig. 240). The forehead is low and long. The color is pale bluish-black

451

Fig. 240. Malay [Pan-tropical spotted] dolphin, Stenella dubia (figure by М.М. Kondakov).

602

above, aSh-gray ог grayish-white below. Numerous minute whitish ог
grayish, sometimes pinkish spots are scattered over the body. A dark
band runs from the flippers to the corner of the mouth. The upper jaw
and lateral sections of the lower jaw are black. The tip of the “beak” is
white. A band runs forward from a black spot around the eye.

The main body measurements (as percentage of body length) of
males with a body length of 165 - 208 cm and females 180-201 cm (Nishi-
waki, 1966) are respectively: from end of upper jaw to blowhole 14.2 - 17.1
and 14.2-17.7, from fork between caudal flukes to posterior edge of
dorsal fin 39.2-40.5 and 39.6-42.8, from caudal fork to anal opening
25.2-26.5 and 25.2-27.2; length of flippers 14.4-15.1 and 12.0- 15.2,
maximum width of flippers 5.0-5.3 and 4.2 -5.3; length of base of dorsal
fin 12.1-14.2 and 12.4-14.2, height of dorsal fin 8.2-9.7 and 7.8-9.4;
and width of caudal flukes 21.6 -22.5 and 23.5.

The main dimensions of the skull (as percentage of condylobasal
length) of these dolphins with a condylobasal length of skull 381 - 428 mm
(Nishiwaki, 1966) are: length of rostrum 58.5 - 61.5 and its width at the
base 21.5 - 23.3; interorbital width 38.8 - 42.8; length of lower jaw sections
84.5 - 90.3, and length of symphysis of lower jaw 16.0 - 17.3. (V.S.)

The presence of this species in USSR waters has not been estab-
lished. Presence in or transgressions into the waters of the Sea of Japan
and around the Kuril Islands are possible.

Outside the USSR, this species is found in the South Atlantic Ocean
up to Cape Horn and Cape of Good Hope, the Indian Ocean, the Pacific
Ocean from New Zealand to the South China Sea, Hawaiian Islands, and
waters of Japan. (V.H.)

Biology is not known.

BRIDLED [ATLANTIC SPOTTED] DOLPHIN?
Stenella frontalis G. Cuvier, 1829

1829. Delphinus frontalis. G. Cuvier. Regne Animal, I, р. 228. Cape
Verde Islands, Atlantic Ocean. (V.H.)

Body length about 180 cm. Color of the dorsal section blackish, changing

over to grayish on the flanks. Abdomen whitish. The dorsum has light-

colored and the abdomen dark spots. A dark-colored band runs from the

corner of the mouth toward the flippers (Fig. 242). The eyes are encircled

452 by black rings. The flippers are small. The length of the rostrum is less

5 The Russian name is given here.

603

452 Fig. 242. Bridled dolphin, Stenella frontalis (figure by М.М. Kondakov).

than double the length of the cerebral section of the skull (Fig. 243).

35-44 37-38
Teeth 32-7 (usually 35-35).

Vertebrae: 7 cervical, 15 thoracic, 19 lumbar, and 36 caudal; total
77. Phalangeal formula: I,-3, По, Шу, 1V3, and V5. (V.S.)

Its residence has not been established in the waters of the USSR.
Habitation in or transgressions into the Sea of Japan are possible.

Outside the USSR, it is found in the Atlantic Ocean (from Cape
Verde Islands to Cape of Good Hope and from North Carolina to

22777 777 a

RUIN SS Us

453 Fig. 243. Skull of the bridled dolphin, Stenella frontalis (figure by М.М. Kondakov).

455

604

Florida), Mediterranean Sea, and Indian Ocean. It has been reported
in the waters of Japan and the Korean peninsula (Cheju-do Island and
Sea of China). (V.H.)

Biology is not known.

LONG-SNOUT [SPINNER] DOLPHIN
Stenella longirostris Gray, 1828

1828. Delphinus longirostris. Gray. Spicil. Zoologica, 1, p. 1. Type locality
not established.

1846. Delphinus alope. Gray. Zoology, Voyage Erebus and Terror, 1,
р. 52.

1846. Delphinus roseiventris. Wagner. Schrebers Sauge thiere, 7, pl. 360.
(У.Н.)

Body length roughly up to 213 cm. Dorsal side of body dark gray.

The abdomen is white with random gray spots. The flippers are small

(Fig. 244). The length of the rostrum is double the length of the cerebral

section of the skull (Fig. 245). The bony palate has very small longitudinal

grooves. Teeth 2-56 (usually 22-25). Vertebrae 72-73. (V.S.)

Its residence has not been established in USSR waters. Habitation
in or transgressions into the Sea of Japan are possible.

Outside the USSR (Fig. 246), it is reported in the Atlantic Ocean
south of the Bahamas and Senegal up to Cape Horn and Cape of Good
Hope, the Indian Ocean (Ceylon), and the Pacific Ocean from Mexico
and Japan to Australia. (V.H.)

Biology is not known (Fig. 247).

Genus of Common Dolphins
Genus Delphinus Linnaeus, 1758

1758. Delphinus. Linnaeus. Syst. Nat., ed. X, 1, p. 77. Delphinus delphis
Linnaeus, 1958.
Small cetaceans with body length up to 2.6 m.

The “beak” is long. The dorsal fin is high and crescent-shaped. The
narrow flippers are also curved.

Body coloration is subject to considerable individual variation but
is usually black above, gray with dark- and light-colored bands on the
flanks, and light-colored below.

The rostrum is 1.5 to 2 times longer than the cranium and bifurcated
anteriorly. Two longitudinal grooves occur on the bony palate. The
pterygoid bones are adjacent throughout their length. The supraoccipital

453

453

605

Fig. 245. Skull of the long-snout delphin, Stenella longirostris (figure by М.М. Kondakov).

crest is asymmetric: its right half is larger than the left. The lower jaw is
slightly longer than the upper. The teeth are conical and number 2=%.
Vertebrae 70-75. Phalangeal formula: I,_3, IIg_,, III;_7, IV>-4, and
V,-2. The 2nd and 3rd digits are the longest. Ribs, up to 15 pairs. The
sternum is complete and T-shaped.

These are pelagic forms living on fish. The females are capable of
annual reproduction. Gestation extends for roughly 10 months and lac-
tation for four months. The females may ovulate and can be fertilized
during the lactation period.

These animals are distributed in all the oceans and most of the seas
in the temperate and warm parts of the world.

The genus comprises a single species, the common dolphin, D.
delphis Linnaeus, 1758. Some scientists (for example, Nishiwaki, 1972)
recognize two more species: D. capensis Gray, 1828 and D. Бата Dall,
1873.

These animals could be of economic importance. (V.S.)

606

=
© 5
——
|
1

ТИТ
и ин
ht 4:4
' Г, о 9
НЕ Nhe

1 wit
И

И

atures
AA yyy

АМАН
НИИНИИ ВОЙ Г

ие

1600 2000 3000 4000 5000 km

Wey rely!
НИИ
yu Vi! У
al il itil

8 =

Fig. 246. Range of the long-snout dolphin, Stenella longirostris (V.A. Arsen’ev).

454

455

456

607

Fig. 247. Long-snout dolphin, Stenella longirostris, at sea (figure by М.М. Kondakoy).

COMMON DOLPHIN
Delphinus delphis Linnaeus, 1758

1758. Delphinus delphis. Linnaeus. Syst. Nat., ed. X, I, p. 77. Waters of
Europe (“Oceano Епгорео”).

1860. Delphinus algeriensis. Loche. Rev. Mag. Zool. Paris, 12, p. 474.
репа.

1873. Delphinus bairdii. Dall. Proc. California Acad. Sc., 5, р. 12. Point
Arguello, Santa Barbara, California.

1883. Delphinus delphis var. curvirostris. Riggio. Nat. Sicil., 2, p. 158.
Mediterranean Sea.

1935. Delphinus delphis ponticus. Barabash-Nikiforov. Byull. Mosk.
Obshch. Ispyt. Prirody, 44, p. 249. Yalta, Black Sea. (V.H.)

Diagnosis
Single species of the genus.
Description

Well-proportioned animals with a moderately elongated body, distinctly
demarcated long beak, and relatively high dorsal fin (Fig. 248).

The body color is a combination of black and white and intermediate
shades of the same. The upper part of the body is dark-colored and the
underside is white; two gray elongated fields and one to three gray bands
extend from the zone of the anal opening to the anterior half of the body
along the flanks. One dark-colored band runs from the chin to each of

608

ИЕ Ms
< ИИ / YI:
Wy = Pas

LY pope ИИ 7772

456 Fig. 248. Common dolphin, Delphinus delphis (figure by М.М. Kondakov).

457 Fig. 249. Variation in coloration of the common dolphin, Delphinus delphis
(figure by N.N. Kondakov).

the flippers. A narrow dark-colored strip joins the eyes at the bridge of
the nose. The flippers, caudal flukes, and dorsal fin are dark-colored.
Color combinations of Black Sea dolphins vary markedly (Fig. 249). Far
Eastern common dolphins are differentiated from their Black Sea and
Atlantic counterparts by distinctly visible lateral and diagonal bands and
by the fact that the dark color of the upper part of the body flanks is
sharply demarcated from the light-colored underside (Tomilin, 1957).

457

609

In general, however, coloration is similar among animals from different
parts of the range.

The skin cover has a thin dermal layer with loose fascicles of collagen
fibers. At the boundary with the subcutaneous musculature, elastin fibers
and even small fascicles are intertwined.

The usual number of vertebrae is: cervical 7, thoracic 14, lumbar 21,
and caudal 31: total 73. Cervical vertebrae are often fused. Ribs 13-15
pairs; their number may differ on the left and right sides.

An instance has been reported of rudiments of the posterior limbs
being detected along the flanks of the urogenital slit among some females
caught in the Yalta region (Sleptsov, 1939). The rudiments are in the
form of triangular lobes with a height of 3.4 cm (right) and 1.6 cm
(left). Each contains two bony members (rudiments of femur and tibia),
three cartilages on the right and two on the left (rudiments of tarsus,
metatarsus, and phalangeal digits).

The main measurements (as percentage of body length) of adult male
common dolphins (six) with a body length of 155 to 190 cm and young
females (three) with a body length of 142 to 161 cm (Tomilin, 1957) are
respectively: distance from tip of snout to anterior edge of dorsal fin 47.1
and 48.0, up to blowhole 17.9 and 19.6 (two measurements each), from
anal opening to notch between caudal flukes 27.7 (three measurements)
and 28.6 (two measurements), length of flippers, 16.3 and 17.0, maximum
width of flippers 6.2 and 6.1, height of dorsal fin 10.3 and 10.1, length of
dorsal fin along the base 15.9 and 14.3, and width of caudal flukes from
fork to corner 11.8 and 12.3.

The most common body length of the Black Sea dolphin is
160 - 170 cm: males 161.8 - 165.8 cm and females 158 - 158.7 cm; maximum
length 210-219 cm. Dolphins with a body length exceeding 200 cm are
extremely rare: of 38,000 or more dolphins examined, only 26 were
longer than 200 cm (Barabash-Nikiforov, 1940). Atlantic and North
Pacific common dolphins are larger than their Black Sea counterparts:
the maximum body length of the former reaches 258 and 259 cm
respectively.

Individual variation in skull dimensions (Fig. 250) is very significant
(Kleinenberg, 1956a). The height and width of the occipital foramen, size
of the parietotemporal fossa, length of the basilar bone [OCHObHbI
KOCTD, basilar process], height of the rostrum, as also the number of
teeth are highly variable. The total and base length of the skull and the
height of the occipital region are relatively stable. Sexual dimorphism
of the skull is poorly manifest (Tryuber, 1937). Among males, the skull
length relative to body length is less, the cerebral section and rostrum
shorter, while the frontal section on the contrary, is greater than among

610

5 YZ

Pw \
. ot
мА й
«Я 2

457 _ Fig. 250. Skull of the common dolphin, Delphinus delphis (figure by М.М. Kondakov).

females. With the jaws closed, the rostral portion of the skull of males
resembles a broad obtuse wedge while in females it is more pointed and
elongated.
The main dimensions of the skull (as percentage of total length of
skull) of 23 adult males with an average total skull length of 39 cm
and 11 adult females with an average skull length of 38 cm (Black Sea;
Kleinenberg, 1956a) are respectively: length of frontal section 72.3 and
71.7, cerebral section 27.7 and 28.3, height of occipital region 31.6 and
31.2, width of cranium 35.7 and 35.5, length of rostrum 60.2 and 60.2,
width of rostrum at the base 21.0 and 20.6, height of rostrum at the last
tooth 8.0 and 7.7, and length of lower jaw 84.7 and 84.7.
Age-related changes in the skull (Tryuber, 1937; Barabash-Nikiforov,
458 1940; Tomilin, 1957) are significant. In the newborn, the cranium does
not have crests, the skull bones are not completely fused, and the teeth
have not cut through. In the first year, sutures between the bones are
closed, crests are formed, and the teeth cut through (45 - 48 in each jaw).
On taking to independent feeding, the muscular apparatus strengthens
due to the greater dimensions of the crests and projections and the
rotundity of the skull disappears. Skull growth is most vigorous in the
first two years; increase in total length of the skull mainly occurs in
the frontal section. At this time the dental system is fully developed. By
three years of age, the skull is finally formed. Crests in the skull attain
maximum development by four to five years of age. (V.S.)

460

611

Geographic Distribution

These animals inhabit the temperate and warm waters of the Atlantic,
Indian, and Pacific oceans in both the hemispheres.

Geographic Range in the USSR

Black Sea, Sea of Japan, Sea of Okhotsk, Pacific Ocean around the Kuril
Islands, Bering Sea, waters of southern Kamchatka, and Commander
Islands (Fig. 251).

These dolphins may be present in our waters of the Baltic Sea since
they transgress (albeit rarely) into the southern part of the sea and even
up to the estuary of the Visla (Koval’skii, 1964). Transgression into the
Barents Sea is also possible.

Geographic Range outside the USSR

In the North Atlantic, it reaches the coasts of northern Norway, Iceland,
the southern coasts of Greenland, and is encountered in Newfoundland
and the Gulf of St. Lawrence (Fig. 252). It is quite common in the
Mediterranean Sea and is encountered on the Canary Islands and along
the entire African coast right up to the Cape of Good Hope and Tristan
da Cunha Island. In the western part of the Atlantic, it is known in the
Bahamas, the Gulf of Mexico, the Caribbean Sea, and in the waters of
South America up to 45° $ lat. It inhabits the entire Indian Ocean and
is known in the south along the coasts of Tasmania and Australia. In the
Pacific Ocean, it inhabits from Kamchatka and British Columbia south
to the Australian and New Zealand coasts and along the coasts of Chile
up to 45° $ lat. (V.A.)

Geographic Variation

This aspect has not been adequately studied. Based on the main dimen-
sions and coloration of the animals, a few subspecies have been described.
Of these, three inhabit or transgress into the waters of the Soviet Union.
1. Atlantic common dolphin, D. d. delphis Linnaeus, 1758.

Largest of the forms, with an average body length of adults at 225 cm.

Color variation is very significant.

Its presence in the Barents and Baltic seas is possible.

Outside the USSR, it is found in the waters of the Atlantic Ccean
in the Northern as well as Southern hemisphere.

612

458

lite
И

Fig. 251. Range of the common dolphin, Delphinus delphis, in the Pacific Ocean
waters of the USSR (V.A. Arsen’ev).

2. Black Sea common dolphin, D. 4. ponticus Barabasch, 1935.

Smallest in skull and body dimensions; body length of adults averages
204 cm.

Color variation relatively minor.

Black Sea.

Outside the USSR, it is found in the waters of Turkey, Bulgaria, and
Rumania.®
3. Pacific common dolphin, D. d. bairdi Dall, 1873.

In body dimensions it occupies an intermediate position between the
two preceding forms. The average body length of adults is 208 cm; thus
it is larger than the Black Sea but smaller than the Atlantic form.

Waters of the Pacific Ocean.

$ The characteristics of the Black Sea and Atlantic common dolphins have been analyzed
and compared. However, no comparison with the Mediterranean populations has been done,

which is essential. (V. H.)

SH ООО АИ И 1 at af (ny Milage via

Perey lity
th | | a | В МИ | ee UI
al ПИ | | И

{lil
и И Mh | a hi! И
NAL i

| 1 И НИ ih
ANIM и м Hea

и м yh Hi nt | i i | ]

rv С hoe
И Pye И i
OSS Ss,
a Sb 0

ПО ar “il |

(ty tlaytl
ian
{1 ЩИ || ait

ТТНТ т]
Avs ЩИ uh ТИ И

| И |

Л
2 a
| mn i | | || и IP
(ih i it т

sill
ute an eight |

Г i НН |
ИИ nite a) I ee it |
И И НИ и т с
И | nH

Fig. 252. Range of the common dolphin, Delphinus delphis (V.A. Arsen’ev).

46

—

614

Outside the USSR, it is found throughout the North Pacific Ocean.
(V.A.)

Biology

Population. The total population of the Atlantic and Pacific subspecies
has not been established. The population of the Black Sea common
dolphin, in spite of the fact that this subspecies inhabits a comparatively
small landlocked water body, apparently reached 1.5 to 2 million, with
the catch in some years in the Black Sea exceeding 100,000. At present,
however, its population is considerably reduced, and probably does not
exceed 200,000 to 250,000 animals.

Food. The food of the Black Sea common dolphin is fish, crus-
taceans, and mollusks. The common dolphin feeds on anchovy (Engraulis
encrosicholus), pelagic pipefish of the family Syngnathidae (evidently,
Syngnathus schmidti and Sighonostomus typhle), sprat (Sprotella spretus
phalaerica), haddock (Gadus euxinus), scad (Trachurus trachurus), mullet
(Mullus barbatus), and bluefish (Tomnodon saltator) [Pomatomus salta-
trix]. From among crustaceans, large isopods (Idothea algirica) and the
shrimp (Crangon crangon) (sometimes stray specimens) form the food of
this subspecies; from among mollusks, Nassa recticulata, Mactra subtrun-
cata, Guoldia minima, Venus gallina, Colyptera hinensis, Modiola adriatica,
Mytilaster sp., and Fellina fabula, constitute the food of this subspecies.

Three types of fish form the food base. These are anchovy, pelagic
pipefish, and sprat, i.e., small schooling fish often forming large concen-
trations. All of the other species are rarely encountered and, in small
numbers, in the stomach of this dolphin, constituting a few tenths of a
percent of the total food. Dolphins swallow mollusks incidentally during
the intake of fish, mainly anchovy.

The food of the Black Sea dolphin changes, depending on the region
of habitation and the season of the year (Table 35). In winter and early
spring months, in the coastal zone, dolphins feed exclusively (or almost
50) оп anchovy. In summer and autumn, pelagic pipefish predominate
in terms of frequency of encounter but are of minor significance in the
entire food. Sprat in this respect plays a major role, although in terms
of frequency of encounter, it is slightly inferior to the pelagic pipefish
(Kleinenberg, 1940).

In the summer months, the common dolphin forms massive schools
in the open sea and the food composition there differs sharply from that
established for the coastal zone (Table 36).

In the coastal zone, this dolphin feeds on many species of fish but
almost exclusively on sprat in the open sea. However, from May through

615

(рзиоэ)
с ysyanig
€ yooppeH
р vayjop]
ЕТ 12145
8 Aaoyouy
ysijodid
OL 218 1эа 007 Aacyouy 001 Аоцму — — = = — ЕЕ
I yOOoppeHH © Уоррен
I вор С vayjop]
ysijedid ysyodid ysiedid
= == 6S asefad bh ева = SZ 218124
Zz Aaoysuy = — — if yeids ZI yeids SZ 1214$ TZ 1е14$
ysijedid
86 osejag = — — 85 Алоцоцу — 0 Kasoyouy — 06 Алоцоцу — 81 Алоцоиу = $Х$185010^0М
@ vaylop]
= = == 0OT — Aaoyouy 86 Алоцоиу 00 — Алоцоиу uoyy лом
С payjiop]
= = 86 Asoysuy = QOT_=— Aaoyouy 001 Алоцочу = goT.—Ss Ano youy rune g
Sa a a ee ao a a А
% wot % шэи 9% wo}jl % Wo}jl % two % mol
poof poo poo poo poo poo
ounr Key judy Че Areniqa.J Атепчег UOIBdYy

(egc61 ‘ЗлэчиэщэГУ) % ‘vag Чо эцу Jo эпох [е}5е0э ay) UY ищаюр поинио» ay} Jo poog ‘sE эает, Z9p

%

616

TINA
yooppeH
vaylop]
ysijadid
218 за

yooppeH
yeids
Aaoyouy
ysyadid
o18e[ag

ша
poo

12490120

58

%

paylop]
1214$

ysijadid
18 |э4

Ч$9эщ&
yooppeH]
Aaoyouy

ysijodid

218 |эа

wo!
poo

Jaquiaidag

яя

+

98

%

yooppeH
paylopy
yeids
Aaoyouy
ysiedid
218 1э4

Ч$3211Я
peos
Aaoyouy
ystjadid
218 |э4

шэи
poo

isnsny

NAA

85

%

Ч$3эЯ
yooppeH
вом
yeids
Aaoyouy
ysijodid
o18e[9g
yooppeH
Ppeos
yeids
AAoyouy’
ysijedid
218124

э
poo

Aqne

eer

YSIISSOIOAON

‘pluoo 5$ этавт,

617

462 Table 36. Food of the common dolphin in the coastal zone of northern Caucasus and
away from coasts (Tsalkin, 1938a)

Percentage of total number of items detected

In coastal zone Away from coasts
(S.E. Kleinenberg)

Pelagic pipefish 72.0 —
Anchovy 15.0 0.1
Sprat =— 99.6
Haddock 1.0 0.3
Idothea (slater) LES —
Scad 8.0 —
Mullet — —

Bluefish 2.0 —

August, sprat serves as the main food not only in the open sea, but also
in the coastal zone, constituting, on average, over 90% of the food for
the season as a whole. In various regions, the age composition of sprat
varies sharply in different months. In May, the stomach of the dolphins
in the coastal zone as well as in the open sea contained both adult and
young sprat, while in June in the open sea, the preference was for young
sprat and in the coastal zone of Crimea (Alushta) and the Caucasus
(Gelendzhik), mainly the adult fish. In July and August, the stomach of
all the dolphins from all the regions contained mainly young sprat. Thus
young sprat constituted the maximum specific proportion in the food of
these dolphins in the summer months (Tarasevich, 1958b).

The seasonal change in the food of the Black Sea common dolphin
is explained by the biological features of the fish consumed by it. Sprat
is a cryophilic fish and its spawning continues throughout the winter and
spring months. In the period of spawning, sprat is scattered over immense
pelagic regions of the sea and does not form significant concentrations.
In summer, however, young sprat as well as adult fish remain sufficiently
concentrated and hence serve as the main food of these dolphins at this
time. Anchovy, on the other hand, is a thermophilic fish and spawns in
the summer months. At this time it is also scattered over a large expanse
in the coastal zone as well as in the open sea and nowhere forms notice-
able concentrations. In the winter months, anchovy gathers at certain
places, 1.е., in regions of steep precipitous drops of coastal waters. In
December, massive collections of anchovy form almost simultaneously
along the southern coasts of Crimea (Balaklava-Sarych region) and the
Caucasus (Poti-Batumi region). It is in this period that anchovy serves
as the main food of these dolphins (Freiman, 1950).

463

618

Thus the main food items of the common dolphin lead a pelagic way
of life, are relatively small in size, and form dense concentrations.

Information on the food of Atlantic and Pacific subspecies of the
common dolphin is very scant. Fish apparently serve as their main suste-
nance. Sardines, flying fish, anchovy, herring, mackerel, and gray mullets
have been indicated as food items. Sometimes crustaceans and mollusks
(bivalves and cephalopods) are encountered. In a dolphin caught close
to Corsica, the stomach contained four specimens of Enoploteuthis mar-
garitifera and three of Chirteuthis veranyi in addition to Loligo vulgaris,
Todarodes sagittatus, Onychoteuthis lichtensteini, Heteroteuthis dispar (?),
and three squids identified as Ctenopteryx cyprinoides (Tomilin, 1957).
The stomach of a dolphin caught on May 22, 1957 in Newfoundland
contained the remains of 20 Atlantic shortfin squids, Шех illecebrosus,
which are quite numerous in these waters in summer (Sergeant, 1959b).

Daily activity and behavior. No changes have been observed in the
daily activity of dolphins. Fast and impetuous movements are charac-
teristic of the common dolphin, as in the case of many other species.
Feeding mainly on pelagic fish, the common dolphin does not go very
deep and short respiratory intervals are characteristic. It usually emerges
onto the water surface every 0.5 -1 min, less often every 1.5-3 min. It can
remain submerged in water for a maximum of 5 min. The duration on
the surface is reckoned at a few tenths of a second. The common dolphin
does not produce blows and splashes water sideways only slightly.

Foraging dolphins move slowly and submerge steeply in the water.
When feeding on fish schools confined roughly at one place, the dolphins
dive almost vertically. They cannot perhaps dive deeper than 70 m since
they never descend below the lower seine ropes of 70-m high fishing
nets.

A “traveling” dolphin swims rapidly, easily overtaking ships sailing
at 15-18 miles an hour. It swims right on the surface of the water and
at short intervals (at the time of inhalation/exhalation) leaps powerfully
out of the water, often over a span of up to 10 m (Fig. 253). The habit
of dolphins running behind ships is interesting. Small schools very often
accompany a ship for a few hours, sometimes on one side, sometimes
on the other, straying off and again rapidly closing in. Dolphins usually
love to play before the very stem, leaping out of the water into the surf
formed by the ship. When moving in schools, the dolphins do not leap
simultaneously; thus at any given moment a few animals can be seen on
the surface of the sea (Kleinenberg, 1956a).

The females are greatly attached to their calves. Two females swam
for a long time around a sweep net enclosing a water body in which their
calves were held.

463

464

619

Fig. 253. Leaping common dolphins (photograph by S.E. Kleinenberg).

The auditory faculty is well developed among dolphins. Common
dolphins are frightened of sharp sounds, which in fact is taken advantage
of when trapping them in sweep nets. A school of dolphins is chased
away from the wall of the net using a “telephone”, 1.е., striking stones
one against the other in the water (Tomilin, 1957). Vision is not as well
developed as hearing. Dolphins can probably see in water no more than
a few tens of meters. They produce extremely diverse sounds—whistling,
squeaking, and crackling. Some sounds are produced at high frequencies,
beyond the range of human perception. Such sounds are used by the
dolphin for echolocation, enabling excellent self-orientation in an aquatic
environment.

The age and sex composition of schools of dolphins vary at different
times of the year. In winter, two types of schools have been noticed.
The first type, called the winter female groups, consists of 40 to 50%
mature females, some gestating, some barren, and those that have only
recently attained maturity (lactating females are very few) and 50 to
60% immature animals of both sexes. Immature females in these schools
represent all age groups while immature males are usually young. Much
larger males, approaching maturity, are found in small numbers. Schools
of the second type are called male groups and contain predominantly
males, to the extent of 90% of the total number of animals in a group in
January-February and about 80% in March-April. The females in these
schools, at best, account for 10-20% and comprise mainly the barren
with only 4-5% being immature. Most of the males in the school are

620

mature animals while the immature are mainly represented by older age .
groups. х

The following types of schools have been established for the spring-
summer months: with the approach of the en masse calving season
(March-April), the gestating females begin to separate from the female
schools and gradually form individual precalving schools.

The maximum number of such schools is formed in May and June but
the precalving schools invariably contain immature animals too, which
sometimes constitute up to 30% and even 45% of the school’s strength.
As the gestating females leave the winter female schools, the remain-
ing dolphins form independent groups. In some cases, they consist of
mainly the immature animals of both sexes and may be called schools of
immature dolphins. Such schools formed in March represent long-time
formations and are seen later throughout the year. If, however, the win-
ter female schools comprise mainly young and barren females, with only
a few gestating ones, after the separation of the latter, residual female
groups are formed with predominantly barren and just recently matured
females. These groups remain active for a short duration, i.e., only in
early spring.

Females of the Black Sea common dolphin in the period preceding
calving concentrate in regions with more favorable meteorological con-
ditions, in the so-called calm zone. The animals find such conditions in
the open sea far away from the coasts. It is here that precalving groups
remain and are gradually transformed into schools of calves. The period
of parturition is very prolonged in this dolphin and hence the number of
gestating females in the precalving groups diminishes gradually while the
number of lactating females concomitantly increases. Later, the lactating
females begin to predominate and such groups can be called nurseries.
Nevertheless, they invariably contain immature dolphins too and their
relative proportion is even slightly more than in the precalving groups.
Schools of calves are encountered mainly in the second half of June
and in July. In August, and more so in September, they become quite
rare.

Females begin to gather with the winter male schools in spring and by
April-May such groups are transformed into breeding schools consisting
of 60-80% mature males and females in roughly an equal proportion.
Such groups are most often seen in the period of en masse mating (in July
and August) and hence are called breeding schools. In the early spring
months, they are formed by the merger of male and female groups (as
soon as the gestating females have left the latter). Therefore, at this time
the breeding schools contain only barren and recently matured females.
In the summer months, lactating females join them in large numbers

465

621

and in July and August, form the bulk of the females in the school. The
growing juveniles form the majority of immature animals present in the
breeding schools. :

The process of reformation of summer into autumn schools and the
order of formation of winter types of common dolphin schools of the
Black Sea population has not been traced (Tarasevich, 1951).

The above extremely schematic description reflects the process of
formation of schools of different types and their transformation from
one type into another. In nature, this process is far more complex.
Such a diversity of schools and the numerous transformations which
they undergo in the course of a year with respect to age and sex com-
position, compel us to estimate very carefully the percentage ratio of
different age and sex groups in a school of dolphins using the data of
hunters. Not only the time, but also the region of catch should be taken
into consideration.

The common dolphin has been observed amidst groups of other
species of dolphins, such as the bottlenose dolphin, the shorthead dol-
phin, and even the pilot whale. The common dolphin does not take
very kindly to captivity compared to the other species and usually does
not survive for more than two or three months in ocean aquariums. Its
reproduction in captivity is not known (Tomilin, 1962).

Seasonal migrations and transgressions. The nature of seasonal
migrations among Atlantic and Pacific subspecies has not been
established. This aspect has been studied somewhat better for the Black
Sea subspecies and it has been assumed that the common dolphin in
general does not perform seasonal migrations, instead its movements
should be regarded as migrations in search of food (Tomilin, 1957). These
assumptions give rise to serious doubts, however.

The common dolphin is a typical pelagic animal avoiding sections
with fresh and turbid waters and evidently, therefore, does not enter the
Azov Sea (Freiman, 1950). It does not dive very deep in search of food
but is satisfied with what is available in the upper horizons of the sea.
It inhabits almost the whole of the Black Sea, in the coastal zone of
Crimea, the Caucasus, Turkey, Bulgaria, and Rumania and also far away
from the coasts. But its distribution in the equatorial seas is very uneven.

In winter months, in the period of spawning, sprat is scattered over a
large water body and is almost not utilized as food by this dolphin. At this
time the Black Sea anchovy concentrations are formed at the wintering
sites in the coastal waters of Georgia (Poti-Batumi) and on the southern
coast of Crimea (Balaklava). The presence of anchovy determines the
wintering regions of the common dolphin, most of which concentrate
on the coasts of Georgia and a small percentage south of the Crimean

466

622

peninsula. In spring, as the water begins to warm up, anchovy begin to
spawn and are scattered over a large expanse without forming concentra-
tions. Simultaneously, adult sprat and their young begin to concentrate
in the coastal waters of Crimea and northern Caucasus and in the open
northeastern waters of the Black Sea. Dolphins in search of food at this
time move northwest where they encounter sufficient collections of sprat
on which they feed in summer.

Depending on the collections of food objects, the common
dolphin forms concentrations of different types: (1) encountered very
rarely—only a few animals or small schools separated from each other;
(2) encountered rarely—some animals live everywhere while schools
are small and sparse; (3) encountered very frequently—numerous small-
and medium-size schools (up to 100 animals each) predominate, large
schools are very rarely seen; (4) encountered very often—large number
of predominantly large schools, of the order of hundreds or thousands
of animals in each, forming dense congregations in an extremely small
water body (Tsalkin, 1938). These dolphins usually form congregations
in the Tuapse-Sochi region along the coasts of the Caucasus; they may
be closer or farther away from the coasts (20 to 60 miles) in different
years. These are not long-time schools, surviving for just one to three
months. Simultaneous with them, schools of different strengths and
some individual dolphins are very widely distributed along the Black
Sea in the coastal zone as well as in the open sea, depending on the
availability of food. In some years, the distribution of the groups and
much smaller formations changes considerably according to seasonal
conditions (Tarasevich, 1958a).

As the dispersal of sprat concentrations increases, schools of dol-
phins begin to abandon the regions of summer habitation and gradu-
ally gather at the wintering sites where they live throughout the win-
ter months. The migrations of the Black Sea common dolphin are very
small but they bear a distinctly manifest seasonal character and are quite
constant in time and direction. There is every reason to consider them
seasonal migrations, therefore, at least in the eastern part of the Black
Sea. The periods and nature of migrations of the common dolphin in
the western part of the sea have yet to be established.

In the Bay of Bosporus, schools of dolphins swimming in different
directions are often noticed. It is possible that the common dolphin
sometimes leaves the Black Sea for the Sea of Marmara or, contrarily,
transgresses from the Sea of Marmara into the Black Sea, but there is no
recorded proof of this (Freiman, 1950). It has been assumed that a more
intense exchange prevails between the Black Sea and the Mediterranean

623

Sea dolphins than is usually acknowledged, which might be reflected to
some extent in the population dynamics of the dolphins of the Black Sea.

Reproduction. Mating and calving in the common dolphin are highly
prolonged but there is no one single view on their duration. Vari-
ous sources indicate that mating extends from June through October
(Sleptsov, 1941), from July to December (Tomilin, 1957), and even from
August to January (Maiorova and Danilevskii, 1934). But assumably most
of these dolphins mate over a much shorter duration—in July, August,
and the first half of September (Sleptsov, 1941); from August to October
with peak activity in September (Tomilin, 1957); or in July (Kleinenberg,
1956a).

The period of calving, according to various authors, extends from
May through November, peaking in June and July (Sleptsov, 1941); from
May to October or from June to November, the majority of females calve
from June to August, with maximum reached in July (Tomilin, 1957); or
from May through September with the peak in May and June (Kleinen-
berg, 1956a). In any case, in most females parturition takes place in the
summer months. Mating of the common dolphin has been observed time
and again from airplanes and hunting craft. A group of mating dolphins
consists of one female and six to eight males which pursue her at high
speed, chase each other, and hold the competitor behind the fins with
the teeth. When the fastest male catches up with the female, she turns
her abdomen upward, mates with him, and the two then disappear into
the water (Golenchenko, 1949b).

The duration of gestation has also been variously determined: about
10 months (Tomilin, 1957), 10-11 months (Kleinenberg, 1956a), and 11
months (Sleptsov, 1941).

The question of the periodicity of calving, 1.е., the rate of replen-
ishment of the population, has not yet been conclusively established. и
was first thought that dolphins produced calves once every four years or
twice in three years (Mal’m, 1936). According to M.M. Sleptsov (1941),
dolphins tend to calve annually but since fertilization can occur only
1.5 to 2 months after birth, mating advances in time every year and the
female becomes ready for fertilization afresh only after the males cease
to mate. From this, it may be assumed that the female dolphins calve
three years in a row but remain barren in the fourth. If this is so, then
one out of four of the eligible females should remain barren every year
(Kleinenberg, 1956a). According to V.E. Sokolov (1954), a calved female
becomes capable of mating again within three to five weeks after par-
turition, 1.е., even in the period of lactation, which would indicate the
absence of advancement in the period of mating in subsequent years.
Thus this dolphin usually reproduces every year and the percentage of

467

467

624

barren females in a group is not high (У.Е. Sokolov, 1954). Оп this
basis, it has been assumed that the sex cycle of the dolphin fits in the
annual cycle, with a gestation period of 10 months (Tomilin, 1957). More
reliable data are required before the question of the reproduction cycle
of the Black Sea common dolphin can be conclusively resolved.

Growth and development. The uterine horns and ovaries are usually
asymmetrical in the common dolphin; in 82.3% of cases, the left horn
was wider and longer than the right one while the left ovary was larger; in
17.7% of cases, the right horn and the right ovary were larger. Probably,
in an overwhelming majority of cases, the embryo (Fig. 254) grows in
the left uterine horn (Sleptsov, 1941).

The newborn calf of the common dolphin reaches, on average, half
the body length of the mother (sometimes more), which apparently is
far more than the corresponding dimensions of newborns among other
whales. The average body length of the newborn is 82-90 cm (Kleinen-
berg, 1956a). As a rule, one calf is born and twins are rare. The calf is
born not head first, as thought earlier, but tail first. The dorsal fin of the
embryo is convoluted and compactly pressed to the body and the caudal
flukes are balled into a “fist” (Kleinenberg, 1956a).

Lactation extends for about four months (Tomilin, 1957) or five to
six months (Sleptsov, 1941; Kleinenberg, 1956a). At the end of lactation,
the stomach of calves contained fish remains along with milk. Milk pro-
duction in the mammary glands of females is very little; thus a 168-cm
long female could yield only 100 ml (Tomilin, 1957). Apparently the calf
feeds often but in small portions. The composition of the milk of the
common dolphin is shown in Table 37.

Fig. 254. Embryo of the common dolphin (figure by N.N. Kondakov).

468

625

Table 37. Chemical composition of the milk of the Black Sea common dolphin
(Ural’skaya, 1957)

Composition of milk, % Author and
i Se a ht te I yeaniog
Fat Protein Sugar Dry Ash Water Та analysis
matter
43.71 5.62 1.45 7.53 0.45 48.76 100 Mal’m, 1932
41.56 48 1.49 6.82 0.45 51.62 100 Ural’skaya,
1954

The growth of the common dolphin in different age groups has not
been studied well. By autumn, the calf reaches a length of 100-110 cm,
1.е., in the period of lactation, it adds 20-30 cm. With cessation of
feeding on mother’s milk (winter), growth slows down and by the next
summer (toward the end of the first year) the dolphins reach a length
of 120-130 cm. The average dimensions of two-year-olds is 140-150 cm
and three-year-olds 165 - 170 cm. The length of most adult dolphins varies
around 170 cm and their average weight 50-55 kg. Sometimes animals
up to 200 cm long and 100 kg weight are encountered (Freiman, 1950).
The common dolphin is characterized by sexual dimorphism: males are
slightly larger than females right from the embryonic stage.

There is no unanimity of opinion regarding the time required to
attain sexual maturity. According to some data, female common dolphins
attain maturity in the third year and males in the fourth year (Sleptsov,
1941). According to other authors, females become mature at the end
of the third year and males in the third or the fourth year (Kleinenberg,
1956a). According to still others, the common dolphin is capable of
mating for the first time in the second year of life (Tomilin, 1957). The
body length of these dolphins at the time of attaining maturity varies
considerably. Mature females 140-150 cm long are encountered while
some 161-170 cm long are still immature (Kleinenberg, 1956a). With a
high degree of probability, it may be said that females with a body length
of less than 155 cm should be regarded as immature and those longer
than 155 cm as mature. This limit for males is put at a body length of
165 cm.

The maximum longevity of the common dolphin has been roughly
put at 20-25 years (Sleptsov, 1941; Freiman, 1950).

Enemies, diseases, mortality, parasites, and competitors. The Black Sea
common dolphin has no enemies. Subspecies of the Atlantic and Pacific
oceans may become prey to the killer whale. There has been almost no
study of the diseases afflicting this‘dolphin. Some instances of stones in
the ureter (Kleinenberg, 1956a) and of skeletal diseases (Tomilin, 1957)

469

626

are known. From among skin parasites, crustaceans (Xenobalanus glo-
bicitis, Penella pustulosa, and Larnaeonema nodicornis) have been found.

Nineteen species of endoparasites are known: these include trema-
todes (six), cestodes (three), nematodes (six), and acanthocephalans
(four). The trematode Brauniina cordiformis Wolf, 1903 is a parasite
of the stomach and intestine of the common dolphin, the bottlenose
dolphin and some species of spotted dolphins. Three species of trema-
todes of the genus Campula have been found in the bile duct and hepatic
ducts of just the common dolphin: C. delphini Peirier, in individuals from
the European waters of the Atlantic; C. palliata Looss, in the Atlantic
subspecies (Europe) and Black Sea residents, and C. rochebruni Peirier,
in specimens from the Atlantic waters of Europe. The trematode Dis-
tomum philocholum Creplin, which localizes in the liver, was detected
in the common dolphin from European waters. The trematode Galac-
tosomum erinaceum Peirier was found in the intestine of the common
dolphin also only from European waters.

The cestode Tetrabothrium fosteri is a parasite of the intestine
of the common dolphin and beaked whales. It has been found in
the Mediterranean Sea and in the southern part of the Pacific
Ocean. Monorygma grimaldii Monier localizes in the abdominal cavity,
mesentery, and diaphragm of the common dolphin and four other species
of dolphins of the Mediterranean Sea and the Atlantic Ocean. The
cestode Phyllobothrium delphini Bosc is quite extensively distributed
among marine mammals. In addition to the common dolphin, it has
been detected in six species of toothed whales, the bowhead whale, and
Weddell’s seal. It localizes in the skin and hypodermic tissue and has
been reported from the Atlantic Ocean, Mediterranean Sea, northern
and southern parts of the Pacific Ocean, and in the Antarctic.

One of the most widely distributed species of nematode parasites,
Anisakis (Anisakis) simplex Rudolphi, infects the gullet, stomach, and
intestine. Apart from the common dolphin, it has also been detected
in ten species of toothed whales, two species of baleen whales, and in
Steller’s sea lion. It has been reported in the North Sea and in different
parts of the Pacific Ocean (Kamchatka, Japan, and New Zealand).
Among the common and three other species of dolphins, Anisakis
(Anisakis) typica Diesing is a parasite of the stomach while Anisakis
dussmeerii Beneden has been found in the stomach and large intestine
of only the common dolphin. The nematode Halocercus (Halocercus)
delphini Baylis and Daubney was detected in the bronchi of the common
dolphin only from the Atlantic Ocean while another representative of
this genus, Halocercus (Posthalocercus) kleinenbergi Delamure, was found
in the lungs of only the Black Sea common dolphin. Yet another species

627

of nematode, Skrjabinalius cryptocephalus Delamure,-was found in the
lungs of only the Black Sea common dolphin.

Of the four species of acanthocephalans, Bolbosoma vasculosum
Rudolphi is a parasite of the intestine of common dolphins and beaked
whales from the Mediterranean Sea and the Atlantic Ocean. Bolbosoma
bellicidum Leuckart has been reported in the common dolphin. Coryno-
soma cetaceum Johnston, a parasite of the intestine of common and bot-
tlenose dolphins, was detected in Australian waters. The specific affinity
of the fourth representative of acanthocephalans, Corynosoma sp., has
not been established (Delamure, 1955; Tomilin, 1962).

Natural mortality has not been studied. It appears possible that the
common dolphin could die from an invasion of the nematode Skrjabi-
nalius cryptocephalus, which has infected the lungs of some animals to
such a degree that they were totally damaged (Kleinenberg, 1956a).

There are almost no competitors of the Black Sea common dolphin.
The Atlantic and Pacific subspecies may be food competitors of other
species of dolphins.

Population dynamics. In the prewar years, when the total catch of the
common dolphin in the Black Sea sometimes exceeded 100,000 animals
per year, a distinct reduction in population was noticed. During the years
of the Great Patriotic War [WW II], hunting activity perforce declined
and the population recouped slightly. Subsequently hunting resumed and
the population decimation was such that hunting these dolphins was
banned. Information on the population of the common dolphin in other
parts of its range is not available.

Field characteristics. This is a rather small (body length of adults
150-170 cm) agile dolphin having a high pointed dorsal fin with a
crescent-shaped notch in the posterior margin [falcate]. The beak is long
and sharply demarcated from the convex corpus adiposum [“melon’].
The dorsum, forehead, and upper caudal crest are black with two long
gray fields stretching along the flanks below these black fields; the ven-
tral side is white. This animal does not blow fountains [1е. its blows
are thin] and exhibits much of its body in normal movements. It often
leaps completely clear of the water. In rapid movement, white surf forms
along the body flanks. Schools of different strengths are common. Some-
times the herds merge into congregations of many thousands of animals.
(V.A.)

Economic Importance

Only the Black Sea common dolphin has been of commercial importance
and in times past was caught in large numbers. In all the remaining

470

628

sections of its range, this species has never attracted special interest and
was caught in very small numbers.

Hunting the Black Sea dolphin is centuries old. For many years
Turkish hunters used an extremely primitive method in killing them.
Hunting activity intensified only after the October Revolution. The first
cartel for this purpose was organized in 1929 in the Crimea. At this
time, along with the then-prevailing method of hunting with firearms,
dolphins were also caught in sweep nets. Hunting with firearms was
carried Out from ships using a smooth-barreled gun and buckshot. Since
in summer the killed dolphins sank rapidly, expert divers were always

onboard who jumped into the water simultaneous with the shot and held

the killed dolphin afloat. Nevertheless, many killed dolphins sank and
others who were injured probably died subsequently. Therefore hunting
with firearms was ultimately banned and further development proceeded
by perfecting the use of sweep nets.

After several improvements in catching dolphins in sweep nets in the
Black Sea, the following method was adopted. The net, made of fabric,
was up to 500 m long (sometimes up to 1,500 m), wall height 60-80 m,
and mesh size (diagonally) about 15 cm. Special rings were provided on
the lower seine rope of the net through which a rope was run for tighten-
ing the bottom of the net (like a purse). A brigade of catchers comprised
30-40 men equipped with one seiner, two or three motorboats, and up to
10 feluccas. The net was held on the stern of the seiner, cast by the mov-
ing ship, and drawn up from the water mechanically. The feluccas going
out for the catch and returning to the coast, were towed by motorboats.
Each felucca had two oarsmen with a sufficient stock of stones.

The seiner and another boat approached a school of dolphins from
two sides with a comparatively long distance between them, leaving the
feluccas with the oarsmen at equal intervals on the trail. Having swept
the school from three sides in the form of a horseshoe, they blocked the
fourth side with a small sweep net. Striking stones against each other
(“telephoning’’) and shouting from the feluccas left along the spread out
net, the catchers prevented the dolphins from leaving it until the ends
of the net could be drawn together. To prevent the net from sagging
or twisting in the water current, the hunters on the feluccas positioned
along the entire course of the net held onto its upper edge and flexed
it in different directions. By drawing the rope running through the rings
on the lower side, a “purse” was formed from which the dolphins caught
were gathered in by the feluccas and later loaded on the seiner for trans-
porting to the coast. Hundreds of dolphins were often caught in one
casting of the net. If another school of dolphins happened to be sighted
in the proximity, the brigade began casting a new net without giving up

471

629

Table 38. Catch of dolphins in the Black Sea (Bodrov, Grigor’ev, and Tver’yanovich,

1958)
Year Number of Year Number of
animals caught animals caught
1931 36,490 1945 3,464
1932 53,858 1946 15,872
1933 67,469 1947 19,400
1934 67,065 1948 20,863
1935 70,448 1949 32,200
1936 62,933 1950 42,535
1937 103,814 1951 20,250
1938 147,653 1952 50,618
1939 81,206 1953 42,757
1940 71,097 1954 82,809
1941 40,250 1955 18,354
1942 1,157 1956 16,082
1943 2,896 1957 29,800

1944 441

the first catch. Thus the brigades sometimes returned to the coast with
catches in several nets. Dolphin hunting was successfully developed in
Bulgaria later.

The results of erstwhile dolphin hunts in the Black Sea (in the Soviet
Union) are shown in Table 38.

When the catch rapidly diminished in subsequent years, hunting of
the Black Sea dolphin was discontinued.

Statistics of the catch of the Black Sea dolphin provide no
specieswise classification and hence the proportion of the common
dolphin is not ascertainable. The ratio of the different species of
dolphins in the Black Sea hunts varied considerably in different years
but the following figures could be regarded as averages: number of
common dolphin, common porpoise (Phocoena), and bottlenose dolphin
(Tursiops) = 200:10:1. Thus the common dolphin invariably represented
the bulk of the animals caught (Tsalkin, 1937).

The average weight of a whole dolphin according to long-term data
is 51 kg; thus the quantum of products recovered from a dolphin is
relatively small.

Weight of body portions and organs of a small female
Black sea common dolphin, kg (Tomilin, 1957)

Total weight Sy
Subcutaneous fat 10.98
Hump flesh 3.85

630

Caudal musculature 2.50
Backbone 2.55
Ribs with musculature in-between 1.85
Adipose body [“‘melon’’] 0.52
Dorsal fin 0.25
Flippers (two) 0.47
Caudal flukes 0.44
Lower jaw 0.48
Tongue Ой
Brain 0.67
Intestine 0.97
Gullet 0.23
Heart 0.17
Liver 0.60
Lungs (two with larynx) 1.00
Kidneys (two) 0.19
Stomach 0.20
Skull, blood, etc. 3.91

Dimensions and weight of common dolphins
(18 complete and 12 partial weighings)
(Dragunov and Kasinova, 1954)

Body length, cm 119-190 x

(159)

Total weight, kg 22.0-90.5 @:=9950:0)
% of total body weight

Blubber 26.0 - 44.8 (< = 34.4)
Carcass without head and 30.0 - 46.8 (x = 37.4)
viscera
Head 8.1-14.8 (x = 10.6)
Tail and fins 25-52 (9
Vicera (entire) 9.2 - 16.2 (х = 12.5)
Flesh with carcass 18.6 - 24.8 (x = 23.4)
Bones of carcass (not cleaned) 10.3 - 17.0 (x = 14.0)
Liver 1.4-3.3 12
Heart 0.39 - 0.86 (= 055)
Lungs 1.9-4.1 =
Stomach (without contents) O7 1:3 C= 10
Intestine (with contents) Zale @="'3'8)
Kidneys 0.40 - 1.02 (= 065)
Spleen 0.05 - 0.18 (> = 0.10)
‘Testes 0.5-1.8 G@ = 14)

472

631

Brain 1120 (GE E15)
Jaw fat 0.15 - 0.34 (=2N0:25)
Blood and unaccounted losses 0105-53 C= 91.2)

The amount of subcutaneous fat varies significantly depending on
the well-being of the animal. In the period of its maximum fattiness (in
April or March), the killed dolphin does not sink. The fat in the animal
is minimal in August. The fat recovered from the blubber is used in
the paint industry, etc. The better quality fat is used as a substitute for
medicinal cod liver oil. The hollow lower jaw of the common dolphin
contains a small amount of very valuable fat of a different chemical
composition. The oil processed from this fat does not solidify in the
cold and is used for oiling fine mechanisms. The carcass of the dolphin
is used as raw material for producing meat-bone meal for feeding farm
animals. High-quality glue is also produced from this raw material. The
skin of this dolphin, after appropriate processing, can be used in the
shoe industry for making leather boots.

From the viewpoint of the most rational use of the raw material,
dolphin hunting is best carried out in the spring and winter months, in
the period of maximum fattiness of the animals. With the same number
of animals caught then, a greater quantity of products can be recovered.

Commencing from 1965, every type of hunting of the Black Sea
dolphins has been banned and all the three species inhabiting this water
body have been brought under conservation. Similar measures have been
adopted in Bulgaria and Rumania. (V.A.)

Genus of Bottlenose Dolphins
Genus Tursiops Gervais, 1855

1843. Tursio. Gray. List Mamm. Brit. Mus., p.p. XXIII, 105. Delphinus
truncatus Montagu, 1821, nom. praeocc.

1855. Tursiops. Gervais. Hist. Nat. Mamm., 2, p. 323. substituted for
Tursio Gray, 1843.

1873. Hemisyntrachelus. Brandt. Мет. Ac. Imp. Sc., Pétersbourg. Sub-
stituted for Tursio Gray, 1843. (V.H.)

Medium-size dolphins with a maximum body length of 3.9 m.

The body build is somewhat heavy, with a “beak” of medium length.
The high dorsal fin has a fairly deep notch along the posterior margin.
The flippers are relatively broad. The lower jaw is slightly longer than
the upper one.

The upper portion of the body is dark gray (sometimes light gray),
the flanks gray, and the abdomen white (sometimes gray).

473

632

The rostrum of the skull is medium in length, longer and narrower
in females than in males. The premaxillae and nasal bones are adjacent
on the right side of the skull but not on the left. Broad processes join
the pterygoid bones which have an oblique notch posteriorly. The teeth
are fairly large (up to 10 mm in diameter) and number 15—28 = 76-106.
They are often completely worn down. The sternum consists of three
fused sections. Vertebrae 63-65. Ribs 12-13 pairs. Phalangeal formula:
I,-2, U7-9, Шб-в› [\>-5, and V,_>. The 2nd, sometimes the 3rd digit,
is the longest.

These dolphins are bentho-ichthyophagous. Periods of mating and
parturition are prolonged. Gestation extends for 11 months while lacta-
tion apparently continues for 4-6 months.

These dolphins are distributed in the Atlantic Ocean from the North
and Norwegian seas to the Mediterranean and Black seas and South
Africa in the east and from southern Greenland to Patagonia in the
west; in the Pacific Ocean, from California in the east and Japan in the
west to Chile and Australia and New Zealand; and, in the Indian Ocean,
from Australia to Africa.

Fossils have been traced to the Upper Pliocene of Europe.

The genus comprises a single species, the bottlenose dolphin, Т.
truncatus Montagu, 1821. Sometimes, one other species, Т. gilli Dall,
1873, or even six more species are recognized. Apart from T. gilli, the
five others are Т. nuuanu Andrews, 1911; Т. aduncas Ehrenberg, 1833; T.
parvimanus Reinhardt, 1888; Т. gephyreus Lahille, 1908; and Т. abusalam
Ruppell, 1842. (V.S.)

BOTTLENOSE DOLPHIN

Tursiops truncatus Montagu, 1821

1780. Delphinus tursio. Fabricius. Fauna groenlandica, p. 49, Greenland.
The name has no nomenclatural importance since its applicability
to this species could not be established.

1804. Delphinus nesarnak. Lacépéde. Hist. Nat. Cétacées, р. XLII, 307.
Northern Atlantic (nom. ргаеосс).

1821. Delphinus truncatus. Montagu. Mém. Wernerian Nat. Hist. Soc. 3,
p. 75. Devonshire, England.

1832. Delphinus aduncus. Ehrenberg. In: Hemprich et Ehrenberg. Sym-
bolae physicae, Mammalia, 2. Red Sea.

1873. Tursiops gillii. Dall. Proc. California Ac. Sc., 5, p. 13. Monterey,
California.

1911. Tursiops nuuanu. Andrews. Bull. Amer. Mus. Nat. Hist., 30, p. 233.
Gulf of California.

633

1940. Tursiops truncatus ponticus. Barabash-Nikiforov. Cetacean fauna.
Black Sea, p. 56. Novorossiisk, Black Sea. (V.H.)

Diagnosis
Only species of the genus.
Description

Two color groups are recognized among bottlenose dolphins of the Black
Sea (Barabash-Nikiforov, 1940, 1960). Type A is characterized by a fairly
distinct boundary between the dark color of the dorsum and the white
color of the abdomen with a light-colored angular patch in the dark
field midbody; the apex of the patch is turned toward the dorsal fin
(Fig. 255). In type B, the boundary between the pigmented upper and
lower surfaces is not distinct and appears as vague straight, wavy, or
broken line, without the light-colored angle near the dorsal fin. The
quantitative ratio of animals of the two types varies in different years.
Among 50% of the bottlenose dolphins of the two types, a frontopectoral
line joins the eyes and gradually extends from their outer corners toward
the flippers. Among the bottlenose dolphins of the Atlantic, the light-
colored patch characteristic of type A and the frontopectoral line are
invariably absent. The latter can, however, be seen in the dolphins of the
Mediterranean Sea.

Cervical vertebrae 7, thoracic 12-14, lumbar 17, and caudal 26-27.
The cervical vertebrae may be fused in various combinations.

The main body dimensions (as percentage of body length) of bottle-
nose dolphins averaged for 52 adult animals with a body length of
270-310 cm (Barabash-Nikiforov, 1940) are: distance from tip of lower
jaw to base of flippers 18.5, from tip of upper jaw to blowhole 14.0, from

473

Fig. 255. Bottlenose dolphin, Tursiops truncatus (figure by N.N. Kondakov).

474

634

Ир of upper jaw to commencement of dorsal fin 42.7; length of flippers
14.6; length of dorsal fin 15.6; and width of the left caudal fluke 9.6.

The average length of males caught in the Black Sea was 228 cm,
of females 222 cm; the largest male caught was 310 cm long. Bottlenose
dolphins from other regions are larger than their Black Sea counter-
parts: in the waters of Japan, the males average a length of 2.7 m and
females 2.8 m; bottlenose dolphins in the Atlantic Ocean attain a length
of 3.1-3.3 m and in the Mediterranean Sea 3.2 m. With advancing age, a
relative reduction (as percentage of body length) is noticed in the length
of the oral slit, length of flippers, and distance from tip of snout to eye
and up to the flippers (Barabash-Nikiforov, 1940).

The average main skull measurements of 10 male bottlenose dolphins
from the Black Sea with a body length of 180-310 cm and five females
with a body length of 214-234 cm (Tomilin, 1957) are. respectively:
condylobasal length 44 and 44, zygomatic width 23 and 22, length of
rostrum 24 and 24, width of rostrum at base 12 and 12, and length of
lower jaw 36 (three measurements) and 40 (two measurements).

The skull of the Black Sea bottlenose dolphin is smaller compared to
bottlenose dolphins residing in other seas (Fig. 257). Vertebral sections
comprise (as percentage of length): cervical 3, thoracic 23, lumbar 30,
and caudal 44 (Slijper, 1936). (V.S.)

Geographic Distribution

Seas of the temperate and warm belts in the Northern and Southern
hemispheres.

Geographic Range in the USSR

Baltic Sea from Gulfs of Riga and Finland, Barents Sea east to Novaya
Zemlya, and the Black Sea; Pacific Ocean Basin, the Sea of Japan, south-
ern part of the Sea of Okhotsk, and waters of the southern half of the
Kuril range (Fig. 258).

Geographic Range outside the USSR

In the Atlantic Ocean, in the zone of influence of the Gulf Stream, the
range extends north up to the Lofoten Islands on the coasts of Nor-
way, coasts of Iceland, Jan Mayen, and south of Greenland (Fig. 259).
In the south, it extends along the African coasts right up to South
Africa and into the Mediterranean Sea. Along the west coast of the
Atlantic Ocean, this dolphin probably inhabits the waters from New-
foundland to Florida and the Gulf of Mexico and beyond, along the

635

Spe At
OY,
ИИ, и,

й А?
= = > Gh, Licey =F
Ё.-—-

„=

\\

Fig. 256. Embryo of a partly albino bottlenose dolphin, Tursiops truncatus (figure

474
by N.N. Kondakov).

2

UE

РИА

2

i

i. Ss
Миле”,
2 Z

Se

474 Fig. 257. Skull of the bottlenose dolphin, Tursiops truncatus (figure by М.М. Kondakov).

coasts of Argentina, Uruguay, and Brazil. The bottlenose dolphin resides
in the waters of the Indian Ocean (Arabian and Red seas, Bay of Ben-

477 gal, and Seychelles Islands), south up to the coasts of Tasmania and
Australia. In the Pacific Ocean, it is encountered on the coasts of the

636

Е
a
8
a
5
3
с
©
8
<

Fig. 258. Range of the bottlenose dolphin Tursiops truncatus т the USSR (М.А. Arsen’ev).

475

4! |
Wing i] Ш " |
nt

wih
|

1
cS, il
Say"

ui ia
ae aaa!
thy pia
IY

° О ПАН й
iy
une

1 ba
pie Quen
— ОНИ 0

Nh J

5 ae
я НН
ани

600 0 1000 2000 3000 3000 5000 km

Fig. 259. Species range of the bottlenose dolphin, Tursiops truncatus (V.A. Arsen’ev).

476

638

Korean Peninsula and Japan, on the coasts of North America (Oregon
and California states), and Mexico. In the south, it reaches Australia |
and New Zealand. No information is available for the Pacific Ocean
coast of South America (Sergeant and Fisher, 1957; Tomilin, 1962).
(V.A.)

Geographic Variation

Geographic variation has not been adequately studied. Apparently three
subspecies are encountered in the waters of the Soviet Union.’ These
are:

1. Atlantic bottlenose dolphin, T. t. truncatus Montagu, 1821.

This is the largest form, with body length reaching 390 cm and skull
length 550-579 mm. The depression on the lower surface of the frontal
bone is inconspicuous. The number of teeth is more than in the other
subspecies, usually 21-26 pairs in the upper jaw and 20-25 pairs in the
lower.

This subspecies is found in the Baltic and Barents seas and in the
waters of the North Atlantic Ocean.

2. Black Sea bottlenose dolphin, 7: Е. ponticus Barabash, 1940.

In body and skull dimensions, it is the smallest. Its body length does
not exceed 310 cm while the skull length may reach 503 mm. The teeth
vary in number from 19-22 pairs in the upper jaw and 18-21 pairs in the
lower. The rostrum is shortened and broadened at the base. Broadening
and dilation in the midportion of the premaxillae are faintly manifest or
altogether absent.

This subspecies is found in the Black Sea.

Outside the USSR, it has been reported in waters south and west of
the Black Sea.

3. Far-eastern bottlenose dolphin, T. г. gillii Dall, 1873.

In dimensions, it probably occupies an intermediate position with
the skull length reaching 540 mm. The articulation for the mandibular
condyle is very large. The lower surface of the frontal bone is deeply
concave anterior to the optic canal. The rostrum is relatively long and
narrow at the base.

This subspecies is found in the seas of Japan and Okhotsk and waters
of the southern part of the Kuril range.

Outside the USSR, it has been reported in the Pacific Ocean, at
least in the northern half. (V.A.)

’Diagnosis of the subspecies after А.С. Tomilin (1957) requires verification and
supplementation.

478

639

Biology

Population. This dolphin is quite rare in the North Atlantic and Pacific
oceans. It is comparatively less in number in the Black Sea, especially
compared to the large schools of common dolphins.

Food. Information on food in the Atlantic and Pacific oceans is frag-
mentary. Fishes serve as the main food, including eels, gray mullets, and
small sharks. Bottlenose dolphins examined from the Mediterranean Sea
contained the remnants of cuttlefish. More complete information is avail-
able about the food objects of the Black Sea bottlenose dolphin. The diet
includes many species of fish, crustaceans, and mollusks: haddock, floun-
der, skate, umbrinas, scorpionfish, anchovy, mullets, gray mullet, Black
Sea shad, and bonito; shrimps and isopods; Nassa reticulata, Cardium
sp., Modiola phaseolina, Syndesmia sp., Cardium simile, and Mytilus sp.
Mollusks are probably not consumed by the bottlenose dolphin but enter
the stomach together with sand and gravel, invariably present in large
quantity. It is also quite possible that the crustaceans found in the stom-
ach of these dolphins had been priorly ingested by the fish the animals
consumed.

Haddock, followed by flounder, is the most abundant fish found in
the stomach of bottlenose dolphins. Anchovies, during the period of
their en masse concentration, also play a significant role in the dolphin’s
diet, sometimes even occupying a favored position. All other species of
fish play only a secondary role. Thus the bottlenose dolphin is, for the
most part, benthophagous since pelagic fish, which even form massive
concentrations (anchovy), are consumed more rarely (Kleinenberg, 1936,
1938, 1956a).

Data on the quantum of food intake of the bottlenose dolphin (under
natural conditions) are not available. A bottlenose dolphin held captive
in a Florida aquarium consumed 20 kg of fresh fish every day while adults
in a New York aquarium could manage with 32 kg of fish each (Tomilin,
1957).

Daily activity and behavior. It was earlier assumed that bottlenose
dolphins feed only at night but the stomach of those caught during the
day contained partly digested food. This led to the conclusion that there
are no obvious changes in their diurnal activity (Kleinenberg, 1956a).

Bottlenose dolphins do not form large schools (Fig. 260). They
are usually found in groups of ten or a few tens, and only rarely in
hundreds. Sometimes the animals produce blows rising to a height of
1-1.5 m. The respiratory pause in an animal moving undisturbed aver-
ages 15-17 sec. Feeding dolphins rise to the water surface at irregular
intervals, ranging from 5 sec to 2 min, while inhalation/exhalation extend

640

478 Fig. 260. Bottlenose dolphin at sea (photograph by О.А. Morozov).

for 1-2 sec. Sometimes the bottlenose dolphin remains under water for
4 or even 6-7 min. It can dive to a depth of 50-90 m (possibly even
to 150 m) at speeds varying from 3-5 to 11-13 km/hr, with a maximum
of 30-40 km/hr. Sometimes the bottlenose dolphin breaches the water,
“rising” to heights of over 3-4 m (Tomilin, 1957, 1962).

Cases of active cooperation have been observed among these dol-
phins. On October 30, 1954, an adult dolphin in a herd of some 25
animals was stunned by a dynamite explosion off the coast of Florida.
Immediately, two large dolphins came to its rescue, lifted it from under
the flippers, and supported it on the water surface. The rescuers peri-
odically dived and surfaced and throughout this period the entire group
moved in a wide arc. The operation gave the impression that the pair of
supporting animals interchanged since the rest of the animals remained
in their proximity. After a few minutes, the affected dolphin began to
recover and the entire school then quickly moved away from the ship.
Every 10-20 sec, the dolphins breached the water, flying in the air for
7-9 м.

479 In another incident, recorded оп November 23, 1954, also in Floridan
waters, some dolphins that had been caught were being transferred from
a ship into a specially constructed tank. Three adult animals had already
acclimatized well while the fourth on being lifted from the deck by the
tail struggled fiercely and hit its head against the wall of the tank. The

641

dolphin was stunned and sank to a depth of 2 т. Two of the already
transferred dolphins quickly rushed, as in the previous case, to hold
and support it on the surface of the water until the affected animal’s
respiration normalized. Thereafter the two rescuers returned frequently
to the bruised animal, abandoning it only after it had begun to swim freely
without their assistance. The third dolphin had meanwhile remained in
the proximity of the others (Siebenaler and Caldwell, 1956).

The bottlenose dolphin can survive in captivity better than other
species, live long in large aquariums, is easily trained, and often raises
offspring in captivity. Sometimes a conditioned reflex can be developed
in a dolphin within a week’s time by repeating the stimulus no more than
two or three times, after which a new stimulus can be introduced. The
bottlenose dolphin learns quickly to jump through a hoop, tug a boat
or a raft, play with a ball, etc. The dolphins are usually held in tanks
in groups, eat well, and sleep well. They often sleep at night and in the
morning, occasionally enjoying a siesta after a meal.

They produce different types of sounds in the frequency range of
7,000 to 170,000 Hz: whistling, howling, barking, clapping of jaws, etc.
Each sound has a particular meaning, relevant to feeding, excitation,
mating, fright, etc. (Tomilin, 1962).

Seasonal migrations and transgressions. On the European and Amer-
ican coasts of the North Atlantic Ocean, bottlenose dolphins proba-
bly perform regular migrations but no direct observations have been
reported. In the Black Sea, they inhabit only the coastal shallow water
zone and are never seen in the open sea. They are most often seen in the
northeastern part of the Black Sea along the coasts of southern Crimea
and northern Caucasus but have been sighted in small numbers in other
parts of the sea as well. During the migration of Azov anchovy, bot-
tlenose dolphins form schools that are larger than at any other time and
concentrate in the fore-channel zone of the Black Sea and are seen even
in Kerch Strait. However, they do not enter the Azov Sea. Apparently,
the Black Sea bottlenose dolphin does not undertake regular migrations
(Kleinenberg, 1956a).

Reproduction, growth, and development. The period of reproduction
is highly protracted although most calves are born in the warm season
of the year. However, in a Florida aquarium, four calves were born in
February, one in March, one in April, and four in May (Tomilin, 1962).

Thirty-four females caught in November were accompanied by suck-
iing calves and the mammary glands of the mothers held milk. Gestation
extends for 12 months. In 1953, in a marine aquarium, a female in the
company of a male from February 23 through March 9 underwent partu-
rition on March 4, 1954. Some other observations confirm the 12-month

642

480 Fig. 261. Parturition in the bottlenose dolphin, Tursiops truncatus (figure by
N.N. Kondakov).

cycle of gestation (Tavolga and Essapian, 1957). Large embryos (Fig. 261)
measured 90 to 130 cm in length while two newborn calves were 122 and
124 cm long (Barabash-Nikiforov, 1940). According to observations on
captive dolphins, calves were born under water tail first, with formed
dorsal fin and caudal flukes. Immediately after whelping, the mother
swirled about in the tank, thereby readily separating the umbilical cord.
The newborn swam slowly along the waterline, inclined to the surface,
and took its first breath 10 min after birth. After 1 hr and 45 min, it suck-
led milk for the first time. According to other observations, subsequent
feedings proceed day and night at intervals of 15-30 min.

The females in an advanced stage of pregnancy segregate from the
herd and form a separate group of pregnant animals. After parturition,
the mother as well as the other females living in the proximity protect
the calf by turns from. possible attack by males. For a few weeks, the
mother does not allow her calf to swim for more than two-three meters

480 out of her sight. The duration of lactation apparently varies markedly.
In one case, the calf was milk-fed for 3.5 months while another calf was
found to partake of a fish by itself just seven months after birth. Yet a
case is known of a young dolphin continuing to suckle its mother for 18
months (Tavolga and Essapian, 1961*; Tomilin, 1962).

. 481

643

Growth rate has not been established. Apparently the newborn grows
rapidly on the high fat content of the mother’s milk, which in composi-
tion contains: fat 46.1%, protein 11.55%, sugar 1.57%, dry residue 13.5%,
ash 0.38%, and water 40.5% (Ural’skaya, 1957).

A female born in a marine aquarium was fertilized for the first time
at six years of age and the first calf born a year later. Considering the
duration of gestation and lactation, it may be assumed that the bot-
tlenose dolphin has a two-year reproductive cycle. A female born in a
Florida aquarium gave birth to four calves over a 15-year period. A male
when caught was approximately 1.5 - 2 years of age with a body length of
183 cm; he lived in the marine aquarium for six years and ten months
and grew in this period to 231 cm, 1.е., added 48 cm in almost seven
years (Sergeant, 1959b).

Enemies, diseases, parasites, mortality, and competitors. The killer
whale can be an enemy of the bottlenose dolphin, as also of
other dolphins. The diseases affecting the bottlenose have not been
investigated. In a Florida aquarium, two dolphins died of infectious
erysipeloid (pathogen: Erysipelothrix rhusiopathiae). In a New York
aquarium, instances of pneumonia were recorded. Skeletal diseases were
also reported (Tomilin, 1957).

A female bottlenose dolphin measuring 266 cm long and weighing
180 kg was caught in the waters of Japan in July, 1954 and placed in
a tank. In December of the same year, many nodular formations were
noticed on her skin, the animal became debilitated, and a month later
died. The affliction turned out to be a bacterial fungal infection caused
by a parasite of the genus Trichophytum Malm. Acute pneumonia was
also reported in this dolphin (Hoshina and Sugiura, 1956).

Eleven species of helminths have been reported in the bottlenose
dolphin: three trematodes, three cestodes, four nematodes, and one
species of acanthocephalan. The trematode Braunina cordiformis Wolf,
a parasite of the stomach and intestine of the bottlenose dolphin,
was also found in the common dolphin and one species of spotted
dolphins in the Adriatic Sea, in the region of Rio de Janeiro, and in
the waters of California and Panama. Another species of trematode,
Synthesium tursionis Marchi, was detected in the intestine of only the
bottlenose dolphine in European waters. In a Californian aquarium, the
bile ducts of the liver of a bottlenose dolphin revealed Zalophotrema
hepaticum Stunkard and Alvey, known earlier from the Californian sea
lion.

The cestode Phyllobothrium delphini Bosc, parasitizing the skin and
subcutaneous tissue, is widely distributed among marine mammals.
Besides the bottlenose dolphin, it is found in the bowhead whale,

644

six species of toothed whales, and one species of seals. It has been
detected in the Atlantic and Pacific oceans, Mediterranean Sea, and in
the waters of Australia and Antarctica. In the bottlenose dolphin and
four other species of dolphins, the cestode Monorygma grimaldii Monier
parasitizes the abdominal cavity, mesentery, and diaphragm; it has been
detected in the Atlantic Ocean and the Mediterranean Sea. The third
species of cestode belongs to the genus Diphyllobothrium (species not
established).

Of the four species of nematodes, Anisakis tursionis Gruz was
detected in the bottlenose dolphin from the Indian Ocean. Crassicauda
crassicauda Creplin, parasite of the urogenital system, was found in
beaked whales and six species of baleen whales, in addition to the
bottlenose dolphin; it was detected in the Atlantic Ocean in the Northern
and Southern hemispheres. The nematode Halocercus lagenorhynchi
Baylis and Daubney is a parasite of the lungs (bronchi) of the bottlenose
dolphin and the white-beaked dolphin; it was detected on the European
coast of the Atlantic Ocean and in the waters of Australia. Stenurus
ovatus Linstow was detected in the blowhole, bronchi, and blood vessels
of only the bottlenose dolphin in the Mediterranean and Black seas.
Forty-one larvae of the nematode genus Anisakis were found in a
female bottlenose dolphin caught in the waters of Japan, probably
the larvae of Anisakis catodonis Baylis (Hoshina and Sugiura, 1956).
The only species of acanthocephalans localizes in the intestine of the
bottlenose dolphin Corynosoma cetaceum Johnston, and has also been
found in the common dolphin in the waters of Australia (Delamure,
1955). 1

The natural mortality of the bottlenose has not been studied. Some
other species of dolphin feed on the same diet and constitute food
competitors.

Field characteristics. Adults usually measure 220 to 250 cm. They
are dorsally and ventrally light in color; the transition in coloration is
gentle, with no sharp demarcation between the two. Body movements
are placid. It neither splashes water while swimming nor does it breach
high. (V.A.)

Economic Importance

In the recent past, several hundreds of bottlenose dolphins were caught
every year in the Black Sea. In other regions, where many other species
of dolphins were caught, the bottlenose constituted a negligible per-
centage of the total catch. Various types of nets were used to catch it
482 along with other dolphins. The weight of the animals ranged from 90 to

645

almost 200 kg, of which the skin with the fat accounted for 30% and the
flesh 33%. An analysis of the seasonal catch of the Black Sea bottlenose
dolphin showed that the average weight of the dolphin was 175 kg, of
which the subcutaneous fat was 50.8 kg (29%) and the skin 7 kg (4%)
(Dragunov and Kasinova, 1951).

The total quantum of products obtained from bottlenose dolphins
caught all over the range was extremely meagre. Hunting of the bot-
tlenose dolphin, and of other species of dolphins as well, has been banned
in the Black Sea. (V.A.)

Genus of Right Whale Dolphins
Genus Lissodelphis Gloger, 1841

1830. Tursio. Wagler. Nat. Syst. Amphibien, р. 34. Delphinus ретопи
Cuvier. Nom. ргаеосс.

1841. Lissodelphis. Gloger. Gemein. Naturgesch, 1, р. 169. Delphinus per-
onii Lacépéde, 1804.

1846. Delphinapterus. Gray. Zoology. Voyage Erebus and Terror, 1, p. 36.
Delphinus peronii Lacépede. Nom. ргаеосс. (V.H.)

Small dolphins, with a body length of up to 2.4 m.

Table 39. Weight of body parts and organs of bottlenose dolphin from the Black Sea (in
March), kg (percentage of total weight shown in parentheses) (Okuneva, 1934)

Total weight 143.53 (100.0)
Head 12.0 (8.36)
Flippers and dorsal fins 2.06 (1.43)
Caudal flukes 2.09 (1.45)
Skin 6.0 (4.11)
Carcass fat 46.0 (32.11)
Flesh (muscles) 39.9 (27.17)
Bones (trunk skeleton) 17.0 (11.83)
Brain 1.46 (1.01)
Tongue 0.37 (0.25)
Lower jaw 0.79 (0.55)
Upper jaw 1.09 (0.76)
Viscera, including: 173 (12.05)
Liver 2.1 (1.88)
Heart 0.67 (0.46)
Lungs 4.5 (3.13)
Stomach 1.55 (1.08)
Intestine 7.2 (5.01)

Kidneys 0.68 (0.47)

646

The body is highly elongated and well proportioned. The head, with
a low and inclined forehead gradually passes into the relatively short
“beak”. The “beak” is set off from the adipose body of the forehead by
lateral furrows. The flippers are crescent-shaped. A dorsal fin is lacking.
Caudal flukes are relatively small. The lower jaw is slightly longer than
the upper.

The body is black above and whitish below and the boundary between
the upper and lower body colors is very sharply defined.

The rostrum is broad and slightly longer than the cranium. The ptery-
goid bones are not adjacent. The premaxillae are flattened anteriorly.
The mandibular symphysis is short, less than one-fifth the jaw length.
The teeth are small and number 73-4. The scapula, almost semicir-
cular in form, has prominent coracoid and acromion processes. Cervi-
cal vertebrae 7, thoracic 14-17, lumbar 29-33, and caudal 35 - 40; total
88 - 92.

The biology of this genus is hardly known but apparently these dol-
phin species feed mainly on fish and cephalopods.

Information on the distribution of this genus is very inadequate and
sketchy: Atlantic Ocean around South Africa and south of southern
Brazil in South America, Pacific Ocean along the coasts of Chile, north
up to 37° М lat., and from Tasmania and New Zealand (evidently Aus-
tralia too) north up to New Guinea, and in the North Pacific Ocean
from the Bering Sea south to California and Japan.

The genus comprises two species: the southern right whale dolphin
(Peron’s dolphin), Lissodelphis peroni Lacépéde, 1804, and the northern
right whale dolphin L. borealis Peale, 1848. Only the latter has been
established in USSR waters but the appearance of the southern species
is quite likely. (V. S.)

NORTHERN RIGHT WHALE DOLPHIN
Lissodelphis borealis Peale, 1848

1848. Delphinapterus borealis. Peale. U. S. Explor. Exped. 8, p. 35, 10°
west of Astoria, Oregon, USA; 46°6’50” М lat. and 134°5' Е long.
(V. H.)

Diagnosis

The body length, up to 2.4 m, is maximum for the genus. For a descrip-
tion of the external features, see under the characteristics of the genus.
Typically, the body is almost entirely black. A narrow light-colored band
runs only on the ventral side from the throat to the tail and forms a

647

483 rhomboid patch between the flippers. The upper side of the beak, part
of the forehead, and tip of the lower jaw may also be white (Fig. 262). For
skull characteristics, see under the description of the genus. Phalangeal
formula: I, -5, Пз, Шо, [Уз, and V,-3. (V.S.)

Description

The upper side of the caudal flukes is black but partly white or gray on
the underside and dark along the margins. The flippers are black.

Teeth 2-77.

Cervical vertebrae 7, thoracic 14-17, lumbar 29-33, and caudal
35-40; total 88-92. The first two cervical vertebrae are fused.

The body dimensions of four males and seven females caught on the
coasts of Japan (Tobayama, Uchida, and Nishiwaki, 1969) are (in cm):
body length 145 - 292 (x 204); distance from tip of snout to forehead 3-6
(x 4.6), to center of blowhole 22 - 34 (x 30), and to flippers 39 -64 (x 53);
distance from anal opening to notch between caudal flukes 40-70 (х 53);
length of flippers 24-31 (х 28), maximum width of flippers 7.3-9.7 (х
8.6); distance between caudal flukes from apex to apex 18-41 (х 33); and
maximum height of body 20-37 (x 30).

The skull (Fig. 263) dimensions of three male northern right whale
dolphins (Tomilin, 1957) of 213, 208, and 246 cm, body length are respec-
tively: condylobasal length 45, 42, and 44 cm; length of rostrum 26, 23,
and 24 cm; width of rostrum at base 11, 11, and 11 cm; and length of
lower jaw 38, 37, and 38 cm. (V.S.)

Geographic Distribution

North Pacific Ocean.

483 Fig. 262. Northern right whale dolphin, Lissodelphis borealis (figure by М.М. Kondakov).

648

arn

ras

SSS Sanaa

{ SESS

483 Fig. 263. Skull of the northern nght whale dolphin, Lissodelphis borealis (figure

by N.N. Kondakov).
Geographic Range in the USSR

Waters of the Kuril Islands (from Shpanberg to Paramushir) but
484 encountered more often from the Pacific Ocean side. Apparently also
occurs in the eastern parts of the seas of Okhotsk and Japan (Fig. 264).

Geographic Range outside the USSR (Fig. 265)

Widely distributed in the Pacific Ocean along the American and Asian
coasts, in the waters of Honshu and Hokkaido islands, on the coasts of
North America from California and Washington states to the Gulf of
Alaska and the eastern part of the Bering Sea; residence in the western
part of the sea has not been established. (V.A.)

Geographic Variation

Not established. Quite likely the accepted species of this genus are only
subspecies of a single species (L. peroni Lac.). (V.A.)

Biology

The population of this species is very small over much of its range.
Махипит concentration in all probability occurs in the waters of Japan.

484

486

649

Fig. 264. Range of the northern right whale dolphin, Lissodelphis borealis in the
USSR (V.A. Arsen’ev).

250 0 250 500 750 1000km
= —————— |
150 160 170

Fish represent the main food while cephalopods are consumed in small
quantities. A female caught on March 28, 1959 east of Honshu Island
contained an embryo 43 cm long.

These dolphins live predominantly in small schools far away from
the coasts. They leap clear of the water. They exhibit a strong mutual
affinity, never abandoning an injured animal and remaining with it for a
long time.

In the waters of Japan, the northern right whale dolphin is caught
regularly along with other species. At least a few hundred of them are
hunted every year. Thus, in May and June of 1949 alone, in the central
part of the coastal Pacific Ocean waters of Honshu Island, a whaling com-
pany caught 465 of them. Hunting is usually donw with rifles, followed by
harpooning the injured animal manually. The shooter and the harpoon
striker operate on a special wooden platform fitted on the bow of the

650

(лэмэзту “W'A) 51122404 siydjapossr] ‘uydjop этеча з48ы way OU эц1 Jo э8иет $э0э4$ ‘695 “31
9

486

651

Fig. 266. Northern right whale dolphin, Lissodelphis borealis, on the deck of a
ship. Pacific Ocean east of Honshu Island, 1959 (photograph by G.M. Kosygin).

ship. Ships with a water displacement of 20 to 30 tons and capable of log-
ging 7 to 10 miles per hour are used. The killed dolphins are processed on
the deck of the ship (Fig. 266). A ship enjoying a successful hunting expe-
dition returns with a catch of 200 or more dolphins of different species.

An experimental catch of these dolphins using woven nets was саг-
ried out near Iturup in the Kuril Islands in September, 1955. The nets
were cast over a herd of 25 animals but only 5 were actually caught.

The products of dolphin hunting are marketed in Japan where the
liver, heart, and kidneys are consumed. Bones and various remnants are
ground into fertilizer meal. Oil is obtained from the blubber. The skin
is used in making leather goods (Wilke, Taniwaki, and Kuroda, 1953;
Sleptsov, 1955, 1961; Klumov, 1959). (V.A.)

SOUTHERN RIGHT WHALE DOLPHIN (PERON’S DOLPHIN)
Lissodelphis peroni Lacépéde, 1804

1804. Delphinus peronii Lacépéde. Hist. Nat. Cétacées, p.p. XLIII, 316.
Tasmanian waters. (V.H.)

These dolphins are smaller than the northern right whale dolphin. The
body length of a male caught in waters of Japan was 227 cm (Tobayama et
al., 1969). In external features and structure and skull proportions, they
in no way differ from their northern counterparts. The typical coloration
of the southern species is: black dorsally and on most of the lateral
surface, and white on the abdomen and the lower parts of the body
flanks (Fig. 267).

487

487

652

The tip of the snout and small sections of the vertical surface of the
caudal flukes adjoining the caudal crest are black. The white field covers
the level of articulation of the black flippers. The edge of the upper jaw
(apart from its apex) and the zone posterior to and below the eyes are
white.

Teeth 4%.

Cervical vertebrae 7, thoracic 17, lumbar 29, and caudal 37; total 90.

The body dimensions of a male caught on the coasts of Japan
(Tobayama et al., 1969) are (in cm): body length 227; distance from
tip of snout to forehead 4, to center of blowhole 36, and to flippers 59;
distance from anal opening to slit between caudal flukes 61; length of
flipper 30, maximum width of flipper 9.5; distance between caudal flukes
from tip to tip 38; maximum height of body 27.

The skull (Fig. 268) dimensions of the same specimen are (in cm):
condylobasal length 47, length of rostrum 26, width of rostrum at base 12,
interorbital width 18, length of lower jaw 40, and length of mandibular
symphysis—[48].

Geographic Distribution

Habitation in the waters of the USSR has not been established but res-
idence in or transgressions into the Sea of Japan or waters of the Kuril
Islands are possible.

Outside the USSR, it is encountered in the southern part of the
range (see under the characteristics of the genus); indicated for the waters
of Japan. (V.H.)

Fig. 267. Southern right whale dolphin, Lissodelphis peroni (figure by N.N. Kondakov).

653
Biology

Not known. Japanese fishermen from Otsu, Ibaraki Prefecture caught a
group of dolphins swimming together on April 1, 1969, which included
the southern right whale dolphin, the northern right whale dolphin, and

the Pacific common dolphin (Tobayama, Uchida, and Nishiwaki, 1969).
(V.A.)

Genus of Shorthead Dolphins
Genus Lagenorhynchus Gray, 1846

1846. Lagenorhynchus. Gray. Ann. Mag. Nat. Hist., 17, p. 84. Delphinus
albirostris Gray, 1846.

1866. Electra. Gray. Cat. Seals and Whales Brit. Mus., p. 268.
Lagenorhynchus electra Gray, 1846. Nom. ргаеосс.

1866. Leucopleurus. Gray. Proc. Zool. Soc. London, p. 216. Delphinus
leucopleurus Rash, 1843.

Sr a

ey, SLE!
Са

^^
В tA
77°

Fig. 268. Skuil of the southern right whale dolphin, Lissodelphis peroni (figure ,
by N.N. Kondakov).

489

654

1966. Peponocephala. Nishiwaki and Norris. Ocean Res. Inst., Univ.
Tokyo, 5, 139. Electra electra Gray, 1846. (V.H.)

Small dolphins, with a well-proportioned body ranging in length from
1.5 to 3 м.

The “ЪеаК” is short, not more than 7 cm; in most of these animals,
it is well demarcated from the forehead by a furrow. The high dorsal fin
is crescent-shaped and located at the center of the dorsum or slightly
anterior to it. A crescent shape is also characteristic of the flippers.
Dermal keels run along the dorsal and ventral sides of the caudal stem.
The coloration of most of 'thé species is a combination of black, gray,
and white fields extending along the body.

The rostrum of the skull is wide at the base and is as long as the
cranium or slightly longer. The anterior section of the ргетахШае is
flattened or slightly concave. The large pterygoid bones adjoin each other
or are slightly separated. The length of the mandibular symphysis is less
than one-fifth that of the lower jaw. The teeth are small and number
2 43. The number of vertebrae varies from 71 to 94 and their centrum
is highly flattened. The vertebrae of the lumbar section have very long,
thin spinous and transverse processes. The first two or three cervical
vertebrae are fused.

Very little is known about the biology of this genus. These dolphins
feed mainly on fish and cephalopods. Mating and parturition occur in
the summer months.

Shorthead dolphins are distributed in all the oceans in the north up
to Greenland and the Barents Sea; in the south they reach the edge of
the Antarctic ice (Fig. 269).

The genus apparently comprises six species, although some authors
recognize up to ten (Nishiwaki and Norris, 1967): 1) L. acutus Gray,
1828; 2) L. albirostris Gray, 1846; 3) L. cruciger Quoy and Graimard,
1824; 4) Г. electra Gray, 1846; 5) Г. obliquidens Gill, 1865; and 6) Г.
thicolea Gray, 1846. L. electra is sometimes considered as an indepen-
dent genus, Peponocephala Nishiwaki and Norris, 1966; this aspect calls
for further study but it would appear to deserve only a subgeneric rank.

The following species inhabit the waters of the USSR: (1) Atlantic
white-sided dolphin, L. acutus Gray, 1828; (2) white-beaked dolphin, L.
albirostris Gray, 1846; and (3) Pacific white-sided dolphin, L. obliquidens
Gill, 1865. Residence of the broadsnout dolphin, L. electra Gray, 1846,
is possible.

These animals are not hunted in our waters. (V.S.)

"(лэ.иэзту А) snyoudysouasvT ‘зищаюр реэчзлоцз Jo $пиэ8 ay] Jo э8ие “697 “314

Ss

wy 000$ 000%

0006 000Z 0001 0 0001

490

656

ATLANTIC WHITE-SIDED DOLPHIN
Lagenorhynchus (Lagenorhynchus) acutus Gray, 1828

1828. Delphinus (Grampus) acutus. Gray. Spicil. Zoologica, I, p. 2. Faroe
Islands.

1843. Delphinus leucopleurus. Rash. Nytt. Mag. Natur., vol. 4, p. 100.
Kristiana Bay (Oslo), Norway. (V.H.)

Diagnosis

Body length up to 275 cm. The body is mainly black dorsally and on the
flanks and whitish ventrally. A broad white band extends along the flanks,
roughly from the level of the dorsal fin and almost up to the caudal stem.
A narrow black band runs from the base of the flippers anteriorly and
upward toward the section between the eye and the corner of the mouth.
The dark-colored flippers are surrounded by a white field.

The pterygoid bones are often adjacent in the skull. Teeth
Vertebrae 77 - 82.

30-37
30/5 3iir

Description

The head is relatively small. The “beak” is not well demarcated from
the inclined forehead. The caudal flukes are ventrally light-colored. The
margin of the lower jaw is dark-colored. A black ring is seen around the
eye and anus (Fig. 270). The embryo bears seven or eight hairs, each
measuring roughly 1 cm, on each side of the beak.

The rostrum is slightly longer than the cranial section. The pre-
maxillae are flat and their outer edge sinuate. The temporal fossa is
elongated.

Cervical vertebrae 7, thoracic 14-15, lumbar 18-22, and caudal
38-41. Phalangeal formula: I, -5, Ij, Шб, [У› -3, and V>.

490 Fig. 270. Atlantic white-sided dolphin, Lagenorhynchus acutus (figure by М.М. Kondakov).

490

‘657

The body length of an adult female Atlantic white-sided dolphin was
225 cm (Schevill, 1956); distance from tip of snout to commencement
of flippers 40 cm, from tip of snout to dorsal fin 87 cm, and from tip of
snout to blowhole 29 cm; height of dorsal fin 24 cm, length of dorsal fin
along base 34 cm; length of flippers 30 cm; and spread of caudal flukes
65 cm.

The skull dimensions (Fig. 271) (Tomilin, 1957) (average of 10 mea-
surements) are (in cm): condylobasal length 41, length of rostrum 21,
width of rostrum at base 11, length of lower jaw (three measurements)
33, and length of mandibular symphysis (two measurements) 43. The
rostrum of the skull is shorter than in the Pacific white-striped dolphin
but longer than in the white-beaked dolphin. The teeth are usually more

numerous than in the other two species but their dimensions are smaller.
(V.S.)

Geographic Distribution

North Atlantic Ocean.

Wi

SSSe fae

Sl

==

oa
<

За.
=

ме:
Ste =

мл,

Fig. 271. Skull of the Atlantic white-sided dolphin, Lagenorhynchus acutus (figure
by N.N. Kondakov).

491

658
Geographic Range т the USSR

Barents Sea up to Spitsbergen and Murman coast of the Barents Sea up
to Kanin (Fig. 272). Transgressions are possible into our waters of the
Baltic Sea.

Geographic Range outside the USSR (Fig. 273)

In American waters from Davis Strait to the Gulf of Maine (Cape Cod).
In the eastern part of the ocean from the Barents Sea (Kanin and Spits-
bergen) and southern Greenland to the southern part of the North Sea
(England, Belgium, Denmark, and Ireland). In the Baltic Sea, only in
the southwest (waters of Denmark, Oslo Fjord, and western coasts to
Sweden), in the east not farther than Penemund close to Oder estuary
(about 18° W long.) and Koloberg (Poland; Kowalski, 1964).

This species is apparently most numerous on the coasts of Norway,
especially in the region of Lofoten Islands and Bergen. Encountered

Fig. 272. Range of the Atlantic white-sided dolphin, Lagenorhynchus acutus, in
the USSR (V.A. Arsen’ev).

492

659

Fig. 273. Species range of the Atlantic white-sided dolphin, Lagenorhynchus
acutus (V.A. Arsen’ev).

rather often on Faroe and Orkney Islands and on the coasts of Ireland
and Iceland (Jonsgard and Nordli, 1952; Tomilin, 1957, 1962). (V.A.)

Geographic Variation

Not established.
Biology

Only fragmentary information is available. Pelagic animals are
encountered more often than benthic animals in the stomach of these
dolphins: salmon, herring, mackerel, and the squid, Шех illecebrosus. On
one occasion, bottom-dwelling hermit crabs (Pagurus bernhardtii) and
mollusks (Buccinum) were found. It has been suggested that gestation
extends for about 10 months and that parturition takes place midsummer
since fully mature embryos were found in June. The newborn is 100 cm
or slightly more in length, reaching 140 cm by November.

A male 180 cm long proved to be three years old while a female
225 cm long was neither pregnant nor lactating. In Kalvag Fjord on the
west coast of Norway, a school of these dolphins arrived on March 10,
1952 and 52 were caught. Visually, many were about 2.5 m long and the
longest about 3 m. About one-third of the dolphins caught contained
embryos 55 to 70 cm long. One female contained two embryos, each
about 65 cm long.

493

660

These dolphins are usually confined to groups of 10 to 50 individuals
but at places where fishes concentrate they gather into large schools of
1,000 to 1,500 animals. Migrations have not been traced.

Beached dolphins have been found on the northeastern coast of
England, northwestern coast of Ireland, and the coasts of Holland. Once
a school of 30-35 animals was found beached on the shoals of Scotland.
Beached animals sometimes include both dolphins and pilot whales.

The helminth fauna of the Atlantic white-sided dolphin is rather
poor. The nematode Anisakis (Anisakis) simplex Rudolphi, wide-ranging
in marine mammals, was detected in the gullet, stomach, and intes-
tine. Two species of cestodes were also found: Strobilocephalus triangu-
laris Diesing, also parasitizing the intestine of the bottlenose whale and
Mesoplodon localizes in the abdominal cavity, the mesentery, and the
diaphragm. Monorygma grimaldi Monier has also been found in three
other species of dolphins (Jonsgard and Nordli, 1952; Delamure, 1955;
Tomilin, 1957, 1962).

The most characteristic external feature is the broad white band
running along the body from both sides below the dorsal fin up to the
caudal stem. This band is distinctly visible since it lies between the dark
coloration on top and the yellowish-gray color below, the latter cov-
ering the lower flanks of the body. The belly of this dolphin is pure
white.

This species is of minimal economic importance on the coasts of
Norway where up to 1,000 animals are caught in some years. In other
regions of the range, it is caught incidentally along with other species.
(V.A.)

WHITE-BEAKED DOLPHIN

Lagenorhynchus (Lagenorhynchus) albirostris
Gray, 1846

1846. Delphinus albirostris. Gray. Ann. Mag. Nat. Hist., 17; p. 84. Near
Great Yarmouth, England. (V.H.)

Diagnosis

Body length up to 304 cm. The color of the upper portion of the body
is grayish-black and the underside whitish. The base of the forehead and
the “beak” are light gray. The premaxillae are broad and flat and their
outer margin is curved. The temporal fossa is ellipsoidal. the pterygoid
bones are often adjacent. Teeth 227 (of these only 3=% are seen above

the gums). Vertebrae 88-92. (V.S.)

661

Description

This species differs from the Atlantic white-sided dolphin in having less
developed keels on the caudal stem, very large flippers, and a very high
dorsal fin. The flippers and caudal flukes are dorsally dark and somewhat
lighter ventrally. The dark color extends down along the flanks of the
body and covers the point of articulation of the flippers (Fig. 274).

The rostrum is almost as long as the cranial section.

Usually, the cervical vertebrae number 7, thoracic 14-16, lumbar
24-27, and caudal 43 - 45. Phalangeal formula: [,_ 3, Пб-7, Ш4-5, [1V,->,
and Уу-1. The body length usually varies from 270 to 300 cm.

The average main measurements of the skull (Fig. 275) (Tomilin,
1957) are (in cm): condylobasal length 44 (eight measurements), length
of rostrum 21 (eight), width of rostrum at base 14 (seven), and length of
lower jaw 36 (four). (V.S.)

Geographic Distribution
North Atlantic Ocean.

Geographic Range in the USSR

Barents Sea, Murman coast, and waters of Rybachii Peninsula; Baltic
Sea, including the gulfs of Finland and Riga (Fig. 276).

493

Fig. 274. White-beaked dolphin, Lagenorhynchus albirostris (figure by N.N. Kondakov).

662

493 Fig. 275. Skull of the white-beaked dolphin, Lagenorhynchus albirostris (figure
by N.N. Kondakov).

194 Fig. 276. Range of the white-beaked dolphin, Lagenorhynchus albirostris, in the
USSR (V.A. Arsen’ev).

495

663
Geographic Range outside the USSR

Along the American coast, it is encountered in Davis Strait, waters of
western and southern Greenland and Labrador in the north to Mas-
sachusetts Bay in the south. In the eastern part of the Atlantic, from the
Barents Sea, Iceland, and Greenland to the coasts of France (Vannes),
Great Britain, and Ireland. Waters of Denmark and southern Sweden in
the Baltic Sea (Fig. 277). Often seen in the western waters of the Baltic
Sea. (V.A.)

Geographic Variation
Not known.
Biology

Large herds of thousands of animals are seen in the summer months in
the northern parts of the range (Barents Sea, waters of Iceland, southern
Greenland, and coasts of Labrador and Newfoundland).

This species feeds mainly on benthic and bottom-dwelling fish and
other animals. The stomach of examined dolphins contained cod, capelin,

494

Fig. 277. Species range of the white-beaked dolphin, Lagenorhynchus albirostris
(V.A. Arsen’ev).

664

navaga, whiting, herring, and less frequently hermit crabs, cephalopods,
and some mollusks.

In summer they usually live in pairs or small groups but some-
times form schools of significant strength. They probably perform regular
migrations since, for example, they are encountered only in spring and
summer but disappear in autumn in Davis Strait; they are more numerous
in summer than winter in the waters of Norway and Great Britain.
Instances of beached dolphins have been reported, although compar-
atively not often, on the coasts of Orkney Islands, Ireland, Sweden,
Denmark, Holland, the Federal Republic of Germany, and France.

The period of mating is somewhat extended but is confined to the
summer months. Most females give birth in midsummer. The embryos
measured were 113 to 122 cm long but a female 305 cm long contained
an embryo 165 cm long. Yet a suckling calf found on the beach was only
122 cm long.

Two species of nematodes are known among the endoparasites.
Anisakis (Anisakis) simplex Rudolphi, wide-ranging among marine
mammals, was detected in the gullet, stomach, and intestine of the white-
beaked dolphin. It has also been recorded in 10 other species of toothed
whales, two species of baleen whales, and Steller’s sea lion in the North
Sea and in the waters of the Pacific Ocean. Halocercus lagenorhynchi
Baylis and Daubney is a parasite of the lungs (bronchi) of the white-
beaked dolphin and the bottlenose dolphin; it has been detected in
dolphins of the European waters of the Atlantic Ocean and in the waters
of Australia (Delamure, 1955).

The white-beaked dolphin is regularly hunted on the Norwegian
coasts but the-volume of hunting is minimal. Previously hunting was
relatively rewarding in Davis Strait but not at present, while the catch
on the coasts of Great Britain is incidental. Hence this species is of very
little economic importance. (V.A.)

PACIFIC WHITE-SIDED DOLPHIN

Lagenorhynchus (Lagenorhynchus) obliquidens
Gill, 1865

1865. Lagenorhynchus obliquidens. Gill. Proc. Ac. Sc. Philadelphia, 17,
p. 177. Near San Francisco, California.

1955. Lagenorhynchus ognevi. Sleptsov (Slepzov). Tr. Instituta Okeano-
logii AN SSSR, 18, р. 60, 15-20 km east of Kunashir Island, Kuril
range. (V.H.)

496

496

665
Diagnosis

Body length up to 230 cm. The body is blackish on top and white under-
neath, the flanks grayish with a dark field in the middle. The premaxillae
are rounded and their outer edge slightly sinuate. The temporal fossa is
large, rounded. The pterygoid bones are often separated. Teeth 3=¥.
Vertebrae 74. (V.S.)

Description

Body coloration is subject to considerable individual variation. Usually
the upper side of the body, end of the snout, anterior portion of the dor-
sal fin, and the flippers are dark-colored. The flanks are lighter in color.
A narrow black band runs on each side from the base of the flippers
anteriorly and upward toward the corner of the mouth and posteriorly
along the body up to the lower caudal stem (Fig. 278). Sometimes two
symmetrical longitudinal narrow white bands run along the dorsum, com-
mencing along the flanks near the blowhole. These bands later run onto
the flanks anterior to the dorsal fin and continue up to the caudal stem.

The integument is characterized by a thin dermal layer with abundant
elastin fibers which are found in small bundles at the border of the
subcutaneous fatty tissue and subcutaneous musculature. The embryos
have whiskers on the snout.

The length of the rostrum of the skull (Fig. 279) is roughly equal to
or slightly more than the length of the cranial section. The usual dental

formula is о. The diameter of the largest tooth does not exceed

Fig. 278. Pacific white-sided dolphin, Lagenorhynchus obliquidens (figure by
М.М. Kondakov).

496

497

666

О <<

<We

ня НЕЕ
CLL ID

SS
ее.

BX
О

SS ‘
22a Sw
2

SS

SS

SS

Fig. 279. Skull of the Pacific white-sided dolphin, Lagenorhynchus obliquidens
(figure by N.N. Kondakov).

5 mm. Cervical vertebrae 7, thoracic 13-14, lumbar 20-24, and caudal
30-34. Phalangeal formula: I, _5, I¢_, II, [V>_3, and V,_>.

- The main body measurements of the Pacific white-sided dolphin
(Wilke, Taniwaki, and Kuroda, 1953) averaged for four females and ten
males are respectively (in cm): body length 172 and 167; distance from
tip of snout to dorsal fin 112 and 106, to anterior margin of base of
flippers 43 and 42; distance from anus to caudal bifurcation 50 and 49;
maximum width of caudal flukes 50 and 44; height of dorsal fin 20 and
18; length of flippers 30 and 28 and maximum width 11 and 10. The
length of the largest male was 222.6 cm and of the female 221 cm.

Females weigh 67 - 93 kg, average 80 kg, and males 58 - 84 kg, average
66 kg (Japanese Hunting; Wilke, Taniwaki, and Kuroda, 1953).

The average skull dimensions (Tomilin, 1957) are (in cm): condy-
lobasal length 40 (ten measurements), length of rostrum 22 (eight), width
of rostrum at base 10 (eight), zygomatic width 20 (six), length of lower
jaw 34 (eight), and length of mandibular symphysis 4 (five). (V.S.)

Geographic Distribution

North ‘Pacific Ocean.

499

667
Geographic Range in the USSR

Coasts of the Sea of Japan, including Peter the Great Gulf and Amur and
Vityaz’ Bay. Waters of Kuril Islands—Shikotan, Iturup, Urup, Kunashir,
and Paramushir on the Pacific Ocean as well as the Sea of Okhotsk side
(Fig. 280).

Geographic Range outside the USSR (Fig. 281)

Coastal waters of Honshu and Hokkaido Islands, Pacific Ocean coast
of North America from the Gulf of California (Mexico) to the Gulf of
Alaska and the Aleutian Islands. (V.A.)

Geographic Variation
Not studied.
Biology

Population. One of the abundant species of Pacific dolphins. In some
parts of the range, it sometimes forms schools of thousands of animals.
It is numerous among our Kuril Islands on the Sea of Okhotsk as well
as the Pacific side.

Food. Small fish and cephalopods which form large concentrations
serve as the main food. Lantern fishes—Scopelidae (Myctophidae)—are
of utmost importance and represent 77% of the total intake by volume,
Japanese anchovy (Engraulis japonica) 9%, Pacific mackerel (Scomber
japonicus) 7%, and squids (“beaks” and eye lenses probably of Watasenia
scintillans) also 7% (Wilke, Taniwaki, and Kuroda, 1953). In the coastal
waters of North America, the stomach of dolphins contained herring,
salmon, sardine (Sardinops caerulea), saury, northern anchovy (Engraulis
mordax), scad (Decapterus paliaspis), squids including Loligo opalescens,
and jellyfish. In Nemuro Strait, the stomach of a female contained squids
(Ommatostrephes sloanei-pacificus) and scad [Japanese horse mackerel]
(Trachurus japonicus) (Scheffer and Slipp, 1948).

Lantern fishes are of major importance in the food intake of these
dolphins in the waters of Japan. Squids were found in almost every stom-
ach dissected but in small quantities. But their importance is undoubt-
edly far greater than revealed by analysis based on the volume ratio in
the intake since the stomach contents contained only the remnants of
squids, such as “beaks” and eye lenses (100 to 200 “beaks” in one stom-
ach). Apparently the small schooling fish and squids constitute the main
food of dolphins in this region.

497

668

0 250 500 750 1000 km

Fig. 280. Range of the Pacific white-sided dolphin, Lagenorhynchus obliquidens,
in the USSR (V.A. Arsen’ev).

Daily activity and behavior. These dolphins are usually more active
in the day. They are encountered in small groups as well as in large
herds. Herds comprising animals of different ages predominate in the
summer months but sometimes separate groups of young dolphins are
also encountered.

These dolphins are more often seen in large bays and gulfs than
in the open sea. They quite often follow a ship but on closing in on
it, dive to a depth of 1-3 m. Sometimes playful dolphins leaping high
above the water land on their side or back with a loud thud and raise

669

‘(ao,uasty А) suapinbygo snysukysouasyT чиц@ор рэр!з-эицл эцоеа ay) Jo э8ие1 $910э4$ ‘185 “S14

ост Ost

86?

500

499

670

much spray (Fig. 282). These dolphins are encountered in groups with
other species also. On one occasion an adult dolphin trying to rescue a
harpooned comrade was observed. It dove frequently between the ship
and the injured dolphin and each time pushed it farther away from the
ship. These animals survive well in tanks, live long, and can be trained
well. They quickly learn to play with a ball, jump through a hoop, etc.,
and to snatch food from the trainer’s hand (Tomilin, 1962).

Migration. Seasonal migrations probably do occur but the informa-
tion available is highly fragmentary. In the waters of California, these
dolphins are noticed in the winter months (November, December, Febru-
ary, March, and April); on the coasts of Washington state in March and
later in July, August, September, and November; on the coast of British
Columbia in July;.and in the waters of the Alaskan peninsula from July
to September.

Judging from the Japanese hunting experience, these dolphins are
encountered in early March on the Pacific Ocean coasts of Honshu at
36° М lat. and in mid-March at 39 to 40° М lat. where they remain
until June, partly advancing farther north toward Hokkaido coasts. In
summer and autumn, large schools are seen in the coastal waters of the

Fig. 282. Pacific white-sided dolphins leaping above the water, East China Sea,
March, 1968 (photograph by A.V. Kucheryavenko).

671

Kuril Islands in the Pacific Ocean and in the Sea of .Okhotsk. A large
school was found on August 1, 1951, feeding on anchovy in the region
of Chetvertyi Kuril Strait; at the end of August, a herd more than 1,000
strong was sighted on the Pacific Ocean side of Iturup Island where saury
concentrations were present. At the end of September, a large herd of
these dolphins was noticed southeast of Shikotan Island where, too, a
school of saury was encountered. Hunting on a small scale is resumed in
the autumn months on the coasts of Japan (Klumov, 1959).

Reproduction, growth, and development. Mating and parturition occur
in summer. An embryo 22 mm long, detected in September, had grown to
370 mm by December and 761 mm by March. A male 124 cm long turned
out to be a suckling calf (weight 29.5 kg). The ovaries of a 171-cm long
female (weight 62 kg) did not contain corpora lutea but the Graafian
follicles were well developed; this female was close to attaining sexual
maturity (Houck, 1961). Females 180 cm or longer were already with
embryos. In oceanariums, mating of female white-sided dolphins with
male bottlenose dolphins has been observed time and again (Tomilin,
1962).

The parasites of these dolphins have not been studied. (V.A.)

Economic Importance

These dolphins are regularly hunted along with other species only in the
waters of Japan. Their annual catch does not exceed 1,000 head. The
weight of the females varies from 67-93 kg (mean 80 kg) and of males
from 58-84 kg (66 kg). They are of little economic importance.

Hunting is carried out using the same small ships (20 to 30 tons)
and the very same methods as practiced in hunting other species of dol-
phins. The dorsal fin, flippers, and caudal flukes are first separated from
the trunk of the killed animal. The skin is then peeled off in two layers
together with the subcutaneous fat and the flesh is separated in large
chunks from the vertebral column. The intestine, heart, and other inter-
nal organs are then cut out and the ribs chopped. The flesh, heart, lungs,
and liver are used as food. The skin is tanned to make low-quality leather
goods. The blubber is melted into oil; the blubber from the head and in
the hollow of the jaws can be converted into a high-grade machine oil.
The skeletal bones as well as all the remnants are used for making fertil-
izer meal (Wilke, Taniwaki, and Kuroda, 1953; Siebenaler and Caldwell,
1956; Klumov, 1959; Tomilin, 1962).

Large schools of these white-sided dolphins prevailing in the waters
of the Kuril Islands suggest the possibility of their organized hunting
here. (V.A.)

501

501

672
BROADSNOUT DOLPHIN

Lagenorhynchus (Peponocephala) electra Gray, 1846

1846. Lagenorhynchus electra. Gray. Zoology. Voyage Erebus and Terror,
1, p. 35. Type locality not established.
1848. Delphius pectoralis. Peale. U.S. Explor. Exped., 8, p. 32. Hawaiian
Islands.
1868. Electra obtusa. Gray. Synopsis Whales Dolphins. Brit. Mus., 7.
Substituted for Lagenorhynchus electra Gray, 1846. (V.H.)
The broadsnout dolphin differs significantly from all other species of the
genus Lagenorhynchus in several characteristics: it has no beak whatso-
ever or such is barely visible and not demarcated from the forehead; a
uniform arcuate line of the profile is formed from the blowhole to the
edge of the mouth. The preorbital depression is larger than in the other
species of the genus. The first three vertebrae, and not the first two as in
the other species, are fused. The general body shape is more elongated,
with a relatively longer caudal stem. The color is monochromatic, dark,
without black, gray, or white fields; a light-colored (white) field is seen
only in the cervical zone and on the breast; sometimes, a similar field is
seen in the posterior section of the abdomen (Fig. 283).

This is a somewhat peripheral form of the genus and reveals fea-
tures of similarity and association with the genera Pseudorca (false killer
whale) and Feresa (dwarf killer whale). (V.H.)

Its range is known only from some fragmentary information.

Its presence has not been established in USSR waters but residence
in or transgressions into the Sea of Japan and waters of the Kuril Islands
are possible.

Outside the USSR, it is found in tropical seas up to north of 10° N
lat. Its presence has been indicated in the waters of Senegal, the Gulf
of Guinea, India, Indonesia, Hawaiian Islands, and Japan (transgression

Fig. 283. Broadsnout dolphin, Lagenorhynchus (Peponocephala) electra (figure
by N.N. Kondakov).

501

673

SS
Va

LN

RS Е W BS

%

у) Я] ИИ
GR ny
Gis il sis
Meat

, ) ys \
И

Fig. 284. Skull of the broadsnout dolphin, Lagenorhynchus (Peponocephala)
electra (figure by N.N. Kondakov).

of a herd of 500 animals in 1965 into Suruga Bay on the ocean side of
Honshu; Nishiwaki and Norris, 1967). (V.H.)

Biology is almost not known. Very rare. In January, 1963, fourteen
dolphins were caught in a net in one of the bays on the Pacific Ocean
side of Honshu Island. All of them showed their heads above the water
most of the time, thus exhibiting near-total fearlessness of man. Held
in an Oceanarium they exhibited no restlessness and were at peace with
man (Nishiwaki, 1965). (V.A.)

Genus of False Killer Whales

Genus Pseudorca Reinhardt, 1862

1846. Phocaena crassidens. Owen. Hist. Brit. Foss. Mamm. p. 516.
1862. Pseudorca. Reinhardt. Overs. Danske Vidensk. Selsk. Forh., p. 151.
(V.H.)
Medium-size dolphins, with body length up to 6 m.
The body is elongated. The small rounded head has a fairly large adi-
pose body [melon]. The obtuse “beak” is barely perceptible. The upper
jaws are slightly longer than the lower. The relatively small dorsal fin

502

674

lies midbody and is deeply notched posteriorly. The о are narrow
and pointed at the tip.

The body is black but the ventral side somewhat lighter.

The rostrum is short, not longer than the cranium, but broad. The
premaxillae are very broad and identical throughout their length. Their
anterior margin is rounded. The reduced pterygoid bones are adjacent.
Teeth a. Vertebrae 50-51. Phalangeal formula: I, 5, Пб_з› III;_¢,
IV,_,, and V,_>. The sternum consists of four sections. Ribs 10 ‘pairs,
of which four articulate with the sternum.

False killer whales feed mainly on schooling fish and cephalopods.
Mating and parturition are protracted.

These whales are distributed in the warm and temperate waters of
the Pacific, Indian, and Atlantic oceans. They are not caught in USSR
waters but might possibly be found along the Kuril Islands and in the
Sea of Japan and the Baltic Sea.

Fossils of false killer whales have been detected in the Upper
Pliocene of Europe and Japan.

The genus comprises a single species: the false killer whale, P. cras-
sidens Owen, 1816. (V.S.)

FALSE KILLER WHALE

Pseudorca crassidens Owen, 1846

1846. Phocoena crassidens. Owen. British Fossil Mamm. and Birds,
p. 516. Lincolnshire, England (subfossil specimen).

1882. Pseudorca mediterranea. Giglioli. Zool. Anzeiger, 5, р. 268.
Mediterranean Sea. (V.H.)

Diagnosis
Only species of the genus.
Description

The body is elongated, spindle-shaped, and the head relatively small. It
somewhat resembles the killer whale but the body build is less compact,
the head more elongated, and the dorsal fin considerably shorter. The
flippers are not rounded but narrow and pointed (Fig. 285).

Very light-colored star-shaped scars are often seen on the black sur-
face of the body. A gray longitudinal band sometimes occurs on the
midabdomen and in the region of the urogenital opening.

Teeth usually number 5. Unlike those of the killer whale, their cross
section is not oval but circular.

675

503

503

Fig. 285. False killer whale, Pseudorca crassidens (figure by М.М. Kondakov).

The width of the rostrum increases with age and can be corre-
lated with the intense growth of the adipose body [melon]. Cervical
vertebrae 7, thoracic 10, lumbar 9-11, and caudal 22-24. The anterior
and sometimes all the cervical vertebrae are fused.

The indices of the main body measurements of the false killer whales
as percentage of the total length of males (three specimens) with a body
length of 279-465 cm and females (two) with a body length of 251 and
267 cm (Tomilin, 1957) are respectively: distance from tip of snout to
blowhole 9.6-12.4 and 11.7-12.3, up to base of flippers 14.5 - 19.6 and
15.3-21.4, up to posterior margin of dorsal fin 51.2 -62.0 and 63.2 - 71.7;
length of base of dorsal fin 15.2- 15.7 and 13.8 - 14.7, height of dorsal fin
8.3-8.6 and 7.4-8.3; length of flippers 11.8-13.2 and 11.5-12.6, maxi-
mum width of flippers 4.7 -5.0 and 4.6-5.0; width of caudal flukes (from
tip to tip) 24.0-24.4 and 20.9-21.4. The largest of the false killer whales
caught measured 596 cm in length. Males are 0.6-1.0 m longer than
females. False killer whales weigh up to 1.5 tons.

The main dimensions of the skull (eight measurements) of adult false
killer whales (Tomilin, 1957) are (in cm): condylobasal length 54-62,
length of rostrum 27 - 30, width of rostrum at base 18-21, and length of
lower jaw 46-51 (Fig. 286).

The vertebral sections constitute (as percentage of length of the ver-
tebral column): cervical 3, thoracic 17, lumbar 31, and caudal 49 (Slijper,
1936). (V.S.)

Geographic Distribution

Encountered in temperate and warm waters of the Northern and South-
ern hemispheres (Fig. 287).

676
Geographic Range т the USSR

Waters of the Kuril Islands (apparently the southern islands) and
probably the Sea of Japan. Transgressions into our waters of the Baltic
Sea are quite possible.

Geographic Range outside the USSR

Encountered in the Atlantic Ocean everywhere from the North Sea and
North Carolina® to the South African and South American coasts (Tierra
del Fuego). It has been reported on the eastern (Baltic) coasts of Den-
mark, western and partly eastern (evidently the southernmost) coasts of
Sweden, and on the Baltic coasts of the Federal Republic of Germany
(Kiel Bay). Not reported on the coasts of Poland.

It has been reported in the Pacific Ocean from Washington state in
the north to the coasts of Peru, probably Chile, New Zealand, Australia,

222

Li

0222222 7 2 2042

==

>
метки

2222

WZ,

TES APE LO

A JAN

EB

ася

о

ЕЕТА ree

2... Ма
7 a
» Ау \\ am
See Meas Saheb ns
м, * a ко т w & |

wv

А Te

`

yf

503 Fig. 286. Skull of the false killer whale, Pseudorca crassidens (figure by М.М. Kondakov).

8 References to habitation farther north (Davis Strait and Aleutian Islands; Tomilin,
1957, 1962) have either not been confirmed (Murie, 1959; Hall and Kelson, 1959; Manvill
and Joung, 1965), or have been questioned, or treated as erroneous (Hershkovitz, 1966).
(V.H.)

OE
ROMO

ily iy
TAH
“Teall | [lay

fil
ryt

9
и

1 | ПА
Ни
СЫН

1 || Wy Ils
hill

‘
$5 We
реет
4 ИН и

1000 0 1000 2000 3000 4000 5000 km

Fig. 287. Species range of the false killer whale, Pseudorca crassidens (У.А. Arsen’ev).

504

505

506

678

and Tasmania in the south; it has been reported in the Indian Ocean in.
the warm and temperate water zones.

Numerous beached animals have been found on the coasts of Great
Britain, Denmark, Holland, the Baltic and Mediterranean seas in the
eastern part of the North Atlantic; in the western part, on the coasts
of North Carolina and Florida states, Cuba, Venezuela, Brazil, and
Argentina. In the Pacific Ocean, beached animals have been found on
the Kuril Islands, the coasts of Japan, Galapagos Islands, California, and
Mexico; and in the Indian Ocean, on the coasts of Sri Lanka, India, South
Africa, and Australia (Tomilin, 1962; Mitchell, 1965; Fiscus and Niggol,
1965; Nishiwaki, 1966). (V.A.)

Geographic Variation

Not established.
Biology

As the range of the false killer whale is very large and instances of
beaching (sometimes in large herds) are well known, this species could
correctly be classed as a relatively abundant one.

Food. Fish (e.g., haddock, cod, and salmon) and some species of
cephalopods serve as the food of the false killer whale; these animals
quite often come close to the coast in pursuit of food objects. It is at
such times that they become stranded on the shoals.

Behavior. False killer whales more often live far from the coasts in
the open sea in small groups although sometimes they do form large
herds (Fig. 288). Instances of large herds of these dolphins being cast
on the beach are known. For example, in Mar del Plata in Buenos Aires
province of Argentina, 835 false killer whales were stranded on a sandy
beach on October 10, 1946 and perished soon thereafter. They suffered
from extreme fatigue and had slowly sunk into the sand (Obruchev,
1949).

Migrations of false killer whales have not been studied.

Reproduction. Judging from the fact that embryos of different sizes
are encountered simultaneously, mating and parturition are undoubt-
edly protracted among these dolphins. An embryo 117 cm long was well
pigmented while another embryo 160 cm long was almost mature. The
largest of the embryos examined was 183 cm long. It has been reck-
oned that females attain sexual maturity at a body length of 366 to
427 cm. An examination of many groups of false killer whales that per-
ished through beaching established a male:female ratio of almost one in
each herd.

505

Fig. 288. False killer whales at sea (figure by М.М. Kondakov).

Enemies, diseases, parasites, mortality, and competitors. Skin parasites
have not been reported but two species of helminths have. The nematode
Anisakis (Anisakis) simplex Rudolphi, parasitizing the gullet, stomach,
and intestine, has been found in ten other species of toothed whales,
two species of baleen whales, and in Steller’s sea lion, as well as in
the false killer whale. It has also been detected in various species of
marine mammals in the North Sea, on the coasts of Kamchatka, Japan,
and New Zealand. The acanthocephalan Bolbosoma capitatum Linstow
parasitizes the intestine of the false killer whale, the pilot whale, and the
sperm whale in the Atlantic Ocean and the Mediterranean Sea. (V.A.)

Economic Importance

There is no special hunting for the false killer whale; it is caught inci-
dentally while hunting for other species. In some cases, the carcasses
of beached animals are utilized. The overall weight of a false killer
whale may reach 1.5 tons (Delamure, 1955; Sleptsov, 1955; Tomilin, 1957,
1962). (V.A.)

Genus of Killer Whales
Genus Orcinus Fitzinger, 1860

1846. Orca. Gray. Zoology. Voyage Erebus and Terror, I, p. 33. Orca
gladiator Gray = Delphinus orca Linnaeus, 1758. Мот. ргаеосс.
—Orca Wagler, 1830 = Hyperoodon Lacépede.

1860. Orcinus. Fitzinger. Wiss. Popul. Naturg. Saugeth., 6, p. 204.

507

680

1870. Gladiator. Gray. Proc. Zool. Soc. London, р. 71. Delphinus orca
Linnaeus, 1758.

1933.? Grampus. Iredale and Troughton. Rec. Austr. Mus., 19 (1), p. 28.
Delphinus orca Linnaeus, 1758. (V.H.)

Dolphins of large proportions, being the largest members of the family.

Body length up to 10 m.

The head is somewhat flattened dorsoventrally. There is no “beak”.
The broad flippers are oval in shape. The dorsal fin is very high, especially
among males (up to 1.7 m). The overall color of the body along the upper
side and flanks is black while the belly and neck are white. A white patch
occurs on each side posterior to the dorsal fin and on the temple above
the eye (sometimes these patches are lacking).

The broad and flattened rostrum is slightly shorter or as long as the
cranium. The maxillae are well developed and very broad. The occipital
crest is absent in the young but high among the adults, especially males.
The temporal fossa is large. The teeth are large, number Я, com-
pressed front to back and have extremely well-developed roots. Verte-
brae 50-52. The sternum consists of three or four sections.!° The length
of the phalanges of the digits and the metacarpals is less than their width.
Phalangeal formula: 15, Il,_7, Ш4_5, [V3_4, and V>_ 3.

The killer whale is sarcophagous and attacks even large whales. The
periods of mating and parturition are protracted. Gestation probably
requires one year.

These whales are distributed in the World Ocean except in glacial
regions of the Arctic and the Antarctic.

Fossil remains have been detected in the Middle Pliocene of Europe.

The genus comprises a single species: the killer whale O. orca Lin-
naeus, 1758.

There is no special hunting of the killer whale. (V.S.)

KILLER WHALE

Orcinus orca Linnaeus, 1758

1758. Delphinus orca. Linnaeus. Syst. Nat., ed. X, I, 77. Eastern part of
the northern half of the Atlantic Ocean (“Oceano Europaeo”).

3 The introduction of the name Grampus to denote the genus of killer whales is asso-
ciated with so many inaccuracies that it must be considered irrational from the viewpoint
of nomenclature. Yet the name (Grampus orca) did prevail in the literature for sometime.
Today, as earlier, it is used only in reference to Risso’s dolphin (Grampus griseus). (V.H.)

10 According to other data, the sternum is not segmented (Sleptsov, 1955).

508

681

1789. Delphinus gladiator. Bonnaterre. Tabl. Encycl. Méth. Cétologie,
p. 23. Spitsbergen, Davis Strait, New England.

1869. Orca ater. Cope. Proc. Ac. Nat. Hist. Philadelphia, 21, p. 22. North-
ern Pacific Ocean (“Northwestern coasts from Oregon to Aleutian
Islands’’).

1869. Orca rectipinna. Cope. I[bid., 21, р. 22. Californian coast.

1874. Orca ater var. fusca. Dall. In: Scammon. Marine Mammals of the N.
W. coast of N. America, p. 298. Coasts of California and Oregon.
(У.Н.)

Diagnosis
Only species of the genus.
Description

This whale has a strongly built body, a fairly large head, a large
mouth, and is a powerful predator. Large rounded flippers (Fig. 289) are
a characteristic feature. Another distinctive feature of this whale is the
high dorsal fin, which is straight along the posterior margin.

This whale exhibits distinct sexual dimorphism in the dimensions of
the dorsal fin, which is considerably larger in the males. The size and
relative dimensions of the flippers in both sexes and the caudal flukes
among males increase with age. Unlike in other whales, the head in the
killer whale (Fig. 290) becomes relatively shorter with advancing age
while the caudal section becomes elongated, i.e., the elongation of the
head is relatively less compared to the caudal section (Ivanova, 1959).

The white coloration on the belly is divided posteriorly into three
tongues, of which the two lateral ones (right and left) terminate behind
the anal opening while the middle one may terminate at the same point
or run along the right up to the caudal flukes (which then merge with
the white coloration), or be altogether absent. Sometimes the right and
left white tongues fuse posterior to the dorsal fin. Wholly dark-colored
killer whales with only a white spot under the eye are encountered in
the Far East.

Unlike most other dolphins, the integument is characterized by a
well-developed dermal layer with a dense network of fascicles of col-
lagen fibers. Small processes project into the dermal papillae from the
epidermal septa. Up to seven hairs are seen on the upper jaw of embryos.

Sex- and age-related variations are distinctly seen in the skull. The
temporal fossa and occipital crest enlarge with age. Among adult males,
indices of the width of the rostrum are slightly higher than in the young.
The lower jaw in males is relatively fonger than in females. The occipital
crest of adult males is larger than that of females.

682

508 ‘Fig. 289. Killer whale, Orcinus orca (figure by М.М. Kondakov), A—male; B—female.

508 Fig. 290. Head of the killer whale, Orcinus orca (figure by N.N. Kondakov).

509

683

The teeth are covered with enamel and are very strong. When the
mouth is closed, the teeth on the upper jaw fall in the gaps between the
lower teeth, thus forming a powerful gripping apparatus. The teeth of
the killer whale, compressed front to back, have apparently adapted to
feeding on large quarry from which the killer whale can tear off large
chunks of flesh. The foreteeth, inclined slightly forward and outward,
are capable of withstanding extremely powerful jerks of the quarry while
the middle and hind teeth perform the function of holding it firmly
(Tomilin, 1957). The surface of the teeth is sometimes worn down to
the pulp cavity. Vertebrae: cervical 7, thoracic 11-12, lumbar 10, and
caudal 21-24. The first two to three vertebrae are usually fused. The
percentages of the different sections of the vertebral column to its total
length constitute: cervical 3.2, thoracic 22.9, lumbar 28.0, and caudal 45.9.
Ribs 11-12 pairs, of which 5-7 anterior pairs articulate with the third
or fourth section of the sternum (Tomilin, 1957).

The maximum length of 320 male killer whales caught in the waters
of Japan during 1948 through 1957 was 9.4 m, of 247 females 8.2 m.
Among these killer whales, males with a body length of 6.3 m and females
6.0 were more common. Males of maximum proportions attain a body
length of 10 m and females 8.2 m (Nishiwaki and Handa, 1958).

The main body dimensions of males (based on measurements of two
to four animals) and females (one to three animals) vary as follows (in
cm): body length 579-830 and 495-670; distance from tip of snout to
commencement of dorsal fin 245-400 and 335, up to blowhole 80-90
and 75, up to anal opening 221-273 and 225; length of base of dorsal
fin 77-91 and 62, height of dorsal fin 121-170 and 52-95; and width of
caudal flukes 195-279 and 99 - 139.

The main measurements of the skull (Fig. 291) of five adult males
and one adult female killer whale (Tomilin, 1957) are respectively (in
cm): condylobasal length 100-112 and 91, zygomatic width 68-78 and
57, length of rostrum 51-57 and 48; width of rostrum at base 34-37 and
27, and length of lower jaw 85-95 and 74. (V.S.)

Geographic Distribution

The killer whale represents a cosmopolitan species. It inhabits coastal
and oceanic waters of the world’s oceans, including the Arctic and
Antarctic seas.

Geographic Range in the USSR

Barents Sea, northwestern and western parts of the Kara Sea (does not
transgress into its eastern part nor into the Laptev Sea), coastal waters of

509

684

SSS SSS

Fig. 291. Skull of the killer whale, Orcinus orca (figure by N.N. Kondakov).

Novaya Zemlya, Murman coast, Kil’din Island, Rybachii Peninsula, and
the White Sea (Fig. 292). Transgressions into our waters of the Baltic
Sea are highly possible (transgression into bay near Gdansk is known).

In the Pacific Ocean waters of the Soviet Union, this species is dis-
tributed everywhere in the seas of Japan and Okhotsk, around the Kuril
Islands, in the Bering Sea, near the Commander Islands, in the Bering
Strait, and in the southern part of the Chukchi Sea (probably north of
70°N lat.).

Geographic Range outside the USSR

From the coasts of Spitsbergen, Greenland, and Baffin Bay in the north
to the southern waters of the Atlantic Ocean. It is quite common in
much of the eastern and western halves of the Atlantic (coastal regions
of Great Britain, Norway, Holland, France, western Africa, Atlantic coast
of North and South America, and the Mediterranean Sea). It inhabits
the westernmost, southwestern, and southern parts of the Baltic Sea up
to Gdansk (Danzig), including the bay (Fig. 293).

In the Pacific Ocean, it is encountered off the coasts of the Korean
peninsula, Japan, Aleutian Islands, and Alaska, including its northern
half (noticed off Cape Barrow, 71°24’N lat.), off the Pacific coast of
North America and Mexico, and along the coasts of South America from
Panama to Tierra del Fuego. It lives in the Indian Ocean and along

‘(лэмэзту У) USSN Э41 Ul ‘9240 5ии1240 ‘ЭТеЧм ло BY1 JO suey “767 “Bly отб

от 691
ах

|
Wy 0001 OSL 005 055 O 052

‘(Ad цэзту УД) 0240 5пи1240 ‘TRUM ты эЧ1 Jo э8иел $э0э4$ ‘©6291 119

wy 000$ 006 0006 0005 COOL 0 0001

512

687

the coasts of Australia and New Zealand. It is widely distributed in the
Antarctic where it reaches the high southern latitudes, right up to the
zone of permanent ice. (V.A.)

Geographic Variation

In spite of the extensive distribution of this species in the World Ocean,
no geographic variation has been established. Sometimes the Pacific
killer whale is regarded as a species (Subspecies), Orcinus rectipinna Cope,
1869, but the characteristics of this form have not been established, albeit
mention has been made that its dorsal fin is not as high as in the Atlantic
species (Hall and Kelson, 1959). (V.A.)

Biology

Population. One of the abundant species of dolphins. It is regularly
encountered in the World Ocean (usually in small groups) and is fairly
uniformly distributed over its immense range.

Food. The killer whale is the only true carnivore in the order of
cetaceans and even consumes warm-blooded animals. Its food is quite
diverse: chum salmon, chinook salmon, coho salmon, cod, capelin, hal-
ibut, skate, shark, herring, smelt, etc. Cephalopods are consumed quite
often and sometimes various types of marine birds. The killer whale
is known to consume seals and small dolphins and to attack fur seals,
Steller’s sea lion, and even walruses (an instance of a group of killer
whales attacking a herd of walruses was observed from an airplane;
Zenkovich, 1938b). There are frequent references in the literature to
the killer whale lying in wait around the coastal rookeries of fur seals
and Steller’s sea lion but the many observations reported in recent years
do not confirm this information (V.A. Arsen’ev). Reports of the killer
whale attacking balsen whales and tearing out their tongues and large
chunks of blubber, and even killing large animals, have been copied
from book to book from the middle of the last century without crit-
ical review. Even a recent and thorough publication (Tomilin, 1957)
describes the methods of killer whale groups attacking large whales, pre-
senting information copied from various sources. Such reports give rise
to considerable doubt.

A study of the stomach contents of several hundreds of killer whales
caught in a ten-year period in the waters of Japan revealed that squids
(octopuses to a very small extent) and fish (including sharks) represent
the main food while marine mammals play a secondary role. The number
of stomachs with remnants of dolphins was more than double the num-
ber of stomachs with remnants of seals. Among fish, cod predominated,

513

688

followed by (using the source terminology) flatfish (probably flounder,
halibut, etc.). Sardines, salmon, and tuna were of lesser importance. Some
stomachs revealed mackerel, greenling, sea perch, and bonito, which have
no practical importance as food for the killer whale. Dolphins of three
species exclusively represent the cetaceans found in the stomach rem-
nants: Dall porpoise (Phocoenoides dalli dalli and Р. 4. truet) found in
the stomach of killer whales on Hokkaido coasts and blue-white dolphin
(Stenella coeruleoalba) in killer whales from the Pacific waters off Hon-
shu. There were some finds of pilot whales, beaked whales, and two cases
of sei whales (chunks of flesh). Young killer whales up to 4.3 m long
feed exclusively on squids and fish. They begin attacking seals and dol-
phins as their body proportions increase and thus remnants of seals and
dolphins are found in the stomach of only older killer whales (Nishiwaki
and Handa, 1958). In July and August, 10 stomachs of killer whales were
found filled with remnants of fish and squids (11 other stomachs in the
same months were empty). No remnants whatsoever of marine mammals
were found (Ivanova, 1961c).

In the eastern half of the North Pacific Ocean, 10 stomachs of killer
whales were investigated. Of these, one was caught close to Kodiak Island
in the Gulf of Alaska, five near San Francisco, and four in Californian
waters. Two of these stomachs were empty while the remaining eight con-
tained the remnants of marine mammals (California sea lion, Steller’s
sea lion, elephant seals, two species of dolphins, and Minke whale), hal-
ibut, ocean sunfish, shark, and squids. The frequency of encounter of the
remnants of these food objects is shown below.

Quantitative characteristics of the food of
killer whales (number of finds)
(Rice, 1968)

California sea lion, Zalophus californianus
Steller’s sea lion, Eumetopias jubatus
Elephant seal, Mirounga angustirostris
Harbor porpoise, Phocoena phocoena
Dall porpoise, Phocoenoides dalli

Minke whale, Balaenoptera acutirostrata
Opah, Lampris regius

Pacific halibut, Hippoglossus stenolepis
Carcharinid, ? Prionace glauca

Squids

BPNFPNPNN IA W

The above eight killer whales with remnants of marine mammals in
their stomachs included six adult males, one adult female. (California),

689

and one immature male (Gulf of Alaska). The stomach of five adult males
contained only the remnants of marine mammals while the sixth addi-
tionally had remnants of squids. The stomachs of the adult female and the
immature male had only the remnants of fish, including sharks. It may be
assumed that male killer whales feed predominantly on marine mammals.

The mammals were represented mostly by the remnants of one, less
frequently two, and only in one case (elephant seal) four animals in the
same stomach. All of them were thoroughly digested, since mostly the
teeth, nails, and sometimes ribs or skin remnants were found. Minke
whale was represented by baleen plates and chunks of blubber. Layer
formations on the teeth helped to establish the age of some of the pin-
nipeds consumed by killer whales. Thus the stomach of a male killer
whale 7.6 m long contained the remnants of a nine-year-old sea lion,
of another male killer whale 6.86 m long the remnants of a one-year-
old Steller’s sea lion (probable weight of the animal 90 kg), and of yet
another male 7.24 m long a two-year-old (probably) Steller’s sea lion
(Rice, 1968).

The hunting of sea lions by killer whales has been described as
follows: a group of five killer whales chased a large male sea lion on
May 25, 1965 in the San Francisco region. The adult killer whale in the
group (it also contained calves) continuously lay under the sea lion, or
above it when the sea lion dived. From time to time, the killer whales
struck the quarry with their bodies or attempted to crush it. The sea
lion jumped from side to side, sometimes leaped out of the water, and
eventually became exhausted. At this point one killer whale swam to its
right, another underneath it, while a third grabbed the quarry with its
teeth and towed it under water. That was the last sighting of the sea lion
on the water surface. This happened very close to a ship following the
attacking killer whales; the latter paid no attention to the ship behind
them. In another case, one of the six killer whales (the group contained
two calves and one adult male) appeared on the water surface holding
a small sea lion in its mouth. The nature of mammal remnants seen in
the stomach of killer whales suggests that they dismember large animals
before ingestion but swallow smaller prey entire (Rice, 1968).

Reports of killer whales attacking large whales require documentary
proof. In the Antarctic, near a whaling base, groups of killer whales
were noticed floating alongside fin whales, humpback whales, and other
species of baleen whales time and again. Yet no attempts by the killer
whales to attack the others were ever recorded. Concomitantly, the other
whales (even when alone) never exhibited signs of restlessness on the
approach of a herd of killer whales. However, killer whales regularly
tore out the tongues of dead and air-filled whales with mouths wide open

514

514

690

(Fig. 294). The carnivores were bold enough to tear out the tongues of
killed whales tied to the board of a whale boat towing them to the base.
Often the killer whales almost wholly consumed the tongues of whale
carcasses stocked as feed for the whale base. Attempts to drive away the
killer whales by throwing various objects at them from the whale boat or
even by opening fire were almost of no avail. The tongue of whales weighs
two to three tons and contains a large amount of fat; thus killer whales
quite often cause serious losses to whale hunters by snatching away the
valuable tongues. Yet there are no reports of killer whales attempting
to tear off chunks of blubber from a whale carcass or to snatch its fins
(V.A. Arsen’ev). Apparently, there is no justification for ascribing the
various damages caused to dead whales to the activity of killer whales.
The killer whale is generally described as an unusually greedy car-
nivore and fantastic figures are cited about the quantum of food found
in its stomach. The stomach of a 6.5-m long killer whale purportedly
contained “five or six seals and semidigested remains of another six or
seven seals and thirteen harbor porpoises; in addition, there were two
nearly whole seals in the gullet” (Tomilin, 1957). If the average weight
of these animals is taken as even 30 kg, the stomach contents of this
killer whale should have weighed at least 600 kg, while the contents of

Fig. 294. Killer whales tearing out the tongues of dead whales (photograph by
V.A. Zemskii).

515

515

691

а closely packed stomach of a 20-m long fin whale would weigh about
1,000 kilograms. The killer whale is no doubt a powerful and dangerous
carnivore but its food habits have yet to be properly studied.

Behavior. Killer whales usually live in groups of a few to 100 or
more animals. They are encountered in the open sea as well as close
to coasts. Of the 567 killer whales caught in the waters of Japan, 391
(69%) were killed within the 30-mile zone, 101 (17.8%) at a distance
of 31 to 60 miles, and 36 (6.3%) beyond 60 miles from the coasts. The
distance at which the remaining 39 killer whales were killed could not be
ascertained. However, the spread of the party hunting for killer whales
(along with other species of cetaceans) is also of some importance in
such cases (Nishiwaki and Handa, 1958).

Some instances are known of the transgression of killer whales into
a river at a distance of a few tens of kilometers from the estuary in
which the animals were confined to a freshwater zone, sometimes for a
few weeks.

While feeding, killer whales move at 5-7 miles per hour and during
migrations at 15-17 miles per hour. It has been noticed quite often that
large groups of killer whales move in a broad front or even in columns
(Fig. 295). They usually spend 2 to 5-6 min under the water and perform
five to six brief spells of exhalation/inhalation in the interval between

Fig. 295. Killer whales at sea. Pacific Ocean, 1948 (photograph by М.М. Sleptsov).

516

692

dives. Sometimes they remain on the water surface for quite a long time,
with their high dorsal fins visible above the sea level throughout this
period. The blow produced by the killer whale is rather low, not more
than 2 m high.

The strong affinity of adult animals for each other and their calves
is a characteristic feature. It is difficult to break up a herd of killer
whales and quite often many animals of a given group can only be
caught on just one side of the ship. The animals are more active in warm
water (20- 25°C) and become sluggish as the water temperature drops to
10- 15°C. It is easy to catch the killer whale at this time and hence most
of those caught in the waters of Japan come from the northern part of
the country. This fact notwithstanding, killer whales do transgress into
polar waters and do not shirk from ice, although none have been sighted
among compact ice.

Migrations. The courses and periods of seasonal migrations of killer
whales have not been studied. They are sighted over much of the range
throughout the year but in the polar sections of the Northern and
Southern hemispheres apparently only in the summer months. The killer
whales seen on the coasts of the Chukchi Peninsula in June abandon
these waters by November-December (Nikulin, 1946). They inhabit
Hudson Bay, Fox Basin, and Davis Strait too only in the summer period.
Killer whales, like other species of cetaceans, are encountered in the
waters of Antarctica only in the summer months and abandon them
altogether in winter. Killer whales may continue to remain in the winter
months even at high latitudes only at such places in the oceans which
fall under the influence of powerful warm-water currents (for example,
the Gulf Stream).

In the waters of Japan, the bulk of the killer whales are caught from
April through November inclusive while the catch is insignificant from
December through April. More killer whales are caught on the coasts of
Hokkaido (northern coast, Sea of Okhotsk; southeastern coast, Pacific
Ocean), fewer on the northeastern coast of Honshu (38- 40° N lat.), and
even fewer on the southeastern coast of this island (33 -35° М lat.). In the
Sea of Japan, there is virtually no hunting of the killer whale (Nishiwaki
and Handa, 1958).

Reproduction. Mating of killer whales was observed in June and July
and births in spring and summer (Scheffer and Slipp, 1948) although the
view is prevalent that parturition occurs mainly in autumn. Embryos 27
to 222 cm long were found in July- August among killer whales of the
Kuril Islands. At this time, simultaneous with gestating females, four lac-
tating mothers were found. They had probably calved in July (Ivanova,
1961с). A nearly mature embryo measured 208 cm in length while the

516

"693

Fig. 296. Killer whale on the deck of a whaling base (photograph by V.A. Zemskii).

largest was 274 cm long. The smallest of the measured calves were 236,
246, 251, and 274 cm. Such measurements are evidently characteristic of
newborn calves. It has been suggested that gestation among the killer
whales can extend for 16 months but the maximum number of births has
been recorded in May to July (an instance of mating was photographed
from an airplane on June 22, 1957) although mating and parturition can
occur in the other summer months as well. Additional documented infor-
mation is required to resolve the questions of the duration of gestation
and periods of mating and parturition.

The duration of lactation has not been established.

By the time it is one year of age, the killer whale calf has reached a
length of about 350 cm. The average length of 320 males caught in the
waters of Japan was 6.4 m, with the largest 9.45 m long. The average
length of 247 females was 6.1 m, with the maximum length at 8.23 m.
Most of the males caught varied in length from 5.5-7.6 m and females
5.5-6.7 m. Of the 567 killer whales caught from 1948 through 1957, 320
were males (56.4%) and 247 females (43.6%). The sex ratio is evidently
close to 1:1 (Nishiwaki and Handa, 1958).

Enemies, diseases, parasites, mortality, and competitors. The killer
whale has no enemies. The diseases known in them are bone tumors

517

694

(detected in skeletons preserved in museums) and dental caries. Ectopar-
asites are not known. Eight species of endoparasites have been reported:
two species of trematodes, one species of castode, three species of nema-
todes, and two species of acanthocephalans.

The trematode Fasciola skrjabini Delamure was found in the bile
ducts of the liver of the killer whale and the Minke whale in the North
Atlantic. The intestine of a killer whale in the fore-Kuril waters revealed
the presence of the trematode Leucasiella subtilla A. Skrjabin. The ces-
tode Trigonocotyle spasskyi Gubanov was found in the small intestine of
whales only from the Sea of Okhotsk (Kuril waters). The widely preva-
lent nematode Anisakis (Anisakis) simplex Rudolphi, parasitizing the gul-
let, stomach, and intestine, was detected in 10 other species of toothed
whales, two species of baleen whales, and in Steller’s sea lion in addition
to the killer whale. The nematode Ана pacificus A. Skrjabin was
found in the stomach of killer whales (and also sperm whales and fin
whales) in the waters of the Kuril Islands. Another species of nematode,
Anisakis sp., found in the killer whale, has eluded precise identifica-
tion. The acanthocephalan Bolbosoma physeteris Gubanov, parasitizing
the intestine was found in killer whales from the Sea of Okhotsk and Bol-
bosoma nipponicum Yamaguti from the Kula Gulf [?] (Margolis, 1954;
Delamure, 1955; A. Skrjabin, 1958, 1959, 1960).

Field characteristics. A very high dorsal fin, rising to a height of over
1 m in males, distinguishes the killer whale from all other cetaceans. At a
close distance, bright white oval patches can be seen on the temples and
a large white patch posterior to the dorsal fin. Killer whales are usually
confined to groups. Quite often, the groups present a broad front or
move in a column. (V.A.)

Economic Importance

In spite of the comparative abundance of killer whales in the World
Ocean, their hunting has acquired no economic importance. Killer whales
are mostly caught casually during hunts for other cetaceans, such as
beaked whales, Minke whales, and various other species of dolphins. A
fairly organized hunting of killer whales exists only in Norway and Japan
with whaleboats specially designed for small whales (Minke whale, bot-
tlenose whale and other beaked whales, pilot whale, and killer whale).
Data on killer whale hunting for several years are given in Table 40.
As mentioned, Norwegian and Japanese hunting is carried out in
specially designed craft equipped with small-bore harpoon guns. In the
Soviet Union, the killer whale is hunted from the large whaleboats used
in hunts for large whales in the Far Eastern waters as well as in the

518

695
Table 40. Magnitude of killer whale hunting (international whaling statistics)
Year Norway Japan USSR Year Norway Japan USSR

1948 27 27 3 1959 69 36 36

1949 34 43 28 1960 82 48 45
1950 12 18 24 1961 111 54 4
1951 24 67 23 1962 124 47 7
1952 13 54 19 1963 90 43 4
1953 9 65 25 1964 77 99

1954 13 109 4 1965 104 169

1955 26 85 15 1966 161 137 9
1956 40 38 67 1967 36 101 4
1957 48 78 30 1968 86 22

1958 39 73 25

Antarctic. In other countries, some stray killer whales are caught inci-
dentally.

The fat of the killer whale, like that of other toothed whales, is used
for commercial purposes. The skin of the killer whale is regarded as
superior to that of other toothed whales and can be used for producing
leather goods. The rest of the carcass is used in the preparation of feed
or in fertilizer meal. (V.A.)

Genus of Risso’s Dolphins

Genus Grampus Gray, 1828

1828. Grampus. Gray. Spicilegia. Zoologica, I, p. 2. Delphinus griseus G.
Cuvier, 1812.
1933. Grampidelphis. Iredale and Troughton. Rec. Austral. Mus., 19,
р. 31. Substituted for Grampus Gray, 1846.!! (V.H.)
Dolphins of medium proportions, with a body length up to 430 cm.
The head is rounded anteriorly and no “beak” is perceptible. The
high dorsal fin, located almost midbody, is notched along the posterior
margin. The flippers are long and narrow.
The body color is basically gray.
The rostrum is broad; its length is shorter than the cranial section
of the skull. The proximal portions of the broad premaxillae form a
prominence. The pterygoid bones are adjacent. The nasal and frontal
bones form a crest posterior to the nares. The mandibular symphysis is
short. In most of these animals, teeth are present only in the anterior
portion of the lower jaw in two to seven pairs (0-2 in the upper jaw).

И See footnote on р. 680.

519

696

Vertebrae 68 - 69. Phalangeal formulae: [5, Пу - ;9, Шов, [Уз - $, and
У..

Very little is known about the biology of these animals. They feed
on cephalopods and live singly or in small groups.

They inhabit the warm and moderate waters of both hemispheres
but are few in numbers everywhere. They are not caught in the waters
of the USSR but might possibly occur in the waters of the Far East.

The genus comprises a single species: Risso’s dolphin, Grampus
griseus G. Cuvier, 1812. (V.S.)

RISSO’S DOLPHIN
Grampus griseus G. Cuvier, 1812

1812. Delphinus griseus. G. Cuvier. Ann. Mus. Hist. Nat., Paris, 19, p. 13.
Brest, France.

1822. Delphinus rissoanus. Desmarest. Mammalogie, p. 519. Mediter-
ranean Sea, Nice.

1866. Globicephalus chinensis. Gray. Cat. Seals and Whales Brit. Mus.,
р. 323. China Sea.

1873. Grampus stearnsii. Dall. Proc. Calif. Ac. $с., 5, р. 13. Monterey,
California. (V.H.)

Diagnosis
Only species of the genus.
Description

Body compact. The highly developed adipose body [melon] imparts a
circular shape to the head while the frontal portion slightly projects
anteriorly. The lower jaw is shorter than the upper. The oral cavity runs
obliquely upward from front to back. The flippers are long, constituting
13.9 to 19.6% of the body length, narrow, crescent-shaped, and pointed
at the ends (Fig. 297).

The body color varies from gray to blackish-gray, gradually turning
lighter from the dorsum toward the belly. The head is also lighter in
color than the dorsum but all the fins are the same color as the dorsum.
Small light-colored spots and bands on the skin are the result of various
damages. Hairs (up to eight) are sometimes preserved on the head of
young animals.

The diameter of the teeth of adult dolphins can reach 1.5 cm and
the height almost 4 cm (root and crown). The loss of teeth in the upper

519

697

ЕЕ,

Е
ке Ty, Lie

И, ИДЯ
Wi Wilf Mt Wife
йе /
И И Hl И, И fil yy Lt Whee
oy Ц AY ma ge ИИ И и ‘A у (er
STEM К и. Ra \ Ч ча He: р ips ii H f
Boas" ve re ‘‹ ЕН Tiss

Fig. 297. Risso’s dolphin, Grampus griseus (figure by N.N. Kondakov).

jaw is a recent phylogenetic phenomenon and apparently the result of
these animals feeding on cephalopods (Tomilin, 1957).

Cervical vertebrae 7, thoracic 12-13, lumbar 18-19, and caudal
30-31. The sections of the vertebral column constitute the following
percentage ratio to its total length: cervical 2.5, thoracic 24.5, lumbar 29,
and caudal 44 (Slijper, 1936).

The basic body measurements of three adult female Risso’s dolphins
(Tomilin, 1957; Pilleri, 1969) are respectively (in cm): body length 330,
320, and 275; distance from tip of snout to blowhole 43 and 48, up to
dorsal fin 127, 119, and 95, up to base of flippers 56 and 64; length of
flippers 54, 61, and 45; maximum width of flippers 22 and 17; height of
dorsal fin 34, 41, and 40; length of base of dorsal fin 56 and 35; and
width of caudal flukes 74 and 79.

The average skull measurements based on nine skulls (Fig. 298) of
adults (Tomilin, 1957) are (in cm): condylobasal length 50, length of ros-
trum 25, width of rostrum at base 20, and length of lower jaw 39. (V.S.)

Geographic Distribution
Warm and temperate waters of the Northern and Southern hemispheres.

Geographic Range in the USSR (Fig. 299)

Coastal waters of the Kuril range, mainly its southern part (Iturup and
Shpanberg Islands)!?, and the Commander Islands.

2 Finds reported for the Kuril Islands (Sleptsov, 1952) have yet to be confirmed with
specimens.

520

519

698
Geographic Range outside the USSR (Fig. 300)

Gulf of Maine and the coastal waters of New Jersey state along the
American coast of the North Atlantic (the range may be more extensive
but precise data are not available). In the eastern part of the Atlantic
Ocean, the range extends from the North Sea (waters of Great Britain,
Schleswig in the Federal Republic of Germany, Denmark, and western
coast of southern Sweden in Bohus Bay) and Ireland down to South
Africa. Mediterranean Sea in the east including the Adriatic. In the east-
ern part of the Pacific Ocean, from British Columbia to Baja California
(Mexico) and waters of the Chilean coast; in the western part of this
ocean from the Commander Islands (?) and Kuril Islands and Japan to
the seas of China. Waters of New Zealand and Australia (New South
Wales), the Indian Ocean, and the Red Sea. (V.A.)

Geographic Variation

Not known.

$
&
к
\

NAY
tiga git
.%

^

(и of

all

ie [О }
Ry)

Fig. 298. Skull of Risso’s dolphin, Статриб griseus (figure by М.М. Kondakov).

522

520

699

Biology

In all probability, this species is not very numerous throughout its range.
Some species of cephalopods constitute its main food. Remnants of other
food items have not been detected in the stomach of Risso’s dolphins.
In search of cephalopods, this dolphin makes comparatively long dives
and remains on the water surface longer than other species. It mainly
inhabits the open seas. In most cases, it swims singly, in pairs or in small
groups. These dolphins gather in herds of a few tens of animals only
occasionally at places of food concentration.

Fig. 299. Range of Risso’s dolphin, Grampus griseus, in the USSR (V.A. Arsen’ev).

‘(Ad UasIy УЛ) 5725148 зпашиоло ‘ичАрюр $,08sty jo э8иел $э10э4$ ‘00$ ‘314 155

SSS SS
Wy 0005 000% 0005 0005 0001 0 0001

701

Migrations have not been established. It is known that these dol-
phins live year-round in the Mediterranean Sea and in European waters,
although some variations in their numerical strength have been recorded
in different months.

According to some fragmentary information, parturition -occurs in
the winter months in comparatively warm waters. The newborn calf aver-
age 150 cm in length.

Three species of endoparasites are known. The cestode Phylloboth-
rium delphini Bosc localizes in the skin of Risso’s dolphin and also six
other species of toothed cetaceans, the bowhead whale, and Weddell’s
seal. It has been detected in these animals at many points in the Atlantic
Ocean, the Mediterranean Sea, waters of the Commander Islands and
Australia, the Pacific Ocean, and the Antarctic. The nematode Crassi-
cauda grampicola Johnston and Mawson was detected only among Risso’s
dolphins in the waters of Australia. Stenurus minor Kuhn parasitizes the
bronchi, tympanic cavity, and blood vessels (Delamure, 1955).

Risso’s dolphin is of no economic importance. (V.A.)

Genus of Pilot Whales
Genus Globicephala Lesson, 1828

1828. Globicephala. Lesson. Nat. Hist. Mamm. Diseaux Dépuis 1788,
Cétacées, p. 441. Delphinus deductor Scoresby = Delphinus melas
Traill.

1828. Globicephalus. Lesson. Férussac Bull. Sci. Nat., 16, p. 116. Substi-
tuted for Globicephala Lesson, 1828.

1884. Globiceps. Flower. Proc. Zool. Soc. London, 1883, p. 508. Substi-
tuted for Globicephala Lesson, 1828. Nom. ргаеосс. (V.H.)

Large dolphins, with a body length of up to 6.5 m.

The head is rounded and a beak faintly perceptible. The dorsal fin
is low with a curved posterior margin. The flippers are narrow and long.

The body is dark, grayish-blue, or black.

The skull is broad and slightly flattened dorsoventrally. The rostrum
is almost as long as the cranium. The premaxillae are very broad. The
pterygoid bones are adjacent. The nasal bones projects into the frontal
and lie above the level of all the other bones. Teeth 7-1, present in the
anterior half of the jaws. With advancing age, the teeth may wear out
and fall out. Vertebrae 58-59. The sternum comprises three to four sec-
tions. Five of the 11 pairs of ribs articulate with the sternum. Phalangeal
formula: 5-4, Ig-44, HI9-1,, [М ->.

These whales feed on large invertebrates (mainly cephalopods)
and schooling fish (teutho-ichthyophagous). Periods of mating and

523

702

parturition are protracted. Gestation and lactation continue for about
a year each. Members of this genus are confined to large herds of a
hundred or more animals.

The range of these whales covers all the seas except the polar. In the
USSR, they might possibly occur in the Barents Sea and in the waters
of the Far East.

Fossil remains have been detected in the Pleistocene of North America.

The genus comprises a single species: the (common) pilot whale,
G. melaena Traill, 1809.

Sometimes pilot whales are classified into two species, G. melaena
and G. macrorhyncha Gray, 1846 (Rice and Scheffer, 1968*), or into three
species, i.e., С. scammoni Cope, 1869, in addition to the two aforesaid.
(V.S.)

PILOT WHALE
Globicephala melaena Traill, 1809

1809. Delphinus melas. Traill. Nicholson’s Journ. Philos., 22, р. 81.
Pomona Island, Orkney Islands.

1812. Delphinus globiceps. G. Cuvier. Ann. Mus. Hist. Nat., Paris, 19,
р. 14. France.

1820. Delphinus deductor. Scoresby. Account Arctic regions, I, p. 496.
North Atlantic.

1824. Delphinus grinda. Lungbye. Kongl. Danske Vedensk. Selsk. Afh. I,
р. XI. North Atlantic.

1846. Globicephala sieboldii. Gray. Zoology Voyage Erebus and Terror,
I, p. 142. Near Nagasaki, Japan.

1869. Globicephalus scammoni. Cope. Proc. Ac. Sc. Philadelphia, 21,
р. 21. California (10°N lIat.).

1871. Globicephalus sibo. Gray. Suppl. Cat. Seals and Whales Brit. Mus.,
р. 85. Japan.

1898. Globicephala melaene. Thomas. Zoologist, (4), 2, p. 99. Mascu-
line form melas for the feminine of the genus, conforming to the
generic name. (V.H.)

Diagnosis
Only species of the genus.
Description

The body is elongated and slightly thickset in the anterior half. The
head is rounded with a projecting frontal part as a result of the highly

524

523

703

developed adipose body [melon]. The oral cavity runs from front to back
with an inclination of roughly 30 - 45° to the longitudinal axis of the body
from the bottom upward. The upper jaw is longer than the lower one.
The blowhole is slightly displaced left of the midline. The long and low
dorsal fin is located almost at the boundary of the anterior third of the
trunk. The length of the fin at the base is not less than 1.5-2 times its
height. Long flippers reach one-fourth of the body length (Fig. 301).

The upper portion of the body is darker in color than the under-
side. A narrow gray band enlarges into a patch between the flippers
midabdomen and on the throat. A newborn pilot whale is gray but soon
turns black. With age, high longitudinal ventral and dorsal keels form
on the caudal stem of males. The adipose body [melon] of the head and
the hump between the head and dorsal fin are larger in males than in
females. Embryos sport three to six hairs on each side of the snout.

Cervical vertebrae 7, thoracic 11, lumbar 12-14, and caudal 28-29.
The first five or six cervical vertebrae are fused.

The main body measurements of an adult male pilot whale caught off
the coast of Virginia (North Atlantic) (Tomilin, 1957) are (in cm): body
length 465; distance from tip of snout to blowhole 54, up to anal opening
318, and up to base of flippers 91; length of flippers 76, maximum width
of flippers 25; length of base of dorsal fin 79; height of dorsal fin 35; and
width of caudal flukes 117. The largest male measured 6.5 m in length
and the female 6.1 m.

The main measurements of six skulls (Fig. 302) of adult pilot whales
averaged (Tomilin, 1957) (in cm): condylobasal length 63 (six measure-
ments), zygomatic width 41 (one), length of rostrum 32 (six), width of
rostrum at base 23 (six), and length of lower jaw 49 (two).

Fig. 301. Pilot whale, Globicephala melaena (figure by N.N. Kondakov).

524

704

Fig. 302. Skull of pilot whale, Globicephala melaena (figure by М.М. Kondakov).

The relative dimensions of the rostrum, the width of the skull at the
level of the orbits, and the width of the rostrum at its base and center
increase with age (Tomilin, 1957). (V.S.)

Geographic Distribution

Almost all the seas and oceans from the Arctic to the Antarctic except
the frozen seas of high latitudes.

Geographic Range in the USSR (Fig. 303)

Southwestern part of the Barents Sea, especially the waters of
Rybachii peninsula (irregular transgressions; Nemirovich-Danchenko,
1877; Chapskii, 1941), waters of the Kuril range, and possibly
southeastern Kamchatka, Commander Islands, and the Sea of Japan (La
Perouse Strait).

Geographic Range outside the USSR

In the North Atlantic it is found along the American coast from Delaware
Bay to Newfoundland, Labrador, Davis Strait (up to 70°N lat.) and the
southeastern coast of Greenland (67°N lat.). In the eastern part it occurs

705

‘(лэ.цэзгу У) USSN Э41 Ul 9720]2\ vjoydasigopy ‘ayeyM 1014 до э8иеч ‘50$ ‘8

97S

525

706

from Iceland and northwestern Norway along the entire coast south up
to Madeira,!? Mediterranean Sea. In the south, evidently inhabits waters
up to 60°S lat. in the Atlantic (Fig. 304).

In the North Pacific Ocean, up to the waters of southeastern China
to the west and south; in the eastern part of the ocean from the Aleutian
Islands and Alaskan peninsula down to Mexico (Baja California). In the
tropical zone, the pilot whale is probably distributed over large expanses
far away from the coasts.

In the Southern hemisphere, it is known on the coasts of India (Bay
of Bengal), Java, Guatemala, Ecuador, Peru, Chile, Australia, Tasmania,
New Zealand, Kerguelen Island, and South Africa. The range apparently
reaches 50 to 60°S lat. (Sergeant and Fisher, 1957; Tomilin, 1957; Brown,
1961; Hershkovitz, 1966; Nishiwaki, 1967). (V.A.)

Geographic Variation

Some authors recognize up to five or six subspecies but their validity
(even of the two subspecies mentioned below, the most sharply mani-
fest) is usually questioned or contradicted (Hershkovitz, 1966). Distant
migrations characteristic of the species result in a thorough mix-up over
the entire extremely extensive but essentially single range. Deviations
in the picture of distribution of the various forms, especially of those
given below, are highly significant (see Tomilin, 1957, 1962; Hershkovitz,
1966).

Two subspecies can be recognized in the waters of the Soviet Union:
1. Common, or Atlantic pilot whale, С. т. melaena Traill, 1809 (syns.
melas, globiceps, deductor).

Thirteen pairs of teeth in each jaw; the white band on the ventral
side of the body is broadened in the region of the flippers, and a large
white, anchor-shaped patch occurs on the throat.

Southeastern part of the Barents Sea, especially waters of the
Rybachii peninsula.

13 References to habitation in the Baltic Sea are very general and neither positive nor
reliable. They are mainly communications from one author to another and it is very difficult
to identify the actual sources. More positive references are available only for the waters of
Denmark (Jutland peninsula) and southwestern Sweden, i.e., the same section of the coast
facing Jutland. Such references usually extend to the entire Baltic Sea. For the waters of the
German Democratic Republic and Poland, there are no direct references to the habitation
of the pilot whale and this species is not included among the cetaceans inhabiting these
waters (Gentshel’, 1937; van den Brink, 1958; Koval’skii, 1964; Siivonen, 1968). Transgres-
sions into the basin of the Baltic Sea proper might occur very rarely but the appearance of
this animal in our waters far into the northeast is highly improbable. (V.H.)

707

“(лэ.чэзгу У’) Duanjau п]244э2190]0 ‘этеЧчм 1014 ay) Jo э8иел $э0э4$ “pog “314

528

708

Outside the USSR, it is encountered in the North Atlantic Ocean
up to southern Greenland (almost 70°N lat.).!4
2. Black, or Pacific pilot whale, С. m. sieboldii Gray, 1846 (syns. scam-
moni, sibo). Seven or eight pairs of teeth in each jaw. Color black, with
no white markings.

Waters of Kuril Islands (not known more accurately), probably Kam-
chatka, Commander Islands, and the Sea of Japan (La Perouse Strait).

Outside the USSR, this subspecies is encountered in the North
Pacific Ocean up to the Yangtze estuary in the west and from the Aleu-
tian Islands and Alaska to Guatemala in the east.

Pilot whales belonging to two or three other subspecies of this
species are sometimes listed as inhabitants of the Southern hemisphere
(Tomilin, 1962; Nishiwaki, 1966). (V.A.)

Biology

Population. The pilot whale can be reckoned as a relatively abundant
species. In some regions of the range (North Atlantic, waters of Japan,
and South Pacific Ocean), this dolphin often forms herds a thousand or
more strong. It is difficult to indicate the regions of maximum population
of the pilot whale.

Food. Squids serve as the main food. In the waters of Newfound-
land, the pilot whale consumes the squid Illex illicebrosus almost exclu-
sively. This squid is remarkably abundant here. In the summer months,
pilot whales approach the island coasts chasing after it. In the absence
of squids, pilot whales consume cod (in small quantities). In a Florida
oceanarium, a young pilot whale initially refused to eat fish but after
sometime was taught to accept it. In the waters in which I. illicebrosus
is not available, the stomach of pilot whales contained: the remnants
of other species of squids, e.g., Ommatostrephes (Todarodes) sagittatus,
and some species of the family Onychoteuthidae. Otoliths of mackerels
(Caranx trachurus), small flounders (Pleuronectidae), and herrings have
been found from time to time.

Young pilot whales feed on much smaller squids than the adults do.
The nearly identical degree of digestion of the food in the stomach of
pilot whales caught at the same time suggests that the entire herd feeds
together. By converting the quantum of the remnants of squids and fishes
into the average weight of intake, it was established that the stomach of
a pilot whale of medium proportions (length 396 cm) held 12-14 kg

14 The following distribution has been given for this form: “Atlantic, Pacific, and Indian
oceans from 70°N lat. to roughly 60°S lat.” (Hershkovitz, 1966).

709

of food. The stomach of a female 247 cm long contained 155 squids
weighing in toto 27 kg. In an oceanarium, a male pilot whale 525 cm in
length daily consumed an average of 45 kg of squids and mackerel, while a
female 396 cm long consumed up to 36 kg; another female (366 cm long)
ate 18 kg three times a day. Apparently, under natural conditions, the
pilot whale may fill its stomach two or three times a day. Such voracious
feeding is only resorted to, however, during the three or four summer
months when immense concentrations of squids are available. During
the rest of the year the daily ration of the pilot whale is quite modest.

At a daily consumption of 27 kg, the annual food requirements are
about 10 tons. Taking the weight of a pilot whale of medium proportions
(396 cm long) as 830 kg, its annual food intake represents 11.5 times the
weight of the animal itself. The daily quantum of food consumed by the
dolphin constitutes 3 to 5% of the animal’s body weight.

Observations on three pilot whales in a basin showed that they fed
only at night and rested during the day. Under natural conditions in
Newfoundland, pilot whales were observed feeding during the day also.
The stomach ofa pilot whale caught between 7:00 and 11:00 a.m. revealed
food recently consumed. It is probable that the long summer day in these
latitudes causes the squids to rise to the upper horizon not only at night,
but also during the day, which enables successful catches by the pilot
whales (Sergeant and Fisher, 1957; Tomilin, 1957).

Behavior. Pilot whales usually live in herds of some 20 animals, on
average, but sometimes do form herds of up to a thousand or even several
thousands of animals. They dive for comparatively longer periods and
surface sometimes 8-10 times in a row, producing low (up to 1.5 m
high) bushy blows. They never display the caudal flukes and never leap
free of the water surface. Large herds of pilot whales are formed from
individual groups of 15-25 animals each.

The composition of the herds varies. Thus a herd of 16 pilot whales
comprised 7 males and 8 females (the sex of one whale could not be
ascertained). Of the seven males, three were mature, three immature,
and one a newborn calf. The eight females comprised six mature, one on
the threshold of maturity, and one newborn. This was a mixed herd. The
predominance of mature females over mature males suggests polygamy.
The ratio between the adult males (472 cm and above) and adult females
(336 cm and above) was close to 1:3. |

Another herd studied consisted of 14 pilot whales which comprised
10 males and 4 females. The length of nine males was typical of mature
animals and all the four females were large, presumably “old”. Such a
herd could be called a “bachelor” group, made up in all probability of

S29

710

animals which did not take part in reproduction in that particular year
(Sergeant, 1962b).

Pilot whales often form mixed herds together with some species of
very small dolphins. In such herds, apart from the distinct presence of
the shorthead dolphin [Lagenorhynchus electra] and the bottlenose dol-
phin, other species of dolphin may also be present. When such a herd
approaches a ship standing in the sea, all the small dolphins disappear
into the water after some time while the pilot whales invariably continue
to remain close to the ship (Brown, 1961).

Sometimes some animals surface vertically from the water almost up
to the flippers, remain in that position for about 0.5 min, then submerge
without altering their upright posture. Some animals strike the water
with their fins (Brown, 1961). This ability to assume a vertical position
is taken advantage of in oceanariums. The pilot whale is gradually trained
to rise higher and higher out of the water to reach food dangled above
it. A large dolphin, such as the pilot whale, can thus be trained to leap
clear of the water and even to “Пу” above it (Fig. 305).

The extraordinary behavior of a herd of pilot whales once came to
notice. This mixed herd consisted of about 150 pilot whales and small
dolphins. Suddenly, some animals assumed a vertical position but with
their head down (under the water) and each struck the water surface
five or six times with its caudal flukes. They then assumed a horizontal
position and rejoined the herd. Such a head-down position has been
reported among pilot whales in other instances too (Brown, 1960).

Pilot whales are beached more often than other cetaceans, both indi-
vidually as well as in groups. On October 7, 1948, as many as 46 pilot
whales were cast ashore simultaneously on a beach in Florida; those
found early in the morning, during low tide, were still alive. Four were
transported by truck while the others perished on the coast in high tide
(Obruchev, 1951). Beached groups usually consist of animals of various
age but sometimes young ones exclusively.

A pilot whale in a Florida aquarium slept only at night in the first
few months of captivity but later continued to sleep during the day also.
While asleep, all of its body was submerged and only the dorsal fin and
blowhole remained exposed above the water.

Pilot whales hear well under water and their aeriai vision is excel-
lent. In captivity, they invariably perceive the approach of the food atten-
dant and swim immediately to the feeding place. One female pilot whale
quickly learned to snatch food from the attendant’s hand; it swam to the
feeding site, stood vertical with its head above the water, and opened its
mouth for the attendant to throw cephalopods inside. Washing of the

529

530

711

SAN See

a ‘Wat ills
\ soot es

ie 3 И tS
nl Ay ae
vl ‘ath , eS

г. т Tae

Fig. 305. Leaping pilot whale (figure by N.N. Kondakov).

pail signalled completion of feeding and the pilot whale would immedi-
ately swim away. A male loved to be combed with a brush and quickly
approached anyone with a brush in his hand (Tomilin, 1957).

A pilot whale (body length 366 cm) caught on February 27, 1957
was housed in a tank 30.5 m long, 15 m wide, and 6.7 m deep. For the
first few days the animal refused food and the attendant had to enter
the tank in a diver’s suit, open her mouth with metal tongs, and push
squids inside which the whale gulped rapidly. After 10 days she began to
swim to the wooden platform affixed to the side of the tank and to take

Ve

food from the tongs held by the attendant. The animal consumed more
- than 20 kg of squids per day. A large fish and two turtles were also kept
in the same tank. During the feeding period sometimes the fish would
snatch the squids from the tongs before the pilot whale and sometimes
even snatched them from the whale’s open mouth. This angered the
pilot whale. She would stop feeding and chase the fish around the tank.
The turtles also swam to the feeding platform, which irritated the whale.
Once she threw herself on them with such vehemence that one suffered a
broken shell. However, the pilot whale soon learned to avoid competition
during feeding: she would stand vertically at the feeding platform with
a third of her trunk projecting above the water, maintaining balance by
slight movements of the caudal ‘flukes, and thus receive the food high
above the water surface.

This offered a basis for teaching the pilot whale various tricks. First,
it was taught to leap clear of the water to reach food dangled high above
its head and next to associate feeding with a whistle. This was followed
by a “hand shake”. As soon as the pilot whale appeared around the feed-
ing platform, the trainer would clasp it around the flippers and whistle
before feeding. Very soon the pilot whale began to anticipate the actions
of the trainer. On hearing the whistle, it would turn on its side, “extend”
the flippers to the trainer, then demand food. By the same method, it
was taught to fetch a plastic dumbbell from inside the tank and eventu-
ally to “sing”. At the command of the trainer, it would quiver its jaws
and simultaneously blow air powerfully through the blowhole. The pilot
whale learned these various tricks quite rapidly. It was not at all afraid
of any unfamiliar object. When in the course of the experiments a large
boat was dropped inside the tank, the whale exhibited no apprehension.
To the contrary, after a few minutes it swam to the boat, fed tranquilly,
and performed its tricks. |

In the first week of February, 1958, the pilot whale lost its appetite.
Next, it began to rub the genital slit against a stone or other projecting
object at the bottom of the basin, including the diver’s abandoned helmet.
This behavior continued for nearly two weeks, after which the animal
resumed normal feeding. Perhaps this was the period of its heat.

For a long time the pilot whale was friendly with the divers who
entered the tank to feed the other animals, even attempting sometimes
to snatch the food from their hands. But after nearly 14 months of living
in the tank, it began to push the diver (or anyone entering the tank) with
its head and to bite his hands while he was feeding the other animals.
One diver barely escaped serious injury when, on the water surface, the
pilot whale tried to pin him to the wall of the tank. Later, it chased three
persons simultaneously, two of whom were photographers. It hit the leg

531

713

of one and zoomed to the second loudly clicking its teeth; all three made
a hasty retreat. It then attacked the large photographic camera set up
on a tripod at the bottom of the tank, hitting it with such force that the
heavy lens was badly damaged. Four days later the pilot whale struck one
of the divers so powerfully that he lost consciousness and barely escaped
drowning. This change in the behavior of the pilot whale was attributed
to its long lonely existence. In July, 1958, it was transferred to a basin
with another small pilot whale and two blue-white dolphins. Initially
the dolphins were frightened and shied away from the large pilot whale,
but soon became emboldened and spent several hours nipping it behind
ihe flippers whenever possible. This irritated the pilot whale who chased
them but the more agile dolphins easily escaped. After about a month the
animals became friends, swimming abreast, and often playing together.
From the time the pilot whale began living with the other dolphins, its
animosity against the divers ceased (Brown, 1960).

Migrations. Pilot whales probably perform seasonal migrations since
they are more abundant in the North Atlantic Ocean (as in the Pacific
Ocean also) in summer than in winter. Norwegian whalers hunting for
small whales in the northwestern Atlantic catch the pilot whales mostly
in the second half of summer. Thus in 1958, of the 216 pilot whales

caught, 64 were bagged in July and 150 in August (Ostby, 1959). Table 41

provides an idea of the duration of residence of pilot whales in the waters
of Newfoundland.

In the first half of January, 1953, a large herd of pilot whales was
sighted at 42°30’ М lat. and 52° W long.; evidently the effect of the Gulf
Stream was more intense in the winter.

In the summer months, pilot whales Е visit the large deep-
water bays of Newfoundland and, concomitantly, though in small num-
bers, are found in the central part of the Labrador Sea. Similarly, pilot
whales are regularly sighted in summer in the eastern waters of the North
Atlantic, initially off the Faroe Islands and the southwestern coast of
Norway, later off northwestern Norway, Iceland, and Jan Mayen Island,
and at the end of summer, in the western part of the Barents Sea, in the
waters of Medvezhii Island, and even at Spitsbergen.

Table 41. Observations on pilot whales in Newfoundland (Sergeant and Fisher, 1957)

Year First observation Subsequent observation
1953 July 8 November 4
1954 July 6 November 10

1955 July 15 October 22

532

714

The pilot whale is purportedly encountered in winter east of the
Grand Bank south of Newfoundland in the waters of the North Atlantic
current. Information on the migrations of the pilot whale in other regions
of its range is so scant that no clear picture of migrations can be drawn.
Apparently migrations are not very distinctly manifest since these ani-
mals are sighted in the northern as well as the southern part of the range
throughout the year.

From 1926 through 1953 (July to October end), 20 instances of
beached pilot whales were recorded on the coast of western Green-
land, most of them in September. In the winter of 1931/32, a herd of
pilot whales wintered on the coasts of western Greenland (Sergeant and
Fisher, 1957).

Reproduction, growth, and development. The periods of mating and
parturition in the pilot whale evidently last for about six months but
most of the animals mate over a brief duration. The maximum number of
matings presumably occurs in the spring months with parturition peaking
in mid-August, although there have been instances of calves born from
May through November. Stray cases of births have been recorded in
almost all the months of the year. Gestation apparently continues for
15-15.5 months. As a rule, a single calf is born and twins are rare.
There is a solitary record of a female with three embryos. The average
length of male calves at birth is 178 cm, of female calves 174 cm. The
teeth begin to cut at a body length of about 213 cm. This process ceases
when the calf attains a length of about 274 cm. The growth of embryos
and newborn pilot whales is shown in Table 42.

The calves begin independent feeding on squids at an average body
length of 230 cm at the age of six- nine months; however, suckling does
not cease and extends for about two years. Lactation of the female contin-
ues for 21-22 months. Instances of fresh impregnation during lactation
are rare; the females usually ovulate at the end of lactation. Thus the
full cycle of reproduction roughly covers 40 months. During the period
of reproductive capability, the female produces, on average, nine calves.

Male calves record a much greater growth tempo than female calves.
A young male living in a Florida aquarium added 30 cm to its length and
put on 45 to 90 kg in four months. The average length of adult animals
is usually about 396 cm. Females ovulate for the first time (attain sexual
maturity) at six years of age at an average body length of 356 cm and
males at the age of 12 years at an average body length of 490 cm. After
attaining sexual maturity, the growth of the animal slows down. Males,
which attain maturity later, are considerably larger than females. Most
of the males in the group measured, fell in the 565-cm group and the
maximum number of females in the 340-cm group. The maximum size

533

5

Table 42. Length and weight of embryos and calves of pilot whales (Sergeant, 1962b)

Embryos Calves
Length, cm Weight, kg Length, cm Weight, kg

16.5 0.08 193 113.4
19.0 0.12 195 953
21.0 0.18 205 145.6
23.0 0.22 208 130.6
24.0 0.25 231 188.7
25.0 0.32 236 202.8
26.0 0.36
31.0 0.54
36.0 0.75
41.0 1.16
43.0 1.45
48.0 1.54
59.0 3.74

114.0 21.00

119.0 28.50

165.0 60.00

of the males was 617 cm and of the females 511 ст. The maximum
recorded age of the males was roughly 40 years and of the females 50
years (Tomilin, 1957; Sergeant, 1959, 1962b).

Table 43 shows the composition of pilot whales caught in Newfound-
land according to their sizes: the animals have been grouped into a series
at 15-cm intervals. The total number of males measured was 1,275, of
females 1,951 and the number falling in each length group is given. The
data show that the size of adult males far exceeds that of adult females
and that animals of all ages fall victim to the hunter.

Table 44 gives data on the age composition of the oldest groups
of pilot whales caught and investigated. The age of the animals was
determined from the tooth layers. From a sample of 518 pilot whales,
males with a body length 549 cm and above (only 11 animals) and females
with a body length 457 cm and above (22 animals) were taken. It can be
seen from this Table that these animals were quite advanced in age, the
females reaching a maximum age of 50 years.

Enemies, diseases, parasites, mortality, and competitors. As in the case
of all other cetaceans, the killer whale is among the enemies of the pilot
whale.

Bone tumors were detected among the diseases suffered by the pilot
whale. The skin parasite Isocyamus delphini Guerin-Meneville was invari-
ably detected around the mouth and in old wounds on the body of the
Newfoundland pilot whale. Conchoderma auritum Linn. was detected on

532

533

716

Table 43. Size composition of pilot whales studied (Sergeant, 19625)
Average length Number of animals Average length Number of animals
of animals in in the group of animals in in the group
the group, cm LANG ERED OM SEROMA TONE) ole a the group, cm в hy se
Males Females Males Females
168 6 10 396 59 169
183 15 22 411 41 259
198 12 17 427 61 278
213 20 16 442 44 221
229 13 12 457 36 170
244 28 32 472 36 50
259 51 48 488 37 10
274 54 61 503 37 5
290 63 66 518 41 —
305 52 78 533 67 —
320 61 83 549 76 —
335 59 83 564 84 —
351 53 59 579 47 —
366 60 73 594 23 —

381 33 129 610 6 —

Table 44. Age of the largest pilot whales from a sample of 518 animals (Sergeant,
1962b)
Age, years Number of animals
Males Females
21-25 4 2
26-30 2 6
31-35 3 8
36-40 2 3
41-45 = 2
46-50 — 1
Total 11 22

the gums and teeth, and Cyanus globicipitis Lutken, Xenobalanus glo-
bicipitis Steenstrup, and Cyrolama globicipitis van Beneden on the body.
Eight species of endoparasites have been recorded: trematodes one,

‘cestodes three, nematodes three, and acanthocephalans one. The trema-

tode Campula gondo Yamaguti was found only in the bile ducts of the
pilot whale from the Pacific Ocean. The cestode Trigonocotyle lintoni
Guiart, parasitizing the intestine, is likewise known only in pilot whales
from the Atlantic Ocean and the Mediteranean Sea. Phyllobothrium del-
phini Bosc, widely distributed among marine mammals, parasitizes the
skin. In addition to the pilot whale, it has been detected in six other

534

И 17

species of toothed whales, in right whales, and in Weddell’s seal from
the Atlantic and Pacific oceans, the Mediterranean Sea, and Antarctic
waters. The cestode Monorygma grimaldi Monier is a parasite in the
abdominal cavity, mesentery, and diaphragm of pilot whales and three
other species of dolphins of the Atlantic Ocean and the Mediterranean
Sea. The nematode Anisakis (Anisakis) typica Diesing localizes in the
stomach of the pilot whale and three other species of dolphins; it has
been found in the North Sea and on the coasts of southeastern Africa.
The nematode Stenurus globicephale Baylis and Dauney, parasitizing in
the blowhole, bronchi, and the circulatory system, has been reported only
in the pilot whale from the North Atlantic. The nematode Torynurus con-
volutus Kuhn parasitizes the bronchi and blood vessels of the lungs of the
pilot whale and also common porpoises and one more dolphin (species
not established). It has been detected in the waters of Europe, the North
Atlantic, Sakhalin, and the Sea of Okhotsk. The only species of acan-
thocephalans, Bolbosoma capitatum Linstow, localizes in the intestine of
the pilot whale, sperm whales, and false killer whales from the Atlantic
Ocean and the Mediterranean Sea (Delamure, 1955).

Field characteristics. This species differs from other dolphins in hav-
ing a round head which projects high above the water, a dorsal fin with a
very broad base and its apex turned posteriorly, and slower movements.
(V.A.)

Economic Importance

The pilot whale is hunted in Newfoundland and the Faroe Islands by
Norwegian ships operating close to Norway as well as in the North and
Barents seas, in addition to other species of small cetaceans; it is also
hunted in the waters of Japan (Table 45).

A relatively large number of pilot whales is caught only in Newfound-
land and in the Faroe Islands; these animals are of considerable local
importance. In all other waters, their catch is too small to be significant.

Hunting is done by various methods. In Norway and Japan, in addi-
tion to other species of small whales (Minke whale, killer whale, beaked
dolphins, etc.), they are shot with small-bore harpoon guns from small
boats. In Newfoundland, most of the pilot whales are caught by chasing
large herds into shallow bays of islands and harpoons used to a lesser
extent.

The flesh of the animals caught is used as food (in Japan) and also
as feed for fur-bearing animals in farms (Newfoundland). Oil is obtained
from blubber. (V.A.)

718

Table 45. World catch of pilot whales (international whaling statistics)

Year Newfoundland Faroe Norway Japan
Islands

1950 172 10

1951 3,100 8

1952 3,155 2

1953 3,584 1

1954 2,298

1955 6,612 13 61

1956 9,799 1 279

1957 7,797 80 174

1958 789 216° 197

1959 1,725 1,422 224 144

1960 1,957 1,680 331 168

1961 6,262 1,892 295 133

1962 150 1,753 43 80

1963 221 2,194 71 228

1964 2,849 1,386 54 217

1965 1,520 1,599 32 288

1966 887 1,488 339 199

1967 739 1,979 117 237

1968 204 1,749 31 166

Genus of Pygmy Killer Whales
Genus Feresa Gray, 1870

1870. Feresa. Gray. Proc. Zool. Soc. London, p. 77. Orca intermedia Gray
= Feresa attenuata Gray. (V.H.)
Small dolphins, with a body length up to 244 cm.

The head is rounded and has no “beak”. The dorsal fin is quite high.

The main body color is dark gray.

The rostrum occupies nearly one-half of the skull length. The ante-
rior part of the premaxillae is flattened; their, inner edges are set off from
each other. The lower jaw has a keel in the zone of the symphysis. Teeth
оц. Vertebrae 68-71.

Biology has not been studied. These dolphins apparently feed on
squids and fish.

There is very little information on the sites where this rare dolphin
is found: in the North Pacific Ocean from Alaska to Guatemala in the
east, on the coasts of Japan, in the Yangtze estuary, and on the coasts
of China in the west.

This dolphin has not been reported in the USSR but it might occur
in the waters of Kuril Islands and in the Sea of Japan. (V.S.)

535

719

PYGMY KILLER WHALE
Feresa attenuata Gray, 1875

1827. Delphinus intermedius. Gray. Philos. Mag. or Annals, 2, p. 376.
Type locality not established. Nom. ргаеосс.

1875. Feresa attenuata. Gray. Journ. Mus. Godeffroy (Hamburg), 8,
p. 184.

1856. Feresa occuleta. Jones and Packard. Proc. Biol. Soc., Washington,
69, p. 167. Substituted for Delphinus intermedius Gray. (V.H.)

Only species of the genus.

Somewhat resembles the false killer whale. The body is elongated
and spindle-shaped. The head is relatively small. The highly developed
adipose pad [melon] projects anteriorly to impart a circular feature to
the head (Fig. 306). The flippers are crescent-shaped, quite broad at the
base and pointed at the tips. The height of the dorsal fin is slightly less
than its basal length. It is arcuately notched along the posterior margin.
The caudal flukes are relatively small.

The body is a monochromatic dark gray. The margins of the jaws
and the region around the anal opening are white. An anchor-shaped
light-colored patch is seen on the breast. Wavy pale-colored bands run
on the flanks (Nishiwaki, 1966).

The skull (Fig. 307) is highly shortened and measures about 16% of
the body length. The short rostrum, broad at the base, gradually, but not
very significantly, narrows toward the tip. The upper side of the rostrum
is Slightly concave (Yamada, 1954). The teeth are fairly large with crowns
about 1 cm high. Cervical vertebrae 7, thoracic 12-13, lumbar 16-17,
and caudal 32 - 34.

Fig. 306. Pygmy killer whale, Feresa attenuata (figure by N.N. Kondakov).

536

720

SY

NINES

Fig. 307. Skull of the Pygmy killer whale, Feresa attenuata (figure by N.N. Kondakov).

The body length of males (Japan; Nishiwaki, 1966) ranges from
214-244 cm, of females 208-227 cm. The maximum weight recorded
was 225 kg for a male measuring 244 cm in length. The body propor-
tions of males with a body length of 214-244 cm and of females with a
body length of 208-227 cm respectively (as percentage of body length)
are: from tip of snout to blowhole 6.8-11.9 and 7.2-11.1, to flippers
16.7 - 21.6 and 18.5 - 22.4; from notch between caudal flukes to anal open-
ing 34.2 -37.2 and 34.2 - 38.6; length of flippers 18.3 -22.1 and 14.7 - 22.2,
maximum width of flippers 6.1-6.7 and 5.8-7.0; length of base of dorsal
fin 12.6- 15.4 and 14.0- 16.6, height of dorsal fin 9.4- 11.6 and 9.6 - 10.9;
and spread of caudal flukes from apex to apex 23.8- 27.3 and 23.0- 28.4.

The measurements of 14 skulls (Japan; Nishiwaki, 1966) with a
condylobasal length of 356 to 390 mm (as percentage of this length)
are: length of rostrum 44.7-49.2, basal width of rostrum 27.7-31.8,
interorbital width 53.7 - 60.7, length of lower jaw 72.2 - 79.9, and length
of mandibular symphysis 8.7 - 10.1.

Information on the geographic distribution of this species is very
scant, with only a few recorded finds and observations (Fig. 308). It has
been noticed in the waters of Senegal and in the North Atlantic. It is
known in the Yangtze estuary, along the coasts of Honshu Island, and
in the North Pacific Ocean. (V.S.)

Biology has not been studied.

721

Fig. 308. Pygmy killer whale, Feresa attenuata (figure by М.М. Kondakov).
A—coloration of underside; B—of upper side.

Genus of Common Porpoises
Genus Phocoena G. Cuvier, 1817

1817. Phocoena. G. Cuvier. Rgne Animal, ed. I, I, p. 279. Delphinus
phocoena Linnaeus, 1758.

1828. Phocoena. Gray. Spicil. Zoologica, I, р. 2. Delphinus phocoena Lin-
naeus, 1758. (V.H.)

Small dolphins, with a maximum body length of 2 m.

These dolphins have no “beak”. The head is obtuse and the forehead
flattened. The anterior margin of the low dorsal fin, short flippers, and
caudal flukes bears small horny tubercles (sometimes these are present
only on the dorsal fin).

The body is dorsally dark, gradually turning white on the ventral
surface.

The rostrum of the skull is equal to or slightly shorter than the
cranium. The posterior section of the premaxillae anterior to the bony
nares forms a prominence. The pterygoid bones are separated by a gap.
The boundaries of the interparietal bones are invariably very prominent.

Teeth 16-39. They have a slightly flattened crown and are set off from
the root by a neck. Vertebrae 62-66. Ribs 12-14 pairs. The sternum is
non-segmented. Phalangeal formula: I, _ 3, II; - 49, III; - 3, [У›-в›апа У -3.

These dolphins are ichthyophagous, surviving mainly on benthic
fish (bentho-ichthyophagous). The females whelp every year. Gestation
extends for nine to ten months and lactation for about four months.

538

722

Females are ready for fertilization two to three months after parturition,
while still nursing a calf.

Common porpoises are distributed in the North Atlantic from the
Barents Sea and Davis Strait to New Jersey in the west and the Mediter-
ranean, Black, and Azov seas, and Senegal waters; in the South Atlantic
from Rio de la Plata (35° S lat.) to Cape Horn and South Georgia; in
the North Pacific Ocean, from the Chukchi Sea to Mexico in the east
and Japan in the west; in the South Pacific Ocean, from Paita in Peru
(5° S lat.) to Cape Horn (Hershkovitz, 1966) (Fig. 309).

The members of this genus are thus absent in the Indian Ocean, in
the western part of the South Pacific Ocean, and in some tropical and
equatorial parts of the Atlantic and Pacific oceans.

The genus comprises four species: 1) P. phocoena Linnaeus, 1758;
2) P. dioptrica Lahille, 1893; 3) P. spinnipinnis Burmeister, 1865; and
4) P. sinus Norris and McFarlan, 1958. Sometimes the genus is divided
into a larger number of species.

Only one species is encountered in the USSR waters: the common
porpoise, P. phocoena Linnaeus, 1758.

The common porpoises are presently not of commercial importance.
(V.S.)

COMMON PORPOISE}
Phocoena phocoena Linnaeus, 1758

1758. Delphinus phocoena. Linnaeus. Syst. Nat., I, p. 77. Baltic Sea
(“Oceano Europeo Balthico’’)

1827. Phocoena communis. Lesson. Man. Mammal, p. 413. Renamed as
Delphinus phocoena auct.

1865. Phocoena vomerina. Gill. Proc. Ac. Nat. Sc. Philadelphia, 17,
p. 178. Puget Sound, Washington State.

1905. Phocoena relicta. Abel. Jahrb. K.K. Geol. Reicheanstate Wien, 55,
p. 387. Crimean waters of the Black Sea. (V.H.)

Diagnosis
Only species of the genus encountered in the USSR.
Description

The external appearance of the common porpoise is typical: the body
build is much heavier than that of dolphins, somewhat stunted, and the

15 Also locally known as “Azovka,” “Pykhtun,” “Svinka” [porpoise], etc. (Black Sea).

|
|

Hl

1f

i
|
th

|

1000 © 1000 2000 3000 4000 5000 km

Fig. 309. Range of the genus of common porpoises, Phocoena (V.A. Arsen’ev).

537

724

7 fe L SS
MODEL Wi LLL
fi \ te И
i i o
“on
af)
539 Fig. 310. Common porpoise, Phocoena phocoena (figure by N.N. Kondakov).

trunk thickset. The head is short. The flippers are oval and the dorsal
fins triangular (Fig. 310).

Horny tubercles 12 - 16, less frequently 18-20, occur along the ante-
rior margin of the dorsal fin in common porpoises (adult animals and
embryos) of the Black and Azov seas (Tsalkin, 1938b). It has been sug-
gested that these tubercles represent derivatives of the thick skin cover
(Kukenthal, 1889-1893). Two or three hairs are seen on each side of
the snout in embryos. The color of the dorsal surface varies from a dark
gray to almost black. The light-colored abdomen is sometimes sharply
demarcated from the dark dorsum but sometimes the transition in col-
oration is altogether imperceptible (Tsalkin, 1938b). A faintly percepti-
ble dark gray band runs from the zone of the ear openings toward the
navel on each side of the body, while another almost black band extends
from the corner of the mouth to the base of each flipper (Barabash-
Nikiforov, 1940). Instances of partial or total albinism are encountered
occasionally.

The form of the tooth crown is variable—from almost conical to
a form with three faintly discernible cusps. The anterior surface of the
crown is slightly turned inside the cavity. When the mouth is closed, the
teeth on the lower jaw fall inward of the upper teeth. Among the Black
and Azov sea common porpoises, the number of teeth in the upper jaw
varies from 44 to 60 (more often, 54) and in the lower jaw 38 to 55 (more

539 often, 44-46) (Tsalkin, 1938b). The teeth of this common porpoise are
characterized by curvature, distortion of the roots, and thickening in
the lower portions (Tsalkin, 1938; Barabash-Nikiforov, 1940). As age
advances, the teeth may wear down or even drop out. Cervical vertebrae
7, thoracic 12-14, lumbar 14-17, and caudal 27 -32. The sections of the
vertebral column comprise (as percentage of its length): cervical 1.9-3,
thoracic 22.1 -24.6, lumbar 30 - 34.2, and caudal 40-44 (Tomilin, 1957).
Six to eight pairs of ribs are articulated with the sternum in the embryos
and usually five pairs in adults.

725

Females are slightly larger than males. The difference in body length
between mature females and males from the Black and Azov seas
averages 7 cm (Tsalkin, 1938).

The main body measurements of this species of common porpoises
(eight animals) from the Black Sea average (Tsalkin, 1938) (in cm): body
length 163; distance from tip of snout to base of flippers 32, to anterior
margin of dorsal fin 73; and length of flippers 22. The length of the
largest common porpoises from the Black and Azov seas is (in cm):
male 167 and female 180; from the North Pacific Ocean: female 178.5;
and from the North Atlantic: male 186.

The average main skull measurements (measurements of five males
and three females; Tomilin, 1957) (Fig. 311) are respectively (in cm):
condylobasal length 25 and 27, length of rostrum 11 and 12, width of
rostrum at base 7 and 8, and length of lower jaw 19 and 21. (V.S.)

Geographic Distribution -

Temperate and partly cold waters of the North Atlantic and the North
Pacific oceans; predominantly the coastal zone.

Geographic Range in the USSR

In the basin of the Atlantic Ocean, this porpoise is commonly
encountered in the Barents Sea where it reaches the coasts of Novaya
Zemlya and Yugorsk Shar and is abundant on the Murman coast; it is
also common in the White Sea (waters of Kanin, Letn’aya and Tersk

539 Fig. 311. Skull of the common porpoise, Phocoena phocoena (figure by М.М. Kondakov).

542

726

coasts, and Gulf of Kandalaksh), Kara Sea (transgresses into its western
section), Baltic Sea, and almost and the entire coastal water body of the
Black and Azov seas (Fig. 312).

In our waters of the Pacific Basin, it is common in the Sea of Japan
and the Sea of Okhotsk, on both sides of the Kuril range, along the
coasts of Kamchatka, Koryatsk land, and the Gulf of Anadyr, around
the Commander Islands in the Bering Sea, in the Bering Strait, and in
the Chukchi Sea where it penetrates north of 70° N lat. (east of Point
Barrow, 71°24’ N lat.). Its range in the west is not known.

Geographic Range outside the USSR

In the western part of the North Atlantic, the common porpoise is known
from New Jersey (39° N lat.), and possibly more southward, up to Davis
Strait, Baffin Bay, and coasts of Greenland in the eastern part of the
Atlantic Ocean, from the waters of Senegal and Dakar in northern Africa
(about 15° N lat.) to the Barents Sea, Iceland, and east coast of Green-
land (Scoresby Sound), i.e., roughly up to 70° N lat. The range covers
the entire Baltic Sea. Transgressions into the Mediterranean Sea from
the Black Sea as well as from the Atlantic Ocean are known (Fig. 313).

In the Pacific Ocean, this species lives along the eastern coasts of
the Korean peninsula and in the waters of Japan. It inhabits the entire
coastal waters of the American continent down to California and Mexico
from the Chukchi Sea (Point Barrow) and the Bering Sea, including
the coastal waters of the Aleutian Islands (Kleinenberg, 1956a; Tomilin,
1957; Hall and Kelson, 1959; Nishiwaki, 1966). (V.A.)

Geographic Variation

In the Northern hemisphere, the species forms three subspecies that
are almost indistinguishable although geographically disjointed. All the
three subspecies are encountered in the waters of the USSR.
1. Northern Atlantic common porpoise, P. p. phocoena L., 1758 (syn.
communis). This is the largest of the subspecies with a body length up
to 186 cm, condylobasal length of skull 255-259 mm, width of condyle
65-71 mm, and height of occipital condyle 38-42 mm. The rostrum
is hardly shortened. Tubercles are seen on the anterior margins of the
flippers, dorsal fin, and caudal flukes.

This subspecies is encountered in the Barents, White, Kara, and
Baltic seas.

Outside the USSR, it has been reported in the waters of the North -
Atlantic Ocean.

727

‘(лэ.иэзту у) USSN э41 Ul

9 и2020ц4 nusos0yq ‘asiodiod иоититоо ay) Jo aduey ‘21$ ‘89

1 OL ov
у : №!
ic wh
se К
1 АИК f
=e $
©. | '
г. A R 2 ,
ый С
7! a =
5 ЧвеЧ3еЯ ay $) E
Г $ ‘ ay
а Ч Ч

728

= 3
АНИ
i Zar ul
И \ |
АИ
an Wea

ah Diet tt
male ИКИ tit

tin | Г ЧОН ИИ

о ОО: HPO

© Я Ин

ИО И к
- ORO а
ites ae
72 НОТ
ee |

т :

AU

НИ
Ва

120

Я Ги
oo И i! iy 1
р ИАИИИ
НО

his |

о
y Wa к Г
хи Nate a
UH i a yh

150
150

1000 2000 3000 4000 £000 km

180

1000 0

[ten ЧИ
КНИГИ

Fig. 313. Species range of the common porpoise, Phocoena phocoena (V.A. Arsen’ev).

541

543

729

2. Black Sea common porpoise, Р. р. relicta Abel, 1905. This form is
of medium dimensions, with a body length up to 180 cm, condylobasal
length of skull in males 245-255 mm, in females 269-274 mm; width of
condyle in males 54-62 mm, in females 58-63 mm; height of occipital
condyle in males 32 -36 mm, in females 37-39 mm. The tubercles on the
anterior margin of the fins are totally reduced.

This subspecies is encountered in the Black and Azov seas.

Outside the USSR, transgressions have been reported for the

Marmora and Mediterranean seas.
3. Pacific common porpoise, P. p. vomerina Gill, 1865. This is the smallest
of the subspecies with a body length up to 178 cm, condylobasal length of
skull 258-293 mm, width of condyle 68-70 mm, and height of occipital
condyle 40-43 mm. The rostrum is highly elongated.

This subspecies is encountered in the Chukchi, Bering, Okhotsk, and
Japan seas, and in the waters of Kuril Islands from the Pacific Ocean
side.

Outside the USSR, it has been reported in the American waters of
the Pacific Ocean from the Chukchi Sea to the Mexican coasts and in
the waters of Japan. (V.A.)

Biology

Population. The species can be regarded as relatively abundant. The least
populous of the three subspecies is the Azov-Black Sea common por-
poise, which is confined to an extremely restricted range. Populations of
the other two subspecies occupy extensive water bodies in which they are
commonly encountered and are perhaps quite abundant.

Food. Information on the food of the Atlantic and Pacific popula-
tions is extremely scant while that on the food of the Black Sea popula-
tion is more widely available (Table 46).

Two species of pelagic fish, viz., smelt (49.5%) and Black Sea anchovy
(18.5%), occupy the primary position in terms of the quantum of fish
found in the stomach of the Azov-Black Sea common porpoise. All the
species of goby (bottom-dwelling fish) comprise only 30.9%. The rest of
the species of fish in the food of the common porpoise constitute only a
few tenths of one per cent. In terms of the number of fish objects encoun-
tered, pelagic fish comprise 68.8% and benthic fish 31.2%. However, in
terms of weight (as percentage of total weight of food), rotan goby occu-
pies the first position (32.4%), followed by round goby (31.2%), smelt
(14.5%), and Black Sea anchovy (11.0%). Thus, the group of benthic fish
constitutes 67.9% of the total weight of food intake and the group of
pelagic fish 32.1%

730

Table 46. Food objects of the common porpoise

Azov-Black Sea
(Tsalkin, 1940)

Fish
Round goby, Gobius
melanostomus
Rotan goby, G. rotan
Mushroom goby, G. cephalarges
Syrman goby, G. syrman
Toad goby, Mesogobius
batrachocephalus
Black Sea flounder,
Pleuronectes flesus
luscus
Black Sea sole, Solea nasuta
Black Sea anchovy,
Engraulis encrasicholus
Black Sea silverside-smelt,
Atherina pontica
Perch, Lucioperca lucioperca
Bream, Abramis brama
Golden gray mullet, Mugil
auratus
Black Sea whiting (haddock),
Gadus euxinus
Black Sea shad (herring),
Caspialosa sp.

Crustaceans
Balanus improvisus
Brachinotus lucassi
Idothea baltica
Leander sp.

Mollusks

Algae
Ulva lactuca

North Atlantic
(Tomilin, 1957, 1962)

Fish
Cod, Gadus morhua
Capelin, Mallotus
villosus
Navaga, Eleginus
navaga
Sand eel, Ammodytes
hexapterus
Whiting, Odontogadus
merlangus
Common mackerel,

_ Scomber scombrus

Pollock, Pollachius
virens

Herring, Clupea
harengus

Sardine, Sardina sp.
Sole, Solea sp.

Eel, Anguilla anguilla
Salmon (small)

Sprat, Sprattus [= clupea}

sprattus
Trout

Crustaceans
Decapods

Mollusks
Cephalopods (Loligo
pealitii)

Algae

North Pacific (Sleptsov, 1955;
Tomilin, 1957)

Fish
Pacific cod, Gadus morhua
macrocephalus
Capelin, Mallotus villosus
Pacific navaga (saffron cod),
Eleginus gracilis
Polar cod, Boreogadus saida
Herring, Clupea harengus
Shad, Alosa sapidissima
Shark, Squalus sp. (?)
Sablefish (black cod),
Anoplopoma fimbria
Whitefish (Leucichtys)

Crustaceans
Shrimps

Mollusks
Cephalopods

Most mollusks probably enter the stomach of porpoises through
the fish they consume. Two species of crustaceans (Idothea baltica and
Leander sp.) can be regarded as incidental food objects as they were found
in only two stomachs. Alga was found in the three groups of porpoise in
large quantities and probably serves as food.

Thus, fish represent the almost lone source of food to the Azov-Black
Sea common porpoise; most of them are benthic and a small percentage

544

731

pelagic. The latter serve as the main food only in the period of massive
migrations (Black Sea anchovy) when they form concentrations of large
proportions.

Seasonal variations in food types have been noticed: benthic fish
predominate in winter, while in summer Black Sea anchovy and smelt
play an extremely important role in the food of common porpoises during
the period of migrations of these fish from the Black Sea into the Azov
Sea and back in spring and autumn (Tsalkin, 1940).

The North Atlantic porpoises feed mainly on fish (benthic as well as
pelagic forms). Cephalopods and other mollusks, crustaceans, and algae
play no significant role in their food. The nature of the food of the Pacific
porpoises is very similar to that of the other populations.

Daily activity and behavior. The porpoises of the Azov-Black Sea pop-
ulations (probably the others too) avoid the open seas and are encoun-
tered more in the coastal shallow waters (Fig. 314). They are usually
confined to small groups of not more than 10 animals. Relatively larger
groups are encountered only in the period of massive migrations of
fish which serve as their food. Exhalation and inhalation occur at not
more than 30-sec intervals although instances of porpoises remaining
submerged for 6 min are known. Their movement is comparatively slow.
Intense flexure of the body while diving is a characteristic feature. The
porpoises of this species, with rare exception, do not trail behind moving
ships.

Fig. 314. Common porpoises at sea (figure by N.N. Kondakov).

545

32

Instances of the transgression of porpoises into rivers are not rare.
They have been noticed not only in the Black Sea rivers (Don and
Danube), but also in the Thames, Seine, Rhine, Elba, Neva, and other
rivers. Sometimes the animals advance up the river for hundreds of kilo-
meters from the estuary (Kleinenberg, 1956a; Tomilin, 1957, 1962).

Seasonal migrations and transgressions. The migrations of common
porpoises have not been studied although some seasonal migrations
undoubtedly do occur. It is quite obvious that these porpoises spend only
the summer months in the northern parts of the range of the Atlantic
and Pacific oceans but abandon these waters in winter. The southward
movement of these porpoises into the Baltic Sea is quite impressive.
Short seasonal migrations have been observed in the Black Sea in which
the animals spend the winter; in spring, they are seen in the Azov Sea
but return again to the Black Sea for winter.

The rare departures of common porpoises from the Black Sea into
the Sea of Marmora and even into the Mediterranean Sea, their trans-
gressions into the Mediterranean Sea from the Atlantic Ocean, and
their passage in the easternmost part of the Baltic Sea (Kronshtadt)
and through Neva even into Lake Ladoga, can be regarded as unusual

transgressions.

Reproduction, growth, and development. Data on reproduction biology
are available mostly for the Azov-Black Sea populations but these can
be extrapolated to the other two populations with sufficient justification.

The periods of mating and parturition, although quite protracted,
occur predominantly in the summer months. The Black Sea common
porpoise mates from the end of June to October with a peak in August.
The duration of gestation has been put at 9-10 months. Births occur
from April to July. Most females whelp in May-June (Tsalkin, 1940) In
the Baltic Sea, mating peaks in July and August while the largest number
of calves are born in May and June. The growth rate of the embryo is
quite high (Table 47). The embryos of common porpoises (Fig. 315) from
the Baltic sea recorded an average weight increment of 5 g per day (total
160 g) in the one-month period from November 15 through December
15, 11 g per day (total 340 g) from December 15 through January 15,
and 30 g per day (total 920 g) from January 15 through February 15.

Usually a single large calf (twins are very rare) measuring half the
length of the mother’s body, sometimes even longer, is delivered. The
average length of a newborn calf is 75 cm (63-86 cm) and it weigh
2.8-7.9 kg. Lactation extends for about four months. The composition
of the milk of the Azov-Black Sea common porpoise is as follows (7%):
fat 33.9, protein 5.22, sugar 1.28, dry residue 7.1, ash 0.6, and water 59.0
(Ural’skaya, 1957). In some cases, the milk may contain up to 45.8% fat.

546

733

Fig. 315. Embryo of the common porpoise (figure by М.М. Kondakov).

Table 47. Monthwise dimensions of embryos, mm (Tomilin, 1957)

Month Azov-Black Sea basin North Atlantic Ocean and North Pacific Ocean
Baltic Sea ;
Number Number Number
of of of

animals Mean Min. Max. animals Mean Min. Max. animals Mean Min. Max.

July ау 5 ПУ ое рае ит еб
Aug. SOS Oe ee ee ee
Sept. 8760 10 0 =
Oct. 67 120 50 200 ее See
Nov. 33 190 80 240 GT оО ee о
Dec. LPO aD 300 ns ee CLS ee ee
Jan. Gi 320190} S80 АГ. 429600225.) 380. 212 248.289. 258
Feb. ae en ere 23 E356 250) 450 ВИ ser all
March 263 460 210 610 20153552001 550 10909 Ronee
April 28 580 370 820 5101650] 1590174011912 14951180510
Мау 35 730 620 840 24) 1675610448740, мили 6б0Зн— В —
June 7 820 780 850 eA VE SS ee
Total 453 10 850 134 92. 1740058 239 510

The calf is capable of swimming and accompanying the mother immedi-
ately after birth. The male to female ratio among newborn calves is close
to one.

At a body length of 130-145 cm, many females of the Black Sea
common porpoise are still immature. More than half the females (59%)
with a body length of 145-150 cm and 79-100% of females 150 cm or
longer were gestating. The smallest gestating female among the Black Sea
common porpoises was 130 cm long and among the Baltic Sea animals
136 cm (weight 50 kg) (Tomilin, 1962).

Enemies, diseases, parasites, mortality, and competitors. In oceanic
waters the killer whale represents the most dangerous enemy of the com-
mon porpoise, which is a food item for the former. It is possible that the

734

porpoise may fall prey to large sharks. Apart from bone tumors, no other
diseases have been reported for these porpoises. Very severe infection
with helminths could be the cause of death in some cases. Among all the
cetaceans, the common porpoise is one of the primary hosts for a large
number of helminths.

Eighteen species of helminths have been registered in the common
porpoise: trematodes four, cestodes three, nematodes nine, and acantho-
cephalans two.

Two species of trematodes have been reported exclusively in common
porpoises. Campula oblonga Cobbold, parasitizing the bile ducts of the
liver, was detected in the European and American waters of the North
Atlantic. Pholeter gastrophilus Kossack, parasitizing the mucous mem-
brane of the pyloric section of the stomach, was found among dolphins
caught in Baltiisk port. The trematode Distomum philocholum Creplin,
localizing in the liver, has been encountered in white-sided dolphins in
European waters as well as in common porpoises. This species para-
sitizes the bile ducts of the liver of common porpoises and three species
of pinnipeds in the Atlantic and northern Arctic oceans; it is also known
in three species of land carnivores.

The three species of cestodes belong to the genus Diphyllobothrium.
Of these, D. stemmacephalum Cobbold was found in the small intestine
of only common porpoises (North Atlantic and Black Sea on the Ruma-
nian coasts); D. lanceolatum Krabbe parasitizes the intestine of common
porpoises and four species of seals (North Atlantic and Pacific oceans);
D. latum L., apart from common porpoise, is found in six species of pin-
nipeds, several species of land carnivores, domestic animals, and man;
it has been detected among marine animals in the North Atlantic and
northern Arctic oceans.

Anisakis (Anisakis) simplex Rudolphi, a nematode of common por-
poises, is very widely distributed among marine mammals (parasite of
the gullet, stomach, and intestine) and has been found in ten species
of toothed whales, two species of baleen whales, and in Steller’s sea
lion in the North Sea and in the Pacific Ocean (Kamchatka, Japan, and
New Zealand). Anisakis (Anisakis) typica Diesing, in addition to common
porpoises, is a parasite of the stomach of three species of dolphins from
the North Sea and waters of South Africa in the Atlantic Ocean. Terra-
nova (Terranova) decipiens Krabbe, apart from common porpoises, has
been reported among many species of marine mammals (17 species and
subspecies of pinnipeds, 2 species of baleen whales, and 1 species of dol-
phins) and localizes in the stomach and intestine; it has been detected in
the basin of the northern Arctic, Atlantic, and Pacific oceans, and around
Antarctica. The nematode Pseudalius inflexcus Rudolphi has been found

547

735

in the bronchi, blood vessels, and heart only of common porpoises from
the North Atlantic coasts of Europe and the Asian coasts of the Pacific
Ocean (not recorded in USSR waters). Of the three species of the genus
Halocerus, Halocerus (Prohalocerus) invaginatus Quekett has been found
in the lungs of only common porpoises from the waters of California, the
Baltic Sea, and the Atlantic Ocean (not found in USSR waters). Halo-
cerus (Posthalocerus) taurica Delamure and Skrjabin and H. (Posthalo-
cerus) ponticus Delamure parasitize the lungs of only the Azov-Black Sea
common porpoise. Stenurus minor Kuhn, aside from Azov-Black Sea and
Atlantic common porpoises, has been found in belugas in the northern
Arctic Ocean, North Atlantic Ocean, North Sea, on the Asian coasts of
the Pacific Ocean, and in the Black and Azov seas. It localizes in the
bronchi, heart, blood vessels, and auditory organs. It is possible that the
large number of the nematode Stenurus minor encountered in all the
Azov Sea porpoises may cause deafness in them. Tozynurus convolutus
Kuhn, parasitizing the bronchi and blood vessels of the lungs of animals
in the European waters of the North Atlantic and coasts of Sakhalin in
the Pacific Ocean, has been detected in pilot whales as well as common
porpoises.

The acanthocephalans Corynosoma semerme Forssell and Coryno-
soma strumosum Rudolphi parasitize the intestine. The former is known
in common porpoises, 6 species of pinnipeds, and 6 species of sea birds;
the latter in common porpoises, 11 species of pinnipeds, belugas, 10
species of birds, and in cats and dogs. These worms have been detected
in almost all the northern seas and in the Caspian Sea (Delamure, 1955).

Field characteristics. These are rather small, predominantly coastal
porpoises. The upper ‘part of the trunk is black and the underside light-
colored. The tip of the dorsal fin is almost rectangular. These animals live
in small groups. Intense flexure of the body while diving is characteristic.
They generally do not breach the water.

Economic Importance

Common porpoises are of almost no economic importance. Danish fish-
ermen catch several hundreds of porpoises a year in the Little Baelt
Strait during their migration from the Baltic Sea. They chase them into
the narrow strait and block them in nets. In other parts of the range, the
catch is only incidental.

Regular hunting of common porpoises is carried out only in the
Azov and Black seas but here, too, this species is not of great economic
importance. In the best years of hunting, the proportions of the differ-
ent species of dolphins in the catch from this water body are: common

736

dolphin 200, common porpoise (from Azov Sea) 10, and bottlenose
dolphin 1 (Tsalkin, 1937). This is based on a catch of 700-800 Azov
Sea common porpoise; in some years up to 4,000 animals are caught and
the proportions vary considerably (Bodrov, Grigor’ev, and ‘Tver’yanovich,
1958).

The technique of catching the Azov Sea common porpoise, although
not different from that used for catching the common dolphin, has cer-
tain special features. The much slower Azov Sea common porpoise is
usually corifined to small groups, distinctly separated, and in the coastal
zone. Therefore, whalers spread the fleet in a broad front, advance for-

548 ward in flanks, and gradually press the animals toward the coast. The
net is cast after the dolphins gather in a very dense group close to the
coast.

Oil, mainly melted from blubber, represents the chief product of
hunting. Further, comparatively high-quality leather goods are produced
and used satisfactorily in the footwear industry. Despite many success-
ful experiments on using the flesh of Black Sea dolphins for human
consumption, it has not found wide acceptance. The material remaining
after obtaining the oil (musculature, skeleton, fins, viscera, etc.) is made
into a flesh-bone meal which can be used as feed for farm animals or
as a fertilizer. The hollow lower jaws of dolphins contain a very small
amount of so-called jaw oil, from which high-quality oil is produced for
lubricating fine mechanisms. In experiments, this oil froze after 10 hr of
exposure to —7 to —8°C; the oil was then filtered at this temperature
and transferred to a chamber at —25°C where it did not freeze for 24 hr
(Okuneva, 1934).

The total weight of the Azov-Black Sea common porpoise varies in
the range 24 to 55 kg (Okuneva, 1934), or an average of 28 kg (Dragunov
and Kasinova, 1951). The weight of the various trunk sections in different
seasons is shown in Tables 48 and 49.

Three very large female common porpoises from the North Pacific
Ocean with a body length of 178.5 cm, 173 cm, and 168 cm weighed
respectively: 75.2 kg, 70.3 kg, and 73.4 kg (Scheffer and Slipp, 1948).

In the overall balance of useful products produced by hunting marine
mammals in our country, common porpoises have no economic impor-
tance whatsoever. As a result of the significant reduction in the pop-
ulation of dolphins in the Black and Azov seas, hunting dolphins of
all species (including the Black Sea common porpoise) has been totally
banned since 1965. There is also no hunting in the other areas of the
range of this species (in the Atlantic and Pacific oceans). (V.A.)

547

548

737

Genus of Dall Porpoises
Genus Phocoenoides Andrews, 1911

1911. Phocoenoides. Andrews. Bull. Amer. Mus. Nat. Hist., 30, p. 31.
Phocoenoides truei Andrews, 1911.
Small porpoises with a body length up to 2 m.
These porpoises lack a “beak”. The low dorsal fin lies slightly ante-
rior to midbody. The flippers are relatively small.

Table 48. Weight of the body sections of Azov Sea common porpoise caught in October
and November (Okuneva, 1934)

Item Male Female
As % of As % of
Weight, kg total weight Weight, kg total weight
Total weight 24.0 - 27.0 100 37.0-55.0 100
Head 8123 775) 2.4-3.6 6.5
Skin Iasi 5 7258) 7.3-8.8 2.0-2.5 5.4-5.9
Trunk fat hs) 92 29.3 -35.4 LES = 7.5 25.3-30.6
Fins and tail 1.0. = 1:1 2.5 -4.1 1.5 -1.8 3.3-3.9
Muscles (flesh) 4.8 - 6.5 19.7-24.1 10.0- 15.5 24.5-27.1
Trunk bones 2.9- 43 10.8 -16.3 3.5- 6.5 9.5-11.8
Brain 0.3 - 0.4 1.2-1.4 — —
Tongue ONS 02 0.5 - 0.6 0.15 0.40
Blood — — 1.0 2.17
Viscera, inclusive of: 3.4 - 4.2 10.6-15.4 6.0-8.5 13.0 - 17.7
Liver 0.5 - 0.6 1.7-2.3 1.1-1.6 2.0-3.3
Heart 0.15 0.62 0.15 0.40
Lungs 0.6 - 0.8 2.1-2.8 1.0-1.6 1.9-3.3
Stomach and intestine 14- 1.6 5.2-6.2 5.1-5.8 10.4-10.5
Kidney 0.1 - 0.2 0.3-0.6 0.20

0.36

Table 49. Weight of Azov Sea common porpoise in the spring catch (2,611 animals)
(Tomilin, 1957)

Total weight, Average weight, As % of
kg kg total weight

Total weight 78,888 30.2 100

Blubber 32,026 12.3 40.6
Muscles (flesh) 24,018 9.2 31.2
Fins 2,956 Л &Ы/
Глуег 1,566 0.6 2.0
Heart 263 0.1 0.3

549

738

The body color is dark (from dark steel to black). A large white
patch occurs on each side of the body. The dorsal fin is partly or wholly
white.

The broad and flattened rostrum is shorter than the cranium. The

pterygoid bones are separated. The teeth are very small and number
19-23-28

55-24-58. Vertebrae 92-98. Phalangeal formula: [_›, II¢-7, Ш4- в

ГУ, - 3, and У, ->. Ribs 15-18 pairs, of which 3-5 anterior ones articulate
with the sternum.

Biology of Dall porpoise has not been well studied. They feed on
cephalopods and fish.

They are distributed in the North Pacific Ocean.

The genus comprises a single species, P. dalli True, 1885.

There is no hunting of this species. (V.S.)

DALL PORPOISE
Phocoenoides dalli True, 1885

1885. Phocoena dalli. True. Proc. U. S. Nat. Mus., 8, p. 95.
1911. Phocoenoides truei. Andrews. Bull. Am. Mus. N. H., 30, p. 32.
Waters of the eastern coast of Japan; Rikuzen Peninsula, Hondo
‚ Island. (V.H.)

Diagnosis
Only species of the genus.
Description

The external appearance is similar to that of other common porpoises.
The body is shortened and the anterior half is somewhat thickset. The
head is short (Fig. 316). The upper jaw is slightly shorter than the lower.
The dorsal fin forms a near-equilateral triangle, slightly concave along
the posterior margin. The length of the base of the dorsal fin is 1.7-2.2
times its height. A few small horny tubercles are present on the anterior
margin of the dorsal fin. High longitudinal keels are present on the
caudal stem, both in the upper and lower parts. These keels increase
with age. The flippers are short and have a broad base.

The color of the head, dorsum, upper anterior and entire posterior
part of the caudal stem is dark, from dark steel gray to black. A large
white patch occurs on each flank and extends anteriorly up to the level
of the anterior margin of the dorsal fin (P. d. dalli) or almost right up
to the eye (P. d. truei). This patch posteriorly extends along the flanks of
the caudal stem to the level of the anal opening but does not reach the

‘(aoyepuoy ‘м ‘М Aq ainsi) чеээо эцюеа ‘эпдт, yop 5эрлоизороца ‘[astodiod пе]
“AI 971d

р
и
yaad

hi

549

550

739

Fig. 316. Dall porpoise, Phocoenoides ааШ (figure by М.М. Kondakov).

abdomen. Individuals with a coloration intermediate between these two
types have been sighted. The dorsal fin is wholly or partly white (dark
only rarely). The flippers are somewhat lighter in color than the trunk,
even white. in exceptional cases. Totally black specimens have also been
encountered (Nishiwaki, 1966). Young animals are usually more deeply
pigmented and newborn calves less pigmented than adult animals.

The teeth are small, chisel-shaped, and barely rise above the gums.
Tubercles of keratinized epithelium are present on the gums around
the teeth. Some teeth may not cut through the gums. Thus, in an adult
male, 5 of 15 teeth in the left half of the upper jaw and 9 of 16 in
the right half had not emerged; 4 of 22 in the left half of the lower
jaw and 4 of 22 in the right half were likewise unexposed (Benson and
Grood, 1942). The interalveolar septa are poorly developed. The teeth
are surrounded by tubercles of keratinized epithelium of the gums which
could rise above the tooth crowns and function as a substitute or them.
Cervical vertebrae 7, thoracic 15-18, lumbar 23-27, and caudal 39-49.
The cervical section constitutes 3%, thoracic 23%, lumbar 33%, and
caudal 41% of the length of the vertebral column. The vertebrae have
a flattened centrum and a long thin spinous process reaching maximum
height in the lumbar section. All the cervical vertebrae are fused.

The minimum and maximum body lengths in 14 adult male and 13
adult female Dall porpoises were respectively (in cm): 172 and 210 (х
186) and 169 and 197 (x 181) (Mizue and Joshida, 1965). The main body
measurements of two male Dall porpoises, one caught off the coast of
Japan and the other from the Aleutian Islands, and a female caught
off the coast of Japan (Tomilin, 1957; Norris, 1966) were respectively
(in cm): body length 191, 183, and 182; distance from tip of snout to
anterior margin of blowhole 25, 23, and 15; the same up to axilla 41, 36,

740

550 Fig. 317. Dall porpoise, Phocoenoides dalli, on the deck of а ship. Sea of Japan,
1965 (photograph by А.Е. Kuzin).

551 and 24; length of flippers 23, 20, and 23; maximum width of flippers 22,
10, and 10; height of dorsal fin 16, 15, and 16; width of caudal flukes
(from tip to tip) 47, 47, and 47; and distance from anal opening to fork
of caudal flukes 62, 58, and 51. The largest male caught to date had a
body length of 210 cm and the female 198 cm.

The skull measurements (Fig. 318) (average of three to five animals;
Tomilin, 1957) were (in cm): condylobasal length 32, zygomatic width
19, length or rostrum 13, width of rostrum at base 10, length of lower
jaw 25, and length of mandibular symphysis 3.7. (V.S.)

Geographic Distribution

This species is encountered in the North Pacific Ocean.

Geographic Range in the USSR (Fig. 319)

Coasts of the Sea of Japan commencing from Peter the Great Gulf,
La Perouse Strait, waters of Sakhalin including the Gulf of Sakhalin and
Kuril Islands, southeastern part of the Sea of Okhotsk, and western coast
of Kamchatka. In the Bering Sea, it is encountered along the eastern
coast of Kamchatka, in waters of Koryask land, Gulf of Anadyr and
Bering Strait, and the adjoining waters of the Chukchi Sea.

550

553

741

Win 20. ААА
а №

"ль *

(Ам, AMY
Ч ty, Woh

ee мили

Fig. 318. Skull of the Dall porpoise, Phocaenoides ааШ (figure by М.М. Kondakov).

Geographic Range outside the USSR (Fig. 320)

Waters of the eastern and western coasts of Japan, Aleutian Islands, and
eastern part of the Bering Sea, Gulf of Alaska, and along the Amer-
ican mainland in the south up to 30° М lat. (McTaggart, 1944; Wilke,
Taniwaki, and Kuroda, 1953; Tomilin, 1957; Hall and Kelson, 1959).
(V.A)

Geographic Variation

Two well-distinguished subspecies, also encountered in our waters, are
recognized in this species. Some researchers, however, do not recognize
these forms while others treat them as independent species. Details of
distribution at the points of contact of the ranges, the numerical ratios
between the two forms at such places, etc. are not available. Their ranges
in Our southern waters overlap.

1. Northern Dall porpoise, Р. d. dalli True, 1885. The white patch on the
flanks and ventral side of the body does not extend to the flippers but
terminates roughly on the vertical of the anterior margin of the dorsal
fin.

This subspecies is encountered in the waters south of Ussuri territory
(Peter the Great Gulf and Pos’et Bay) and the southern islands of the
Kuril range to the southern part of the Chukchi Sea inclusive.

Outside the USSR, this subspecies is reported in the waters of the
Korean peninsula and Japan in the south up to roughly 38° N lat. and
in American waters from the Chukchi Sea up to roughly 34° М lat.

742

2. Southern Dall porpoise, Р. 4. гие Andersen, 1911. The white patch
on the flanks runs anteriorly beyond the base of the flippers and almost
reaches the eye; on the abdomen, it runs from the base of the caudal stem
to the base of the flippers. A small white patch occurs on the throat.

This subspecies is reported in the waters of the southernmost parts of
Ussuri territory (Primor’e) in the north up to 42 to 43° N lat. (Pos’et Bay,
Peter the Great Gulf, and more eastward), and possibly in the southern
part of the Kuril range.

Outside the USSR, it is reported in the waters of the Korean penin-
sula and Japan between 38° and 43° N lat. Along the American coast, it

hy
НИ

551 Вр. 319. Range of the Dall porpoise, Phocoenoides dalli, т the USSR (V.A. Arsen’ev).

743

(лэмэзту У‘)

yjop 52рлоизо2оц4 ‘asiodiod ива эцз Jo э8иел $90245 ‘00$ “314

И

ИМ:
п Е

GSS,

744

has been indicated from 28 -30° М lat. to 45-50° М lat. (Klumov, 1959;
Nishiwaki, 1966).'° (V.A.)

Biology

Population. Probably, one of the relatively abundant species, forming at
times large herds of hundreds of animals. It is apparently quite abundant
in the western parts of the Pacific Ocean along the coasts of Japan and
the Kuril Islands.

Food. Pelagic schooling fish and cephalopods constitute the main
food. Food composition varies in different parts of the range. The stom-
ach of six animals of the northern subspecies caught in the waters of Cal-
ifornia and Oregon states contained hake (pike) (Merluccius productus),
jack mackerel (Trachurus symmetricus), and squids (Loligo opalescens);
the stomach of four animals from British Columbia contained herring,
while capelin (Mallotus villosus) was found in the stomach of two ani-
mals from the Gulf of Alaska. Lantern fish (few species) are of overriding
importance in the waters of Japan and squids and other species of fish
are of lesser importance (Wilke and Nicholson, 1958). In the Bering Sea,
along the islands of the western part of the Aleutiian range and on the
coasts of Kamchatka, squids predominate in the food (Fig. 321) while
smaller fish and shrimps are consumed to a lesser extent. In most of
the cases examined, various species of animals were found together in
the stomach of porpoises (Mizue, Joshida, and Takemura, 1966). The
species of cephalopods and fish found in the stomachs have not been

- Stated.

The food composition of northern and southern Dall porpoises
compiled from specimens collected in the waters of Japan from March
through June is compared in Table 50.

Fishes of the family Sudidae are considered very rare and only rare
specimens are found in ichthyological collections; nevertheless common
porpoises apparently feed on them quite regularly. The stomach of
the porpoises caught in the Sea of Japan contained saury and squids
(Sleptsov, 1955).

Behavior. Dall porpoises are most often encountered in small groups
of 2 to 18 head but herds of 20 to 25 animals have been encountered

16 Hall and Kelson (1959) do not recognize this form and regard Long Beach and Los
Angeles at 34° N lat. as the southern point of this range.
According to the data of A.G. Tomilin (1957), our northern form covers the Sea of
Okhotsk and Bering Sea as also the Sea of Japan. His more recent references correspond
to those cited in the text. (V.H.)

555

745

Fig. 321. Squids in the stomach of a Dall porpoise. Pacific Ocean, east of Honshu
Island, 1960 (photograph by G.M. Kosygin).

in the fore-Kuril waters (Fig. 322). Sometimes herds of these dolphins
contain 100 or more animals.

While in motion, these porpoises breach the water often and sharply
so, flying in the air over quite a long distance. On finding themselves in
the vicinity of a ship, they run rapidly ahead of its bow, quickly changing
from side to side, or swim along the board, easily overtaking the ship.
The speed of these porpoises exceeds 20 km/hr.

These porpoises are sighted close to the coasts as also far away from
them in deepwater regions. The dolphins studied off the coasts of North
America were caught at depths of 180 m and off the coasts of Japan at
depths of 2,700 m.

Migrations. Information on migrations is extremely fragmentary. In
the northern parts of their geographic range, especially at the places of
ice cover, these dolphins move from south to north and back, spending
the winter season outside the ice-covered regions. Apparently, compar-
atively small seasonal migrations are a feature of these porpoises in the
other parts of their range also.

554

746

Table 50. Food of Dall porpoises (Wilke and Nicholson, 1958)

Food item Southern subspecies Northern subspecies
(86 animals) (7 animals)
Number of % by _ Number of % by
stomachs vol. stomachs vol.
Fishes
Lantern fishes:
Notoscopelus sp. 27 21 2 37
Diaphus sp. 6 2 — —
Taletonbeania taylori 5 1 3 11
Lampanychtus sp. 1 Traces — =
Unestablished species 44 49 3 35
Sudidae—Paralepis sp. 7 4 2 10
Hake, Laemonema longipes 5 11 — —
Mackerel, Scomber japonicus 2 1 — —
$4 14$
Ommatostrephes sloanei-pacificus 22 5 1 2
Watasenia scintillans 35 6 3 3
Unestablished species 5 Traces 2 Traces

|

An
\
№

<
\
№

С
WS

Fig. 322. Dall porpoise, Phocoenoides dalli, at sea (figure by N.N. Kondakov).

In the Pacific Ocean waters along the Canadian coast, Dall porpoises
were encountered in summer mainly in the bays between the islands
north of Vancouver Island. The animals prefer bays which are wide open
on both sides with characteristic ebb-tide currents.

In the waters of Japan, early in March, Dall porpoises are caught in
the southern part of the range; large herds of these animals are seen in
mid-March in the waters of Iwata Prefecture (about 40° N lat.). Hunting
here continues up to June. Later, most of the porpoises migrate to the

556

747

coasts of Hokkaido and are caught again in autumn along the coasts of
Honshu.

In the coastal waters of the islands of Japan, hunting is carried out
within the 30-mile coastal limit but most of the animals are concentrated
in the summer at a distance of 10 to 15 miles from the coasts (Wilke,
Taniwaki, and Kuroda, 1953).

Reproduction. In May, 1950, embryos weighing over 6 kg were found
in the females of the northern subspecies (Wilke er al., 1953). A fully
formed embryo was found in a female porpoise in the waters of Queen
Charlotte Island in November, 1926. It is possible that the period of
parturition is greatly extended among these porpoises. South of Queen
Charlotte Island, newborn calves were often sighted in August and not
even once encountered before August 7.

A large number of animals was studied over several years in the
southwestern part of the Bering Sea from the second half of May to
early August. Fully mature testes were not seen in the male porpoises
during this period. Evidently, their mating occurs later, probably not
earlier than August end. This is also supported by the results of studying
the female reproductive systems: corpora lutea were not detected in the
ovaries of females.

The period of parturition has tentatively been placed in the second
half of July to mid-August. Gestation among animals of this species thus
extends for not less than a year. Among all the pregnant females, without
exception, embryos lay exclusively in the left horn of the uterus; the state
of the left horn indicated that the embryo had formed in it and the corpus
luteum was present in the left ovary. Graafian follicles in the ovaries of
all the nonproducing females, without exception, had developed in the
left ovary while the right one appeared to be immature. Among the
embryos, no difference was detected between the left and right ovaries
but in a newborn calf the left ovary was already larger than the right
one.

Based on a small number of samples, it was tentatively determined
that the body length of a calf at birth is 85-90 cm. Most females become
mature after two full years and a body length of about 170 cm. Males
achieve sexual maturity at the age of three years or even older and a body
length roughly of 185 cm. The ratio between the males and females in
the population is close to 1:1. Of the 54 animals studied, 26 were females
and 28 males (Mizue and Joshida, 1965; Mizue, Joshida, and Takemura,
1966).

A slightly different distribution of animals of different sexes has been
noticed among the southern subspecies. As established from the results

557

556

748

of spring hunting (from March through May), males remain in the south-
ern and females in the northern half of their range (Wilke et al., 1953).

Enemies, diseases, parasites, mortality, and competitors. The nema-
tode Halocerus (Prohalocerus) kirbyi Dougherty, not known among other
marine mammals, has been detected in the lungs of animals from San
Franciscan waters. Н. dalli and Irukanema аа are also known. In the
stomach of a lone dissected animal from waters of the Kuril Islands,
an immature Anisakis sp. was detected (Delamure, 1955; A. Skrjabin,
1960).

Field diagnosis: Sharply demarcated large white patches (“‘wings”’) on
the sides and a white smear on the dorsal fin are characteristic of this
dolphin (Fig. 323). (V.A.)

Economic Importance

This species is of limited hunting importance only in Japan where up
to a thousand or even more porpoises are caught each year. No catches
have been reported from other parts of the range.

Dall porpoises are hunted along with other species of dolphins and
small whales, using small craft (20-30 tons) with a crew of 10-12 per-
sons. These vessels are used in spring to catch these porpoises and sharks
and swordfish in summer; these vessels are again used in spring to catch

Fig. 323. Dall porpoise, Sea of Japan, 1965 (photograph by А.Е. Kuzin).

749

porpoises. These ships cruise at 7-10 miles/hr. Shooting with guns and
harpooning the killed animals manually is done from a special wooden
platform constructed on the bow. From time to time, when the por-
poises approach the vessel quite close, they can be harpooned without
using firearms. In most cases, the killed porpoise floats on the water
surface for a few minutes in a vertical position with the head above the
water. Killed porpoises have even been found floating a day or two days
after shooting. Nevertheless, a large number of killed animals escape the
hunter’s bag. Thus, in 1950, about one-third of the killed Dall porpoises
were lost.

With the growing salmon fisheries in the open sea, Japanese fisher-
men have begun catching a large number of Dall porpoises every year.
The regions of porpoise catching have been identified as regions in which
the salmon are caught using gill nets. This catch is practiced in the north-
western part of the Pacific Ocean and in the southwestern part of the
Bering Sea, including coastal waters of the Aleutian island range and
Kamchatka coasts. Porpoises fall into salmon nets in coastal waters more
often than in the open sea. It has been assumed that porpoises are also
abundant east of this region of salmon catching, where salmon are not
fished. Porpoises probably become entangled in the nets when they feed
on the fish caught in them. However, an analysis of their stomach con-
tents did not reveal a significant number of salmon. In some years, some
tens and even more than a thousand Dall porpoises fall into salmon nets,
but until recently these porpoises were not utilized and fishermen threw
them overboard (Mizue and Joshida, 1965).

The output of Japanese porpoise hunting is disposed mainly in the
market. The carcass is prepared for sale as follows. The flippers dorsal
fin and caudal flukes are dismembered. Two longitudinal sections are
made through the skin (along the dorsum and belly) from the tip of
the snout to the tail. Later, blubber with the skin is separated from the
trunk in two pieces. The flesh is separated from the vertebral column,
also in two pieces, the ribs cut out, and the abdominal cavity cleared by
removing the intestine, heart, liver, and kidneys, which are sold as edible
products. The skin is tanned but the product is of rather poor quality. Oil
is rendered from the blubber; fat from the head and jaw regions serves
as raw material for producing excellent machine oil. Skeletal bones and
all other remnants are processed into a fertilizer meal. The flesh is quite
often used as food.

Females weigh 91 - 115 kg (М 95 kg) and males 84-118 kg (М 102 kg)
(McTaggart, 1944; Wilke, Taniwaki, and Kuroda, 1953; Klumov, 1959).
(V.A.)

558

750

Genus of Black Finless Porpoises
Genus Neophocaena Palmer, 1899

1846. Neomeris. Gray. Zoology. Voyage Erebus and Terror, I, Mamm.,
p. 30. Delphinus phocaenoides G. Cuvier. Nom. praeocc. (Neomeris
Costa, 1844, from the group Vermes).

1847. Meomeris. Gray. List Osteol. Spec. Brit. Mus., pp. XII, 36. Not
important from nomenclatural viewpoint.

1899. Neophocaena. Palmer. Proc. Biol. Soc. Washington, 13, p. 23. Sub-
stituted for preoccupied Neomeris Gray, 1846. (V.H.)

Maximum body length, up to 190 cm.

The head is rounded. No “beak” is evident. A dorsal fin is lacking.
A band of fine horny tubercles runs along the middorsal line.

The body is lead-black but the abdomen light-colored.

The rostrum of the skull is rounded anteriorly, broad, and short
(much shorter than the cranium). The premaxillae are broad and not
pinched anteriorly. The small pterygoid bones are widely separated. The
teeth have broadened and flattened crowns and number #=12. Vertebrae
60-63. Phalangeal formula: 15, II,;-7, III,;_;, [V3, and V>. Ribs, usually
in 14 pairs.

Almost nothing is known about the biology of these animals. They
feed on bottom-dwelling crustaceans, fish, and cephalopods.

They inhabit the warm waters of the Indian Ocean and the western
part of the Pacific Ocean. They usually live near the coasts and transgress
into rivers. They have been reported from the Far Eastern waters of the
USSR but none have been caught.

The genus comprises a single species, N. phocoenoides G. Cuvier,
1829. (V.S.)

BLACK FINLESS PORPOISE
Neophocaena phocaenoides G. Cuvier, 1829

1829. Delphinus phocoenoides. G. Cuvier. Régne animale, I, p. 291. Cape
of Good Hope, South Africa. (V.H.)

Diagnosis
Only species of the genus.
Description

Similar to the other common porpoises in outer appearance. The body
build is compact. The trunk is somewhat stunted and thickened in the
anterior portion. The head is relatively short (Fig. 324). The adipose

559

ПЭ

Fig. 324. Black finless porpoise, Neophocaena phocaenoides (figure by М.М. Kondakov).

body (corpus adiposum) [melon] on the head is highly developed and
projects forward in the frontal part. The mouth section is small and its
corners are upturned. The flippers are quite broad and crescent-shaped.
Along the middorsum, from the cervical section to the anus, a band of
horny tubercles, 3-6 cm wide, occurs, which broadens slightly anteriorly
and narrows posteriorly. Much larger tubercles (up to 0.5 mm high and
2-2.5 mm in diameter) occur along the margins of the band. In the
embryos, these tubercles are disposed one each on quadrangular plates
closely adjoining each other. The upper lip of the embryo sports eight
whiskers (four on each side).

The lead-black color on the upper side of the body gradually light-
ens notably on the abdomen. A gray patch is visible on the thorax and
abdomen. The flippers and caudal flukes sometimes have light-colored
sections of varying size, Cervical vertebrae 7, thoracic 13-14, lumbar
11-14, and caudal 28-31. The first five cervical vertebrae are fused.

The main body measurements of an adult female black finless por-
poise (Tomilin, 1957) are (in cm): body length 124; distance from tip of
snout to anal opening 8.5, to base of flippers 23, to posterior margin of
blowhole 9; width of caudal flukes from corner to corner 43; maximum
length of flippers 26 and maximum width 8.

The measurements of the skull (Fig. 325) of a male black finless
porpoise with a body length of 116 cm are (Tomilin, 1957) (in cm):
condylobasal length 19, zygomatic width 12, length of rostrum 7, width
of rostrum at base 6, length of lower jaw 13, and length of mandibular
symphysis 1.5.

In the skeleton of one male (length of vertebral column 85 cm)
there were 7 cervical vertebrae, 12 thoracic, 14 lumbar, and 25 caudal;
the cervical section constituted 4.5%, thoracic 27.6%, lumbar 37.2%, and
caudal 30.7% of the vertebral length (Tomilin, 1957) (it is possible that
some caudal vertebrae were lost). (V.S.)

Geographic Distribution

Warm waters of the western part of the Pacific Ocean and the Indian
Ocean.

752

Fig. 325. Skull of the black finless porpoise, Neophocaena phocaenoides (figure
by N.N. Kondakov).

Geographic Range in the USSR (Fig. 326)

Pacific Ocean waters of the southern Kuril Islands; transgressions are
possible into the waters of the Sea of Japan.!’

Geographic Range outside the USSR (Fig. 327)

Waters of Japan, Korean peninsula, China (South China Sea), Kaliman-
tan and other islands, and also Malaccan peninsula. It is known in the
Indian Ocean in the Bay of Bengal and the Persian Gulf, the Arabian Sea,
and along the coasts of Africa up to the Cape of Good Hope (Sleptsov,
1961; Tomilin, 1962). (V.A.)

Geographic Variation

Not established.
Biology

Information on biology is extremely scant. Crustaceans (macrura group:
Penaeus, Palaemon, and especially P. japonicus), cephalopods (Loligo
sp. and Sepia sp.), and some species of fish serve as food objects. They
consume predominantly benthic animals and hence the black finless por-
poise can be regarded as a coastal form. According to observations made

17 Once, in 1951, two porpoises were encountered 20 to 25 miles off Shpanberg Island
in the southern Kuril Island range (Sleptsov, 1952). This species has not been caught in
our waters.

561

753

Fig. 326. Range of the black finless porpoise, Neophocaena phocaenoides, in the
USSR (V.A. Arsen’ev).

on the Kuril Islands (Sleptsov, 1961), it is encountered in open waters,
mainly in small groups (five or six animals) and singly. Sometimes the
animals gather far away from the coasts into very large herds consisting
of several tens of animals. Quite often, they transgress into rivers in
which they spend much time, ascending hundreds of kilometers upstream
(along the Yangtze up to 1,800 km; also sighted in Lake Dongting Hu).
Their movements are gentle and they rarely breach the water.

Migrations have not been studied. It has been assumed that parturi-
tion occurs in October since a female 119 cm long with an embryo 52 cm
long was encountered in this month.

Seven species of helminths have been registered among black fin-
less porposes: trematodes three, cestodes one, and nematodes three. The
trematodes Campula folium Ozaki, Orthospianchus elongatus Ozaki, and

560

754

Nasitrema spathulatum Ozaki were detected in the waters of Japan and
are known only in the black finless porpoise. The first parasitizes the
liver, the second the intestine, and the third the nasal cavity. The ces-
tode, Diphyllobothrium fuhrmanni Hsu, detected in waters of Japan and
China and parasitizing the small intestine, in addition to the black finless
porpoise, has been found in spotted dolphins. All the three species of
nematodes were found only in the black finless porpoise from the waters
of China. Halocercus pingi Wu localizes in the lungs, Sternus auditivus
Hst and Hoeppli in the auditory organs, and Onchocerca fulleborni Hoep-
pli and Hsu in the musculature. Most of the species of helminths found
in the black finless porpoise have not been detected in other marine
mammals.

The black finless porpoise has no economic importance whatsoever;
it is not hunted except for a few stray animals caught in the rivers
of China (Delamure, 1955; Sleptsov, 1955, 1961; Tomilin, 1957, 1962).
(V.A.) ’

Fig. 327. Species range of the black finless porpoise, Neophocaena phocaenoides
(V.A. Arsen’ev).

562

755

Family of Narwhals
Family MONODONTIDAE Gray, 1821

Cetaceans of medium dimensions, with body length up to 610 cm.

A dorsal fin is absent although a long low fold occurs on the dorsum.
The head is rounded, small, without a beak, and demarcated from the
trunk by a distinct neck. The flippers are short and broad.

Body coloration is subject to age-related variations: it is dark in
young animals, turning light with advancing age.

The rostrum is relatively short and broad. The frontal and interpari-
etal bones form a crest. The petrous temporal bone grows toward the
skull. The broad pterygoids articulate with the squamosals. Teeth ум.
Vertebrae 49-55. The first and second cervical vertebrae are not fused
whiie the rest may be fused in varying groups. The ulna has.no olecranon
process.

The geographic range of these animals is restricted to the waters of
the northern polar region (Fig. 328).

The family comprises two genera: (1) belugas or white whales, Del-
phinapterus Lacépéde and (2) narwhals or unicorns, Monodon Linnaeus.
Belugas are of economic importance. (V.S.)

Genus of Belugas or White Whales
Genus Delphinapterus Lacépéde, 1804

1804. Delphinapterus. Lacépéde. Hist. Nat. Cétacées, р. XLI, 243. Del-
phinapterus beluga Lacépéde = Delphinus leucas Pallas, 1766.

1815. Beluga. Rafinesque. Anal. Nat., p. 60. Substituted for Delphi-
napterus Lacépéde, 1804. (V.H.)

Dimensions are the smallest in the subfamily, with a body length up to

6 m.

The body is markedly elongated. The relatively small head has a very
prominent frontal projection. The monochromatic body is dark in the
young, lightening to white (ivory) or yellow in the adult. The epidermis
is remarkably well developed.

The relatively narrow skull is flattened dorsoventrally. The rostrum,
broad at the base, is roughly 1.5 times longer than the cranium. The
maxillae dorsally adjoin the bony nares and extend far backward, almost
up to the occipital bones. The pterygoids are well developed. Powerful
crests are typical. Teeth 5-1, often irregular in form, and sometimes with
supplementary cusps. Vertebrae 49-54. Some of the cervical vertebrae
are fused. Of the 11-12 pairs of ribs, 5-7 pairs are fused to the sternum.

756

Fig. 328. Range of the narwhal family, Monodontidae (V.A. Arsen’ev).

561

563

ПЭ

The sternum is either non-segmented or consists of two or three sections
(sometimes even six). Phalangeal formula: I,_4, Пв-э, Шб-в› [V5-7, and
V,-.- Digits III and IV are almost equal in length. The number of pha-
langes of some digits in embryos is reduced in adult animals. Sometimes
the digits are split and resemble claws.

Belugas are ichthyophagous; they feed mainly in the coastal zone of
the sea. Gestation extends for 11-12 months. Females calve annually.

These animals are distributed in the seas of the North Atlantic, Arc-
tic, and North Pacific oceans.

Fossil remains have been traced in the Pleistocene of North America.
The genus comprises a single species: the beluga, D. /eucas Pallas, 1776.

These animals are of commercial importance. (V.S.)

BELUGA OR WHITE WHALE!’
Delphinapterus leucas Pallas, 1776

1776. Delphinus leucas. Pallas. Reise durch versch. Prov. Russ. Reiches,
3 (I), p. 497. Ob’ estuary.

1776. Delphinus albicans. Muller, Zool. Danicae prodromus, p. 7. Green-
land.

1804. Delphinapterus beluga. Lacépéde. Hist. Nat. Cétacées, pp. XLI,
243. Davis Strait.

1935. Delphinapterus dorofeevi. Klumov and Barabasch. Zh. “Rybnoe
Khozyaistvo SSSR,” No. 11. Sakhalin Bay, Sea of Okhotsk.

1935. Delphinapterus freimani. Klumov. [bid., no. 7. Dvina Bay, White
Sea.

1935.- Delphinapterus leucas maris-albi. Ostroumov. Zh. Za Rybnuyu
Industriyu Severa, no. 11. Onega Bay, White Sea. (V.H.)

Diagnosis
Only species of the genus.
Description

Age-related changes of body proportions are known among belugas
(Arsen’ev, 1936a; Vladykov, 1943, 1944; Kleinenberg et al., 1964). The
changes pertain to an increase in ratio of body length to radius of the
trunk and relative (to body length) reduction of length of the flippers

18 The local Russian people (coastal population) call the animal “beluga”. “Belukha”
represents an artificial (bookish) modification of “beluga” to distinguish the whale from
“beluga fish” (giant sturgeon) (Heptner, 1930; Chapskii, 1937).

564

758

as well as narrowing of the caudal flukes. The form of the flippers
of belugas is typical: their proximal margin is flexed, the flexure very
distinctly manifest in old animals (Vladykov, 1943). Such a form of the
flippers (Fig. 329) is possibly important for braking during sharp turns
(Kleinenberg et al., 1964).

The absence of a dorsal fin is explained by the adaptation of the
animal to living amidst ice (the fin could interfere in floating in such
conditions; Heptner, 1930) and, further, enables the animal to bend along
the longitudinal axis of the body, which is important in catching fish
(Yablokov, 1959). A rather low leathery crest replaces the missing dorsal
fin in some animals.

The epidermis (up to 7-11 mm) and the horny layer [stratum
corneum] (up to 1-6 mm) are highly thickened; the dermal layer is
very well developed while the subcutaneous adipose tissue is relatively
thicker compared to other whales (Bel’kovich, 1959; Kleinenberg et al.,
1964).

Age-related changes of body color are significant. The light gray
color of the newborn changes after a few days or a week into almost black
in suckling calves, but lightens gradually thereafter (Bel’kovich, 1959).
Whitening is more rapid on the dorsum, body flanks, and abdomen. How-
ever, the dark-colored edge of the caudal flukes undergoes no depigmen-
tation. The body surface of most of the animals is covered in numerous
scars, spots, and stripes.

The white color of the belugas is considered a protective adap-
tation (Kukenthal, 1900). It has also been suggested that such a col-
oration frightens schools of fish, which facilitates their catch by the whale
(Yablokov, 1956). Another view (obviously erroneous) holds that the
white coloration reduces heat dissipation from the body of the animal
(Chapskii, 1941). Four color groups of belugas (Fig. 330), corresponding

en
Meee typ pple et

и,
ИЛИ,
77 277:

й

Fig. 329. Beluga, Delphinapterus leucas (figure Бу М.М. Kondakov).

564

159

to definite age groups (Dorofeev and Klumov, 1936), are usually recog-
nized (for more details, see p. 782). However, it has been opined that
the variations in coloration of the beluga body should not be taken as an
accurate criterion for determining the age of these animals (Bel’kovich,
1959).

The simple peg-shaped teeth of belugas are characterized by a typical
cutting and supporting form developed as a consequence of the wearing
down of the teeth (Yablokov, 1958a, 1959). The maximum length of the
teeth in the upper jaw is 5.8 cm and thickness 1.2 cm; in the lower jaw,
5.0 cm and 1.8 cm respectively (Yablokov, 1959). The largest teeth occur
in the anterior part of the upper jaw and midpart of the lower jaw. Teeth
are usually lacking on the premaxillae. Teeth are cut in year-old calves.
Cervical vertebrae 7, thoracic 11-12, lumbar 6-12, and caudal 21-26.

The anterior portion of the digestive tract of the beluga, right up to
the second chamber of the stomach, is covered with Keratinized stratified
epithelium in which (up to the first chamber of the stomach inclusive)
a network of tonofibrillae, reinforcing the epithelium, is well developed.
Such adaptations help the animal to swallow a large quarry whole without
mastication (Kleinenberg, Yablokov and Tarasevich, 1958). The stomach
consists of five chambers. The intestine, length varying from 22.5 to
37 m, is not divided distinctly into a small and large intestine due to the
absence of a cecum. The duodenum is highly developed; the last third of
the rectum is covered by stratified epithelium forming low longitudinal
nonexpanding folds, with a powerful muscular cover supplemented by a
powerful anal sphincter.

Sexual dimorphism is manifest in the dimensions of the animals;
males are larger than females. Moreover, the ratio of body length to
radius of its cross section is more in males than females, underscoring
the excellent hydrodynamic form of the male body; the caudal flukes
of males are relatively larger while the flippers, on the contrary, are

Fig. 330. Age-related color groups of the beluga, Delphinapterus leucas (figure
by N.N. Kondakov).

565

564

760

relatively smaller. In males, the number of teeth is slightly more than
in females (Yablokov, 1959). The frontal prominence in most males is
more powerfully developed (this is particularly true of aged males).

The maximum length of belugas in the Far East is 600 cm among
males and 500 cm among females; in the Kara Sea 472 and 413 cm; Gulf
‘of St. Lawrence 447 and 409 cm; western Greenland 572 and 474 ст;
Hudson Bay 448 and 400 cm; and the Beaufort Sea 460 and 383 cm
(Kleinenberg et al. 1964).

The main body measurements of Kara Sea belugas (Tomilin, 1957)
(average for 13-14 males and 11 females) are respectively (in cm): body
length 410 and 365; length of head 52 and 46; distance from tip of snout
to base of flippers 30 and 71; distance from anal opening to fork between
caudal flukes 94 and 89; length of flippers 44 and 39, width of flippers
31 and 26; and width of caudal flukes (from tip to tip) 88 and 77.

The main measurements of the skull (Fig. 331) of belugas from the
White and Kara seas and the Far East (Tomilin, 1957) respectively aver-
age (in cm): condylobasal length 52 (10 measurements), 57 (10), and 59
(10); zygomatic width 28 (8), 30 (10), and 32 (4); length of rostrum 25
(10), 28 (10), and 29 (4); width of rostrum at base 17 (9), 18 (10), and
19 (4); and mandibular length 42 (4), 42 (8), and 44 (9). (V.S.)

Fig. 331. Skull of the beluga, Delphinapterus leucas (figure by N.N. Kondakov).

761

Geographic Distribution

Predominantly the cold waters of the Northern hemisphere where it is
very widely distributed almost all around the polar region.

Geographic Range in the USSR

Apparently no changes have taken place in the range over the historic
past. In the Barents Sea, beluga is quite regularly encountered along
the Murman coast (mainly in winter); it is sighted almost year-round at
Kanin and Kolguyev, and in the Chesha Bay. It is quite common in the
Pechora Sea, Vaigach, and along the west coast of Novaya Zemlya. At
the end of summer, it is not infrequent around Franz Josef Land. It is
most numerous probably in the southeastern corner of the Barents Sea
where is is sighted throughout the year. It lives mainly along coasts and
has not been sighted away from them. There is no doubt of its absence
in the open Barents Sea (Fig. 332).

The beluga inhabits the entire White Sea. As an exception, it trans-
gresses into the Baltic Sea where it has been sighted in our waters, espe-
cially in the Riga (Parnu) and Finland gulfs. In the Kara Sea, it pre-
dominantly covers the southwestern and western parts. It is encountered
along the east coast of Novaya Zemlya right up to Cape Zhelaniya, at
Vaigach, in Vaidarask and Gulf of Ob in Yenisey Gulf and also along
the coast east up to Vil’kitskiy Strait. The beluga inhabits the Kara Sea
mainly in summer. It can be encountered in autumn in its northern parts.
At some places (Gulf of Ob) small herds are sometimes encountered in
winter.

In the Laptev Sea, the beluga is seen in the summer months along
much of the coast from Severnaya Zemlya islands to Novosibirsk. It is
most common in the southwestern part of the sea (Pronchishcheva Bay,
Begichev Island, etc.), and also in the Lena estuary. From the Novosibirsk
islands, the beluga is seen in the waters of the East Siberian Sea. In the
winter months it is probably confined to the middle and northern parts
of the Laptev Sea.

It inhabits the Chukchi Sea predominantly in the summer months.
Distribution along the Chukchi peninsula is restricted to the coastal
section from Cape Dezhnev to Cape Shmidt but sometimes transgresses
into the waters of the East Siberian Sea. Under favorable ice conditions,
it moves far north. It is quite regularly found close to Wrangel Island
and probably reaches 74 to 75° N lat. It is encountered in Bering Strait.

In the Bering Sea, it inhabits the Gulf of Anadyr in which it is
most numerous close to the Anadyr estuary. Its distribution southward

762

(2
es

у <
Ly
и (.
2,
—' aaa” ES

a:

OL 09
a x
>> ©
г 5
Se Males
) Wemeg 23

a
<

567

763

is limited to Cape Navarin or only slightly farther. Distribution in the
open waters of the northern part of the Bering Sea has not been studied.

It is found almost all along the coast of the Sea of Okhotsk. Depend-
ing on the period of the year, it is common along the Sakhalin coasts (east
coast, Terpeniya Bay, La Perouse and Tatar Straits, Amur liman, and
the Gulf of Sakhalin). In the southwestern part of the Sea of Okhotsk,
it ranges from the Amur estuary to Uda and Ayan bays in the west. It
is common along the northern coasts of the Sea of Okhotsk, including
Shelikhov Gulf, and the west coast of Kamchatka, mainly its northern
part. It is rare along the west coast of the Sea of Okhotsk from Ayan Bay
to Okhotsk. It has not been reported in the waters of the Kuril Islands.

Geographic Range outside the USSR

In the basin of the Atlantic Ocean, it is widely distributed in the Cana-
dian Arctic archipelago (Fig. 333). It is encountered in the Gulf of
St. Lawrence, along the Labrador coasts, in Ungava Bay, Hudson Bay,
Hudson Strait, Foxe Basin, Davis Strait, along the coasts of Greenland
and Baffin Island, and in Baffin Bay where it reaches north of Smith
Strait. It is encountered in the bays of Lancaster, Jones, and Barrow,
and in the Beaufort Sea. It has been sighted in McClure Strait, Amund-
sen Gulf, and at many points along the coast in the region of Mackenzie
Island. The beluga is also encountered east of the Canadian coast close
to Iceland, Jan Mayen Island, Spitsbergen, and Finmark. Its distribution
along the east coast of Greenland has not been clearly established.

In the Bering Sea, it is known along the coasts of Alaska from Bristol

_ Bay to Bering Strait and farther, in the Chukchi Sea up to Cape Bar-

568

row on the northern coast of Alaska. It is also encountered east of this
point (Arsen’ev, 1939; Vladykov, 1944; Tomilin, 1957; Sergeant, 1962a;
Kleinenberg, Yablokov, Bel’kovich, and Tarasevich, 1964). (V.A.).

Geographic Variation

The monotypic state of the genus Delphinapterus has been reviewed time
and again but the new species described are not usually recognized while
the probability of some subspecies very recently described has invited
serious dispute. Preliminarily, the existence of three subspecies of belugas
may be recognized in our waters:

1. White Sea beluga, D. | maris-albi Ostroumov, 1935. Smallest of the
forms; adults 253-376 cm (x 312).

This subspecies is found in the waters of the White Sea and the
southern part of the Barents Sea.

Not reported outside the USSR:

‘(дэ иэзту У ‘A) злэлы ш зе8эа Jo зриц Jo says ay} 1иэзэл4эл $10 502721 smuaidnuiydjag ‘e8njaq эцз JO э8иел $10946 ‘65$ ‘8 195

765

2. Kara Sea beluga, D. /. leucas Pallas, 1776. Occupies an intermediate
position among the three subspecies in body dimensions. Adults vary in
length from 318 to 464 cm (х 390).
This subspecies is found in the Barents, Kara, and Laptev seas.
Outside the USSR, it apparently lives in the remaining waters of the
North Atlantic.

3. Far Eastern beluga, D. /. dorofeevi Klumov and Barabasch, 1935.

Largest of the subspecies. Adults vary in length from 320 to 600 cm
(х 424).

This subspecies is found in the Okhotsk, Bering, and Chukchi seas.

Outside the USSR, it lives in the American waters of the aforesaid
seas.

A systematic analysis of belugas from the western Atlantic and east-
ern Pacific Oceans in relation to the above forms has not been attempted
and the classification is tentative. It is possible that the populations in
some of our Far Eastern water bodies are not wholly identical from the
viewpoint of systematics.

Each of the subspecies occupies an immense range and it is not
unlikely that many local populations exist within these ranges. Proba-
bly, belugas of the Laptev Sea form a distinct local population. Belugas
from the Okhotsk and Bering seas definitely represent isolated popula-
tions, distinctly separated from each other by a large distance. Within
the Okhotsk population per se, two relatively isolated herds have been
postulated (see “Seasonal Migrations and Transgressions”’).

Belugas inhabiting waters outside the limits of the USSR also
probably form several local populations. One such occupies the
zone Iceland—Greenland—Spitsbergen. In Canadian waters, three such
populations can be recognized: in the Gulf of St. Lawrence, in the
eastern Canadian Arctic, and in the western Canadian Arctic or the
Beaufort Sea. The first of these, even if it mixes at all with the eastern
Arctic population, does so very minimally. The two arctic populations
of America are separated by 20° along the longitude in the zone of
Melville Strait (Sergeant, 1962a). However, there is as yet no adequate
information to resolve this problem. (V.A.)

Biology

Population. The beluga may be regarded as a moderately populous
species. According to approximate calculations, the population of White
Sea beluga comprises 8,000 - 10,000 and of the Kara Sea 40,000 - 50,000
animals (Klumov, 1939). No less than several tens of thousands of
animals live in the Sea of Okhotsk but the Bering Sea population is

766

probably somewhat less. Information is not available for judging the
beluga population in other parts of the range within USSR waters. The
commercial utilization of all these populations is so insignificant that
hunting has no bearing whatsoever on the natural population dynamics.

Food. Fish and crustaceans serve as food objects. The extensive range
occupied by the beluga provides quite а large number of species of food
objects which differ in various parts of the range. However, fish every-

571 where occupy a predominant position. The large number of organisms
found in the stomachs contain some which enter incidentally or represent
the food of animals on which the beluga feeds (Table 51).

Fish (pelagic and benthic) which form massive schools and bottom-
dwelling crustaceans constitute the main food. The least number of empty
stomachs among the belugas of the Sea of Okhotsk is seen during the
massive arrival of salmon (Arsen’ev, 1939); in the northern seas, the
beluga feeds most intensely in the period of arrival of arctic cod (Klumov,
1936a; Tarasevich, 1960a, 1960b).

Based on the frequency of encounter and the number of animals
found in the stomach of belugas, their main food items are: in the
White Sea—herring, capelin, and shrimps; in the Kara Sea—arctic cod,
cisco, and other whitefish; in the Sea of Okhotsk—salmon (chum salmon
and humpback salmon), navaga, herring, and isopods; in the Gulf of
St. Lawrence—capelin, sand eel, cephalopods, and Nereis; and in Hudson
Bay—capelin, cephalopods, and Nereis. Arctic char (Salvelinus alpinus;
Daan and Douglas, 1953) was additionally encountered in the stomach
of belugas of the Canadian Arctic.

A seasonal change of food intake, also having a bearing on the age
of belugas, has been noticed. For the White Sea beluga inhabiting the
waters of Kanin, the main food objects in June are lumpsucker_ and floun-
der, changing to navaga, cod, and shrimps in mid-July, and herring and
flounder at July end (Klumov, 1936a). In the Gulf of Sakhalin of the
Sea of Okhotsk, young belugas at the end of the milk suckling period
consume mainly crustaceans, which constitute up to 50% of their ration,
and relatively small fish. As the animal grows, the importance of crus-
taceans and small fish gives place to large fish (Salmon) whose specific
proportion in the intake increases. Large fish serve as the main food
of adult animals in which crustaceans account for less than 9% of the
intake (Arsen’ev, 1939). Belugas of the Gulf of St. Lawrence also exhibit
differences in food intake relative to the regions, seasons, age, and even
sex groups (Vladykov, 1947).

The food of the beluga has been studied everywhere only in the
summer months. No information is available on its winter sustenance.

569

767

Table 51. Food objects of belugas

White Sea (Klumov, 1936a)

Fish
Atlantic herring, Clupea harengus L.
West Atlantic capelin, Mallotus villosus
Mull.

European smelt, Osmerus eperlanus dentex

Steind
Arctic lamprey, Lampetra japonica
tentrionalis Berg
Atlantic navaga, Eleginus navaga (Pall.)
Lumpsucker, Cyclopterus lumpus (L.)
Atlantic cod, Gadus morrhua L.
Haddock, Melanogrammus aeglefinus
(Hodd.)
Flounders: Pleuronectes flesus Р.
P. platessa L.
Liopsetta glacilis (Pal.)

Crustaceans
Shrimp, Crangon crangon L.

Barents and Kara seas (Klumov, 1936a)

Fish
Arctic cod, Boreogadus saida (Lepech.)
Arctic cisco, Coregonus autumnalis (Pall.)

Siberian cisco, Coregonus sardinella Val.
Muksun, Coregonus muksun (Pall.)

Siberian sturgeon, Acipenser baeri Brandt

Nelma, Stenodus leucichthys nelma Pall.

Pike, Esox lucius (L.)

Siberian powan, Coregonus lavaretus
pidschian (Gmel.)

Atlantic herring, Clupea harengus L.

Smelt, Osmerus sp.

Arctic char, Salvelinus alpinus

Crustaceans
Isopod, Mesidothea entomon
Mesidothea sabini
Gammaridae

Sea of Okhotsk (V.A. Arsen’ev, 1939)

Fish
Chum, Oncorhynchus keta Walb.
Humpback salmon, Oncorhynchus
gorbuscha Walb.
Navaga, Eleginus navaga gracilis Til.
Pacific herring, Clupea harengus
pallasi Val.
Flounder, Pleuronectidae
Rudd, Leuciscus brandti Dyb.

Smelt, Hypomesus sp.

Goby, Myoxocephalus sp.

Pacific eelpout, Zoarces elongatus Kner.

Blenny, Blenniidae

Arctic lamprey, Lampetra japonica
(Martens)

Sakhalin char, Salvelinus leacomaenis
Pall.

Okhotsk whitefish, Coregonus ussuriensis

Berg

Gulf of St. Lawrence (Vladykov, 1947)

Fish
Haddock, Melanogrammus aeglefinus (L.)
Ocean pout, Zoarces anguillaris (Peck)

Blenny, Blenniidae
Snailfish, Neoliparis atlanticus (Mull.)

Capelin, Mallotus villosus Mull.

Sculpin, Myoxocephalus scorpius groen-
landicus (Cuvier and Valenciennes)

Staghorn sculpin, Gymnocanthus tricuspis
(Reinhardt)

American smelt, Osmerus mordax (Mitchill)

Sturgeon, Acipenser oxyrhynchus Mitch.

Atlantic herring, Clupea harengus L.

Sea lamprey, Petromyzon marinus L.

American sand lance, Ammodytes americanus
De Kay

Atlantic tomcod, Microgadus tomcod
(Walbaum)

Contd.

768

Table 51 Continued

Sea of Okhotsk (V.A. Arsen’ev, 1939) Gulf of St. Lawrence (Vladykov, 1947)
Crustaceans Red hake, Urophycis chuss (Walbaum)
Shrimp, Crangon septemspinosa Say Baltic cod, Gadus callaris L.
Isopod, Mesidothea entomon orientalis Greenland cod, Gadus ogac Reinhardt
Gurjanova
Shrenck’s crayfish, Cambaroides Witch flounder, Glyptocephalus cynoglossus
schrenckii Kessler (L.)
570 Hermit crab, Pagurus capilatus Smooth flounder, Liopsetta putnami (Gill)
Mud shrimp, Upogebia issaeffi Balss. Winter flounder, Pseudopleuronectes
americanus (Walbaum)
Cumacea Atlantic lumpfish, Cyclopterus lumpus L.
Macruran crabs Thorny skate, Raja radiata Donovan
Smooth skate, Raja senta Garman
Mollusks Skate, Raja sp.
Mytilus edulis Linné Atlantic salmon, Salmo salar L.
Lamellibranchiata Crustaceans
Rhizopoda Shrimp, Pandalus montagui
Nemertini Scud, amphipods
Argis
Copepods
Cumacea
Schizopodes
Cephalopods

Bathypolypus obesus (Verrilli)
Atlantic shortfin squid, Шех illecebrosus
(Le Sueur)

Gastropods
Whelk, Buccinum undatum Linné
Periwinkle, Littorina sp. (?)

Bivalves
Crenella sp. (?)
Cystodaria silgua Spengler
Macoma baltica Linné
Mesodesma arctata Conrad
Mesodesma deaurata Turton
Mesodesma sp. (?)
Mya sp. (?)
Yoldia limatula Say

Polychaeta
Cistendes gouldii Verrill
Nereis virens Sars

572

769

Daily activity and behavior. The beluga is а typical herding animal,
found usually in groups of various strengths. Among numerous obser-
vations, lone animals were encountered in 16.1% of cases (Bel’kovich,
1960). A herd of belugas irrespective of its strength, invariably con-
sists of small groups (Fig. 334) of two to four or five to eight animals
(Golenchenko, 1949a). The age and sex composition of the herd can
vary while the males within a herd often form independent groups. In
other cases, mixed groups are formed in which young ones can also be
seen along with mature animals. However, immature animals never form
independent groups. In the period of reproduction, groups are noticed
within a herd. These groups consist of three or more belugas, among
which is a female with a suckling calf and one or two adult animals.
The female in such groups probably had given birth only recently and
was preparing for fertilization afresh. At the end of the reproduction
period, adult males quite often segregate themselves from females and
form independent herds.

The herd size varies markedly. The most frequently noticed herds
comprise tens (54.9% cases) or several tens of animals (25.8%). Herds
running into a few hundred animals are rare (3.2%) but herds running
into a few thousands of animals can be seen in the period of massive con-
centrations of fish (salmon in the Far East). Small herds are invariably
formed of animals chasing huge schools of fish. An analysis of hunting
data shows that the male to female ratio in a herd is invariably close to

77. 2

CL 4 NOG

ААА 72
LEE GLB.
Ley a GEL РА,
ZZ wy gilli LE gz

Z РЕНА
=. 2 > у os
CELL ee ety bee LFA “Ss 222222,
ЕН = 2 Е SS РНР,
о р Se Аее. CELA В НК

НА
<> LL: LL
22% р

=
A;
==

ZA

Fig. 334. A group of belugas at sea (figure by N.N. Kondakov).

573

572

770

one but differentiation into age- and sex-related groups is noticeable шо
the large migrating herds. In such cases, the front of the herd consists
exclusively of adult males followed by females with calves and immature
animals (see Table 52).

A varying composition of animals (age- and sex-related) is noticed
in some parts of a large herd not only in the Far Eastern waters, but
also in the European north along Novaya Zemlya and in the White Sea
(Bel’kovich, 1960).

The speed of belugas depends on the state and behavior of the herd.
Feeding animals are confined to a relatively small region, dive in different
directions, often at the same site, and remain under water for a long
time. Such herds move in any direction at a speed of not more than 1
mile/hr. “Migrating” herds follow a distinct direction, are more compact,
swim more rapidly, and dive for a very short duration. A migrating herd
moving undisturbed travels at 2-5 miles/hr but a frightened herd can
gather speed, perhaps up to 10 miles/hr; they, however, cannot sustain
this high speed for long. At high speeds the females and calves begin to
lag behind since, apparently, they cannot cover more than 7, or in an
extreme case, 8 miles/hr. Sleeping belugas have been observed several
times. They lie on the sea surface almost immobile, usually close to the
coasts, passively drifting with the waves and the current (Arsen’ev, 1939;
Bel’kovich, 1960; Kleinenberg et al., 1964).

Table 52. Distribution of belugas in a herd (Gulf of Sakhalin) (Arsen’ev, 1939)

Front of herd (44 belugas)

White Blue Gray Suckling calves

Male Female Total Male Female Total Male Female Total Male Female Total

22 6 28 4 4 8 1 3 4 2 1 3

65.1% 18.6% 9.3% 7.0%

Total: males 29 (67.4%) and females 14 (32.6%).

Rear of herd (23 beluges)

White Blue Gray Suckling calves
Male Female Total Male Female Total Male Female Total Male Female Total

— 6 6 1 6 7 5 4 9 1 — 1

26.1% 30.4% 39.1% 4.4%

Total: males 7 (30.4%) and females 16 (69.6%).

71

Belugas feeding predominantly on pelagic life usually do not sub-
merge deeper than 8-10 m. In any case, a herd surrounded by a sweep
net usually does not escape from the lower edge even when the height
of the net does not exceed 5 m and falls short of reaching the bot-
tom. It is quite possible, however, that should the need arise, the beluga
could submerge to several tens of meters. During feeding, belugas dive
for 3-5 min but can remain submerged up to 15 min. After relatively
long submergences in search of food, they surface three or four times in
a row. Migrating animals surface regularly at 20-40 sec intervals with
inhalation/exhalation extending for about a second while the back of the
animal can be seen above the water for about four seconds.

While surfacing, the first to be seen above the water is the upper
part of the head with the blowhole. The head is then submerged as
the dorsum arces above the sea surface. This semicircle, moving around
its own axis, gradually disappears under water. Caudal flukes are never
displayed. The blow is rapid, like a small bush, and is seen like a white
flash in sunny weather (V.A. Arsen’ev).

The beluga does not avoid ice. In the north and the Far East, at
places of its summer habitation, it is seen on days when the ice is broken
and the animal remains free for the first time between ice chunks. In the
north the beluga is common along the edges of ice and seen quite often
in air holes (in the ice) and open water pools among drifting ice up to 9
to 10 points in density. In such cases, small herds are more common. The
animals can overcome considerable masses of dense, apparently compact
ice, past which they enter the open water in which their food is concen-
trated. Such instances are common in the Shantarsk archipelago in the
Sea of Okhotsk. Instances are known of encountering small groups of
belugas in large air holes among stationary ice in which they sometimes
even winter.

The beluga does not avoid freshened or even fresh water. In the
freshening waters of the estuaries of large rivers, belugas are regularly
seen and are hunted. Almost all over the range, dozens of large and small
rivers are known in which the animals regularly transgress chasing fish. In
most cases, they rise up along the river for several tens of kilometers and
are Seen quite often even at a distance of 100-200 km from the estuary. In
some large rivers, these animals were sighted at a very long distance away
from the estuary: in the Yenisey almost near Podkamennaya Tunguska
at over 800 km, in the Amur at Khabarov at about 1,000 km, in the
Pechora at 900 km, in the Ob’ at 1,500 km, and in the Yukon almost at
1,500 km.

Vision and audition are well developed in the beluga but the former
is effective only at a close distance. The animals can orientate themselves

574

UL

in many Situations through their sense of hearing. They can perceive
various sounds, including those produced in a very wide range by other
belugas. These sounds are in the form of clicks and creaks and various
types of whistles; they also resemble barking, roaring, and gnashing, and
a sound somewhat similar to a trill. The sound frequency varies from a
few kHz to 10,000 and even 20,000 kHz. Sounds play the role of var-
ious signals and are used for echolocation. Hunters well know that if
even one beluga finds an exit from the net surrounding the animals, the
entire herd escapes through that very exit in a very short time. It is also
known that the animals well perceive all the sounds produced not only
in water, but also in air (on the coast or on the ice). Moreover, belugas
can very well orientate themselves to sea-level changes (high and low
tides) and are almost never cast on the coast (Arsen’ev, 1939; Tomilin,
1962; Kleinenberg et al., 1964).

Experiments in raising the beluga in an aquarium have yielded pos-
itive results. A male survived in an aquarium for several years and was
taught during this time many “acts” of a “dolphin circus”. This, together
with the external appeal of this white whale, was greatly appreciated by
numerous visitors to the aquarium.

Seasonal migrations and transgressions. Large regular migrations rep-
resent a characteristic feature of the biology of the beluga. However, the
periods and courses of migration of individual populations have been
studied very schematically and opinions are often contradictory.

White Sea. The beluga is generally seen at May end to early
June after the sea becomes free from ice. In the northern portion, mainly
along the west coasts of Kanin Peninsula, it has been sighted even earlier,
early in April, but such instances are generally rare and the herds few. In
summer the beluga is encountered regularly almost all along the entire
coast of the sea; it is most numerous in the Dvina and Onega bay’s
and along the west coast of Kanin. In August, the number of belugas
begins to decrease and by October most of the herd goes into the Bar-
ents Sea (Klumov, 1936; Provorov, 1957). Some belugas overwinter in
the White Sea, as can be seen mainly in the Kandalaksha and Dvina bays
and also in the air holes in Voronka and Gorlo regions. From the air
individual animals or small groups were sighted in winter among dense
drifting ice in the central part and in Voronka (Khuzin, 1960; Kleinen-
berg et al., 1964). Thus the bulk of the White Sea beluga population
inhabits the White Sea from early spring to late autumn, then enters
the Barents Sea before its total freezing, where, apparently, they winter
in its southeastern section. Spring migrations of animals commence as
soon as the icy environment permits, sometimes even when a large part

575

773

of the sea is packed with dense drifting ice. In summer belugas are dis-
tributed throughout the sea and transgress into the numerous gulfs and
bays (Klumov, 1934; Provorov, 1957).

Barents Sea.Asmall number of belugas is seen throughout the
winter at different places along the Murman coast, which they probably
abandon in March. The beluga remains in the region of Kolguev Island
and the Chesha Bay throughout the year: in winter, in open water pools
and along the edges of ice, moving toward the coasts in summer. In
the region of Vaigach Island and along the west coast of the southern
island of Novaya Zemlya, belugas are most numerous early in winter
but, when the ice recedes, herds of belugas are seen close to Novaya
Zemlya throughout winter and even in spring. Early in summer large-
scale migrations of animals into the Kara Sea are noticed through the
strait as also around the northern extremity of Novaya Zemlya. By mid-
July, belugas have almost disappeared from the western coasts of these
islands. In autumn large herds of belugas are once again seen here, now
moving in westerly and southwesterly directions.

At the end of June herds of belugas are seen in the Straits and
bays of Franz Josef Land archipelago, where they remain until autumn
but disappear again with the formation of an ice cover (Klumov, 1936a;
Kleinenberg et al., 1964).

Thus belugas arriving from the White and Kara seas winter in the
Barents Sea. They live along the edges of drifting ice in the region of
Kolguyev Island—Chesha Bay and also along the western coast of Novaya
Zemlya depending on ice cover conditions. Small individual herds of
belugas move toward the Murman coast. At the beginning of summer,
most of the animals leave the Barents Sea for summer feeding in the
Kara and White seas. Some small herds remain throughout the year in
different regions of the sea (Chesha Bay—Kolguyev Island; southern part
of Novaya Zemlya—Vaigach Island; Franz Josef Land archipelago); for
wintering, they go beyond the edge of ice or inhabit open water pools.

Kara Sea. Herds of belugas are seen in Kara Strait and in Yugorsk
Shar Strait from the middle to the end-of May and even in early June and
travel into Baydarata Bay. At this time the animals enter the Kara Sea
and encircle Cape Zhelaniya (northern tip of Novaya Zemlya). In some
years the periods of movement of belugas vary considerably depending
on the ice situation; further, in some cases, belogas were sighted even in
April. In June, in Amderma region (east of Yugorsk Shar Strait), large
herds of belugas move eastward and some are seen along the southeast-
ern coast of Novaya Zemlya throughout July. Belugas reach the estuary
of the Gulf of Ob early in July. On entering the gulf, the animals gener-
ally travel along the right bank since the prevailing northeastern winds

774

press the ice toward the left bank. Belugas return from the gulf mostly
along the left bank or along the midsection of the bay. The Gulf of
Ob, mainly at the point of its confluence with Tazovsk Inlet, belugas
remain until the formation of an ice cover, i.e., until early November
(Dukhovnyi, 1933, 1934; Chapskii, 1937; Kleinenberg et al., 1964).

In the estuary of Yenisey Gulf, belugas are seen in the first few
days of June (waters of Dickson Island) and even earlier in years of
little snow. In most cases, herds of belugas arrive there from the east,
from Pyasinsk Bay, and sometimes from the side of the open sea. Later,
the animals remain there regularly and enter Yenisey Gulf and some-
times rise high up along the Yenisey River. In the strait, belugas traverse
mainly along the eastern coast. ш’ the September the number of animals
decreases but individual herds are seen here throughout this months.
In the second half of July and in August, belugas are common along
the coasts of Pyasinsk Bay and are sighted quite regularly in August and
September in Vil’kitskiy Strait and along the coasts of Severnaya Zemlya
where en masse arrivals of arctic cod occur at this time (Heptner, 1930;
Zikov, 1934; Klumov, 1936a; Kovalev, 1938; Kleinenberg, Bel’kovich, and
Yablokov, 1960).

The migrations of belugas in the Kara Sea can be sketched as fol-
lows. In spring and early summer herds of belugas enter the Kara Sea
through Novaya Zemlya Strait and around its northern extremity. The
animals moving through the strait are seen in Baydarata Bay, later move
eastward, and reach the Gulf of Ob by mid-July. By this time Matochkin
Shar Strait has become ice-free and animals passing through this strait
are seen along the eastern coasts of Novaya Zemlya from where, per-
haps in part, they move also toward the southern coasts of the Kara Sea.
The animals encircling Novaya Zemlya move eastward along the edges
of drifting ice or along open water pools. Some move toward .Severnaya
Zemlya islands and others reach the mainland coast somewhere north-
east of Pyasinsk Bay. These belugas, moving southwest later, are seen
in Pyasinsk Bay, around Dickson Island, and in Yenisey Gulf; however,
the animals sometimes reach Dickson Island not from the northeast, but
from the north. Later, this group of belugas disperses and the animals
are sighted along the coasts of Taimyr and in Vil’kitskiy Strait. In August
and September and in years of little snow even in October, herds of bel-
ugas depart by the same routes (in a reverse direction) to the wintering
sites. Therefore, the autumn courses of belugas in the estuary of Yenisey
Gulf have a northeasterly direction and in the Gulf of Ob a westerly
direction. Small herds of belugas winter in the Kara Sea, probably reg-
ularly, in the air holes in the Gulf of Ob and Yenisey Gulf, and in the
large air holes among drifting ice in various parts of the Kara Sea.

576

775

The periods of migrations of belugas in the Kara Sea depend ритаг-
ily on the period and concentration of the arctic cod, which represents
the main food of belugas. Many major rivers enter the Kara Sea and bring
with them masses of warm water in spring and summer. These waters,
tich in biogenic elements, promote the growth of phyto- and zooplankton
and thus provide extremely favorable conditions for the habitation of the
arctic cod, in search of which herds of belugas follow. Thus the periods
and magnitude of arrival and distribution of belugas depend to some
extent on the magnitude and distribution of river waters in the Kara Sea
(Tarasevich, 1960a, 1960b).

Laptev Sea. The distribution and migrations of belugas in
this sea are not quite clear. Herds of belugas (sometimes large ones)
have been sighted in Vil’kitskiy Strait, along Cape Chelyuskin, in Pron-
chishcheva Bay, in Khatanga Gulf, in the estuaries of the Anabar and
Olenek rivers, in the Lena delta, near Tiksi and, what is more, belu-
gas have been sighted every day at some of these points. Thus the ani-
mals inhabits the entire southwestern coast of the Laptev Sea and some-
times herds of belugas are seen in July to September along the west
coasts of Lyakhovsk and Kotel’nyi Islands (Klumov, 1936a; Kleinenberg,
Bel’kovich, and Yablokov, 1960).

In all probability, an isolated but comparatively small population of
belugas resides in the Laptev Sea and spends the winter in the large
open water pools and air holes of this shallow water basin. In the sum-
mer months the animals inhabit the southern and western coasts of the
sea including Vil’kitskiy Strait, Severnaya Zemlya coast, and possibly
the adjoining Kara Sea waters. During their habitation in the waters of
Novosibirsk Islands, belugas are sometimes seen in adjoining regions of
the East Siberian Sea. Late in autumn the animals leave the coasts Юг’
the central part of the sea where they winter. ©

Sea of Okhotsk. The maximum collection of belugas
in summer is seen in the western (Gulf of Sakhalin, Amur liman, and
Shantarsk archipelago) and northeastern (Shelikhov Gulf with Penzhina
and Gizhaga bays) parts of the sea. In the northwestern part of the sea,
from Shantarsk Island to the region of Ayan-Okhotsk and along the
western coast of Kamchatka, belugas are encountered in small numbers.
They have not been noticed in the central deep water portion of the
sea. Probably, two local herds of belugas spend the summer in the Sea
of Okhotsk and could be called Amur and Penzhina herds (Arsen’ev,
1939).

The earliest appearance of large beluga herds in the southwestern
part of the Sea of Okhotsk (evidently in April) was recorded in Terpeniya
Bay along the southeastern coast of Sakhalin (Polyakov, 1884). At the

S77

776

end of April to early May, small numbers of animals arrive in Tatar Strait
where they remain on the edges of stationary ice in the northern section
of the strait. Early in May many belugas are sighted in the narrow part
of Tatar Strait (Pogibi-Lazarev) and the animals approach the coasts as
soon as the ice breaks up along Sakhalin. As the strait becomes clear,
belugas gradually move northward into the Amur estuary but are con-
fined to the narrow part of the strait generally up to mid-July, leaving
only after the fish (herring and humpback salmon) disappear there. The
animals then move into the Amur estuary and into Sakhalin Strait.

Concurrently, at May end to early June, large herds of belugas
approach from the southeast to the northern extremity of Sakhalin and,
encircling the island from the north, reach the Gulf of Sakhalin, most
of which at this time is still covered by compact ice. On the Sakhalin
coast of the bay, ice begins to break in the first half of June and herds
of belugas are seen immediately thereafter in the Gulf of Sakhalin and
later in the Amur estuary. These animals live there until late autumn.

Roughly at the same time (or slightly later), belugas are seen in the
bays of the Shantarsk Sea while the sea itself and partly the large gulfs
are still ice-bound. However, the dense drifting ice is no obstacle to the
belugas moving in the upper courses of the bays which are free of ice
(Tugursk, Ulbansk, and so оп).

The migrations of Amur belugas can be sketched as follows. In April-
May, herds of belugas appear in Terpeniya Bay and later a part of this
herd moves through La Perouse Strait into Tatar Strait and feeds in its
northern section on herring and humpback salmon. By mid-July, belugas

‘leave for the Amur estuary and the Gulf of Sakhalin. Almost concur-

rently, at the end of May, most of the herds approach the northern
extremity of Sakhalin from the southeast, encircle Sakhalin from the
north, enter the Gulf of Sakhalin, and later Amur estuary in the first ten
days of June. Here the animals feed throughout the summer. Roughly at
the same time, a part of the herd from the northern extremity of Sakhalin
turns not into the Gulf of Sakhalin, but moves west and arrives in the
large bays of the Shantarsk Sea over large masses of dense ice. There
they are encountered throughout July, August, and September. Individ-
ual small herds may arrive from the Shantarsk Sea along the coasts in
the northeast and are sometimes seen in the region of Ayan Bay. Some
animals may even reach the estuaries of Okhota and Kukhtui rivers.
The nature of migrations of belugas changes in autumn. Instead of
large compact herds, small isolated groups scattered over a large water
body are seen. With the formation of a compact ice cover at the end of
October - November, belugas abandon points of summer habitation. A
small part of the herd arrives October end or early November from the

НИ

north into Tatar Strait and apparently exits through La Perouse Strait
into the Sea of Okhotsk. Most of the herd from the Shantarsk Sea and
the Gulf of Sakhalin encircle Sakhalin from the north at October end
and in November and head southeasterly. During autumn migrations very
large herds are not seen. The periods of migrations cover a long time
frame. Probably, with the disappearance of schooling fish, some animals
begin to leave the summer feeding waters even before ice formation;
formation of dense or stationary ice cover then compels the remaining
animals to leave. The wintering sites of belugas have not been established
(Dorofeev and Klumov, 1936a; Arsen’ev, 1939).

According to the latest observations, groups of belugas have been
recorded north of Sakhalin in winter in air holes among dense drifting
ice (G.A. Fedoseev). It is possible that some herds spend winter amidst
ice in the midpart of the Sea of Okhotsk.

In Penzhina and Gizhaga bays of Shelikhov Gulf, belugas are seen
close to the coasts at the end of May immediately after the thawing of
coastal fast ice in the bays and gulfs. The animals remain there through-
out summer feeding on schooling fish (herring and salmon). They per-
form regular local migrations, their periods and directions depending on
the powerful low and high tides (at some points during low tide, the
coast is exposed for a distance of 5 km or more). Belugas inhabiting
Shelikhov Gulf reach west up to Tauy Bay and some animals or small
groups have been encountered even more westward, right up to Okhotsk.
A few animals are encountered in the summer months along the west
coast of Kamchatka (predominantly in its northern section). With the
appearance of floating anchor ice and the subsequent freezing of coastal
waters in October-November, belugas leave the coasts and are no longer
seen in these waters in the winter months (Arsen’ev, 1939). The points of
wintering of this herd of belugas and the courses of their migrations have
not been established. The winter ice cover of Shelikhov Gulf is typical.
Thus in Gizhaga Bay coastal shore ice 20-25 miles wide is formed while
the midpart of the gulf is covered with drifting ice. In Penzhina Bay only
small straits freeze while the entire remaining water body is covered by
floating ice of varying density. It is quite possible that herds of belu-
gas may winter in the regions of sparse ice without undertaking distant
migrations. There are, however, no observations on winter movements.

Migrations of belugas in the Bering and Chukchi seas have been very
poorly studied. In summer the animals concentrate in the Gulf of Anadyr
and the Anadyr estuary. Many schooling fish entering this river serve
as food for the belugas and hence beluga arrivals are often en masse.
Belugas are seen scattered throughout this bay in July. At about this time

578 also, belugas are seen at many points in the Chukchi Sea: Cape Serdtse

778

Kamen’, Kolyuchin Inlet, Cape Shmidt, and probably somewhat later оп
Wrangel Island and north of it (up to 72 to 73° М lat.). Simultaneously
they are also seen on the coasts of Alaska where they are known from
Bristol Bay to at least Cape Barrow. In Bering Strait, spring (April-
May) migrations of belugas occur in the north while autumn migrations
(September-October) are observed in the south (Arsen’ev, 1939; Nikulin,
1946).

The migrations of belugas in these waters can tentatively be described
as follows. The animals winter in the eastern and northeastern parts of
the Bering Sea. In April-May, with the disappearance of ice, they begin
to move north and northwest. Some belugas exit into the Chukchi Sea
and are distributed over an immense water body, from the boundaries
of the East Siberian Sea to Beaufort Sea and Wrangel Island, depend-
ing on the disposition and density of ice. Other, probably more, animals
remain at the inlet into the Gulf of Anadyr and with thawing of ice, are
scattered over this water body, where they spend the summer months. In
autumn (October-November), with the compaction of ice and the for-
mation of young ice, animals from the Chukchi Sea move southward,
cross Bering Strait, and leave for their wintering sites. The ice displaces
the belugas from the Gulf of Anadyr also, compelling them to move east
and southeast.

In most cases, the dispersal of migrating marine mammals (whales,
belugas, and walruses) in the Chukchi Sea is restricted in the west by the
line Cape Shmidt—Wrangel Island since De Long Strait almost year-
round is covered by a broad strip of very dense ice (Arsen’ev, 1935).
On encountering this massive ice obstacle, the animals change course
northward toward Wrangel Island and travel farther west into the waters
of the East Siberian Sea only in years of little ice.

In winter months belugas are noticed at many points in the Bering
and Chukchi seas, where they remain in large air holes and open water
pools formed among the floating ice. Wintering belugas are noticed
almost every year close to Provideniya Bay (northeastern part of the
Gulf of Anadyr); animals are sighted at different points along the coast
of the Chukchi Sea: Nunyamo, Lesovsk, Dezhnev, Serdtse-Kamen’, and
other capes (Nikulin, 1951; Uspenskii, 1958; Kleinenberg er al., 1964).

In the Beaufort Sea, belugas are encountered in summer in Amund-
sen Gulf and McClure Strait, to the west of Banks Island, at many points
in the MacKenzie delta, and to the west of the estuary of this river. It has
been suggested that, in the Beaufort Sea, belugas even winter far from
the coasts since they are seen in spring concurrently at many points
(Vladykov, 1944; Sergeant, 1962a).

579

dag

In the eastern part of the Canadian Arctic, belugas similarly under-
take seasonal migrations, which are most distinctly manifest in Davis
Strait and Baffin Bay. In summer they sometimes reach 80°N lat. (Hall
Basin), descending in winter to the middle of Davis Strait: Cumber-
land Sound—Sukkertoppen. Regular migrations in spring and autumn
are noticed in Hudson Strait but a small number of belugas winters
in the western and northern sections of Hudson Strait, eastern part of
Cumberland Sound, and in Lancaster and Jones sounds. In the Gulf of
St. Lawrence, as it frees from ice, belugas are seen in the estuary of the
river (often transgressing into the river) and at other points on the coast
but abandon these regions in autumn.

The population living east of Greenland probably overwinters to the
north and northwest of Iceland. In spring the animals migrate northward,
reaching Spitsbergen, often in large numbers, and return to the wintering
sites in autumn (Vladykov, 1944; Daan and Douglas, 1953; Sergeant,
1962a).

Transgressions of belugas beyond the limits of their normal range
are not very rare. In the Bering Sea, belugas were noticed in the waters
of Karagin (58°N lat.) and Commander Islands (54°N lat.) (Tomilin,
1957) although 46° М lat. serves roughly as the southern boundary of the
distribution of belugas in the Sea of Okhotsk. From the Chukchi Sea
some animals or small groups reach the estuary of the Kolyma and the
central part of the East Siberian Sea, although they may reach there later
from the Novosibirsk Islands. Animals of the two populations may be
encountered there.

Transgressions into the Atlantic Ocean usually occur in particularly
cold winters. In the western part of the North Atlantic, the southernmost
transgression was noticed in Massachusetts Bay (42°N lat.) and in the
eastern part along the coasts of Great Britain, Holland, Schleswig coasts
in the Baltic Sea, Estonia, Finland, and even France (von den Brin, 1958;
Kleinenberg e7 al., 1964). During 1964 to 1966, eleven sightings of belugas
were recorded in the waters of Central Europe. The well-known case of
the long residence of a beluga in the Rhine is particularly interesting.
Close to Nijmegen town, Holland, a beluga was sighted on May 18, 1966.
Gradually, ascending against the current, it crossed Deventer, Kampen,
Duisburg, Dusseldorf, and Bonn, and on June 13 reached Honnef, which
is 400 km from the sea (50° 40/S lat.). Here the beluga turned back and
entered the sea on June 18 after moving downstream for 400 km in five
days. Attempts to catch it in a net or by roping or using an injector gun
were not successful. This adult beluga with a body length (visual) of 5
to 6 m spent a month in fresh water, which is rather unusual (Slijper,
1967).

580

780

Reproduction. ш the Sea of Okhotsk mating of belugas was recorded
in April-May (Nikol’skii, 1936); in the Gulf of Ob in July (Zaikov, 1934);
in the Barents and Kara seas from May through August, peaking in May-
early July (Kleinenberg and Yablokov, 1960); in the Gulf of St. Lawrence
from February to August with a large number of animals mating in May
and June (Vladykov, 1944); and in Hudson Bay from March through
September, peaking in May (Daan and Douglas, 1953). Thus the period
of mating extends for about six months but the majority of females are
fertilized over a relatively short duration, 1.е., from April end to early
May. Only some individual animals mate at other times.

In the White Sea, calves with the umbilical cord intact were encoun-

tered in mid-July (Provorov, 1957) in the Gulf of Ob (Zaikov, 1934), in

Hudson Bay (Daan and Douglas, 1953) on August 15, and in the Sea of
Okhotsk from July 18 to 26 (Kleinenberg et al., 1964). However, judging
from the dimensions of the suckling calves caught in the summer months,
their births may have taken place much earlier (Kleinenberg et al., 1964).
In all probability, the period of parturition, like the period of mating, is
protracted and births may occur from early spring throughout the sum-
mer months. The period of maximum births has not been ascertained.
Thus gestation in belugas extends for 11-12 months and, according to
one view, might be extended 13-14 months (Sergeant, 1962a).

Information on the duration of lactation is quite contradictory, from
5-6 months to 12 months or even longer. Until the end of September
(close of the hunting season and specimen collection), calves were found
feeding exclusively on milk. In the stomach of animals of the next color
stage, i.e., gray belugas, milk was also found together with food gathered
independently, but the age of these animals could not be established. It
would probably be more correct to assume that the period of lactation
extends for six months (Kleinenberg et al., 1964). At the same time, the
teeth of young belugas emerge roughly at 10 months of age and hence it
is possible that milk suckling of the calves ceases in this period.

The female usually delivers a single calf although, as among other
cetaceans, twins are sometimes encountered. One female contained three
embryos (two males and one female). It has been suggested that female
belugas are capable of mating at a short interval (two to four weeks) after
parturition. This suggests the possibility of annual births but investiga-
tions carried out on various beluga populations have shown that some
percentage of mature females do remain barren every season. The ratio
between barren and gestating females in a herd varies considerably in
different years as also in different regions of habitation. It has not been
possible to establish the pattern of barrenness and hence the reproduc-
tion cycle of this animal can only be surmised. In some cases, it appears

781

probable that а female may give birth for five to six or more consecutive
years, after which she rests for a year (or more). In other cases, such rest
periods might set in after two or three years (Tomilin, 1962; Kleinen-
berg et al., 1964). It is also possible that the reproduction cycle varies
for females of different age groups. Therefore, the tempo of population
replenishment has yet to be established and recommendations for the
rational commercial exploitation of the beluga are difficult to make.

In the period of mating it was sometimes observed that two or more
males chased a single female while groups of 10-20 adult males, isolated
from mixed groups, were encountered concurrently. This suggests the
possibility of polygamy among belugas (Sergeant, 1962а).

Growth and development. Measurements of beluga embryos from
Greenland waters showed that in the period of embryonal growth, devel-
opment is relatively uniform (Table 53).

Since the period of births is very prolonged, embryos of different
size may be encountered at any time of the year (Fig. 335). Thus in
the northern seas of the Soviet Union, the length of measured embryos
varied from 7 to 168 cm, the embryos investigated in July being larger
than those found in August and September (Kleinenberg and Yablokov,
1960). The maximum dimensions of embryos differ in different regions of
the species’ range. In the Kara Sea, the largest embryo was 150 cm long,
in Hudson Bay 151 cm, Barents Sea 170 cm, Sea of Okhotsk 175 cm,
waters of western Greenland 183 cm, and in the Gulf of St. Lawrence
213 cm. Newborn calves are often smaller than the larger embryos. Thus
the length of the smallest calves (from among those measured) was (in
cm): in western Greenland 116; Sea of Okhotsk 130; Kara Sea 140; Bar-
ents Sea 147; Gulf of St. Lawrence 147; and Hudson Bay 153. The average
body length of a newborn calf, covering all the regions, is assumed to be
150 cm (Kleinenberg et al., 1964).

The growth rate of belugas is the highest in their first year. The
average monthly increment has been put at 30 cm; in the lactation period

Table 53. Body length of beluga embryos, cm (Nielson and Dagerbol, 1930; after
Tomilin, 1957)

Month Number of © Body length Average
specimens

November 36 32-78 60.2

December 40 46-104 87.3

January 11 88-112 96.3

February 19 122-142 129.9

March 19 113-183 142.2

April 6 142-158 149.1

782

Fig. 335. Embryo of beluga (figure Бу М.М. Kondakov).

the increment should be more than after changing over to independent
feeding. By the end of the first year the animal reaches a length of almost
300 cm. The dimensions of belugas at the age of up to one year from
the Sea of Okhotsk varied from 140 to 280 cm, the peak of the growth
graph falling in the 180-200 cm interval.

Further growth of belugas is identified from the color changes of the

581 animals in three stages. Animals of the first color group, called “gray”
animals, have a bluish-gray body and are immature. The next color group,
the “blue” group, covers animals transiting from gray to white. Mature
as well as immature animals of both sexes are encountered in this group.
Finally, the “white” group comprises adult mature animals of various
sizes with a pure white body color. The age of transition from one color
group to another has not been established.

It has been assumed that the average annual increment in the second
and third years is 75 cm and in the fourth year about 45 cm among
males and about 30 cm among females (Sergeant, 1959a). The males
soon outgrow females and hence the average size of adult males is more
than that of females (Table 54).

Age determinations of females based on dentine layers in the teeth
showed that the immature group comprised animals of three to four years
(body length 202-317 cm) but females aged two to five years could be.
placed in the group of potentially mature animals. The minimum age of
mature females is three years but the maximum number of northern bel-
ugas mature at age four to five years or more (body length 315 - 435 cm).
Among mature belugas, the growth rate decreases sharply. In the first
year after sexual maturity, the female adds an average of 20 cm, in the
second year 5-7 cm, and in the third year 2-3 cm. In the fourth to the
fifth years of maturity, growth ceases and the animals attain physical
maturity. By then, they measure 355 -360 cm in body length and fall in
the age group seven to nine years. The body length of females in the
Far East in the first year of maturity rose by 28 cm, in the second and
third years by 7-9 cm, and in the fourth and fifth years by 2-4 cm. Their

582

783

Table 54. Body dimensions of belugas of different color groups, cm (Kleinenberg ef al.,

1964)
Region Color Males Females
ge) ee ИЕ ВНИИ. ИЕ ВЕЧЕ
Number of Body Average Number of Body Average
specimens length specimens length
Gulf of St. Lawrence Gray 8 226-263 247 13 203-282, 239
Вше 12 282-345 311 18 282-343 318
White 59 329-447 406 38 310-409 372
Kara Sea Gray 1) 239-340 276 15 224-341 272
Вше 20 300-393 336 17 274-365 322
White 35 343-472 400 26 326-390. 353
Sea of Okhotsk Gray 34 195-310 252 28 170-357 286
Blue 173 187-479 351 98 251-422 18531
White 136 359-500 447 53 360-418 388

physical maturity set in in the fifth to the sixth year after sexual maturity.
The average body length of physically mature females in the Far East
was 390 cm (Kleinenberg её al., 1964).

Male belugas from Hudson Bay attain sexual maturity at six to nine
years (12-18 dentine layers) at a body length of 275 -320 cm and females
at six years (12 layers) at a body length of 275 cm. Belugas on the eve
of the “white” stage have 13 to 18 dentine layers. Up to 35 layers are
seen in very old animals. The maximum number of layers counted is 50
(Sergeant, 1959a). Longevity of the belugas has not been established.

Enemies, diseases, parasites, mortality, and competitors. The killer
whale and the white polar bear may be regarded as enemies of the beluga
albeit they inflict little damage on the population. Apart from diseases
caused by parasites, no other has been recorded. Ectoparasites are not
known.

Thirteen species of helminths are known among belugas: trematodes
three, cestodes one, nematodes eight, and acanthocephalans one. None
of the three species of trematodes has been reported in any other ani-
mal. The trematode Odneriella seymouri Price was found in the Bering,
Okhotsk, White, and Barents seas. Leucasiella arctica Delamure and
Kleinenberg, parasitizing the rectum, has been detected in the Barents
Sea. Leucassiella mironovi Krotov and Delamure was found in the small
intestine of belugas from Aniv Bay and La Perouse Strait.

The cestode Diphyllobothrium lanceolatum Krabbe, reported before
for three or four species of pinnipeds and possibly common porpoises,
is evidently a facultative parasite of the beluga.

Four of the eight known species of nematodes have been exclusively
reported in the beluga. Anisakis (Anisakis) kukenthalu Cobb, parasitizing

784

the stomach, was found in the Arctic Ocean (Spitsbergen), the Barents,
Kara, North, and Okhotsk seas, and in Aniva Bay (Sakhalin). Sztenu-
rus arctomarinus Delamure and Kleinenberg was detected in the lungs
of belugas from the northern Arctic Ocean, and White and Barents
seas. Otophocaenurus oserskoi Skrjabin, localizing in the auditory organs,
was found among belugas from the waters of Sakhalin, northern Arc-
tic Ocean, and White and Barents seas. Stenurus pallasii van Beneden,
parasitizing the auditory organs, bronchi, and circulatory system, was
detected in the Arctic. Anisakis (Anisakis) simplex Rudolphi, very widely
distributed among marine mammals, was found in the gullet, stomach,
and intestine of belugas and also in ten other species of toothed whales,
two species of baleen whales, and Steller’s sea lion. It is known from
various places in the North Sea, eastern Kamchatka, Japan, and New
Zealand. Terranova decipiens Krabbe, parasitizing the stomach and intes-
tine, has been detected in the beluga only once (Kara Sea), whereas
it occurs in 12 species of pinnipeds, 2 species of toothed whales, and
2 species of baleen whales in the North Sea, the Atlantic and Pacific
oceans, and in Antarctica. The nematode Stenurus minor Kuhn, found
among belugas in the northern Arctic Ocean, North Atlantic, North,
Black, and Azov seas, and along the Asian coast of the Pacific Ocean,
parasitizes the auditory organs, bronchi, heart, and blood vessels of only
common porpoises in addition to the beluga. Finally, Crassicauda gili-
akiana Skrjabin and Andreeva parasitizes the same organs of the beluga
and common porpoise. It has been detected in the Amur estuary and in
the waters of Kuril Islands.

The lone species of acanthocephalan parasitizing the intestine of
beluga, Corynosoma strumosum Rudolphi, is known in 11 other species
and subspecies of pinnipeds, common porpoises, the sea otter, and also
land carnivores and ichthyophagous birds. Among marine mammals, this
helminth has been found along the Atlantic coast of Europe, in the Baltic
Sea, Lake Ladoga, along the Canadian coasts, in almost all the northern
waters of the USSR, around the Kuril Islands, in the waters of Sakhalin,
along the coasts of California, and in the Caspian Sea (Delamure, 1955;
Tomilin, 1962).

Population dynamics. Changes of beluga populations relative to nat-
ural factors are not known. The comparatively low scale of hunting per-
haps does not influence the numbers of this species.

Field characteristics. The pure white color of adult animals distin-
guishes this species from all other marine mammals. Further, the beluga
has no dorsal fin. The corpus adiposum [melon] is prominent, reaching
almost the end of the broad snout; a “beak” is absent. The teeth are
conical and develop in both jaws; canines are absent. In all the color

583

785

groups, coloration is monochromatic. Belugas usually stay together in
groups which gather into herds of various strengths. In the period of
en masse arrival of fish which serve as their food, belugas form herds
of thousands of animals. This species is essentially a coastal form but is
also encountered amidst ice at times of high density. (V.A.)

Economic Importance

Although beluga hunting has been carried out for several centuries in
our country, it has never been extensive. In the 1920s to the 1930s,
beluga hunting in the seas of the Soviet north and Far East expanded
but was not intensive. In the years of maximum catch (1931-1935), the
number caught in the Soviet Union did not excede 5,000. The maximum
catch occurred in the Far East up to 1940 but thereafter dropped to
a few hundreds per year. After 1950, hunting intensified somewhat in
the White, Barents, and Kara seas, but nonetheless the annual catch
has never exceeded a couple thousand animals. Outside the limits of
our waters beluga hunting is also not significant. Formerly, the most
productive area was the Gulf of St. Lawrence where some 2,000 belugas
were caught annually, dropping to a few hundreds in 1932 - 1938; hunting
ceased altogether in 1940. In Hudson Bay, from 1948-1960 more than
1,000 belugas were caught only in three years, the catch varying from 600
to 900 in the other years. In the waters of Greenland and Spitsbergen,
the annual catch used to vary widely, from a few tens to several hundreds.
In the rest of the Canadian Arctic regions, the catch was invariably low.
Statistics of beluga hunting are far from perfect and the figures cited
should be regarded as highly tentative. Nevertheless, the catch of beluga
throughout its range has never been much and hence the products of
beluga hunting have never been of commercial or economic importance.

The techniques of catching the beluga are diverse and specific to
each region. The oldest equipment for catching the beluga in the Euro-
pean north is the White Sea sweep net whose length depends on the
capability of the cooperative (number of operators and karbases* avail-
able) and may go up to almost 2,500 m./In principle, this is a dynamic
method of catching but is extremely primitive in practice. A composite
net 2,400 m long is cast in the open sea, not farther than 7 to 8 km from
the coast; the operation involves 24 boats and 170 people. The depth
of casting should not exceed the wall height of the sweep net. Two sec-
tions each of the net set up on two boats are stitched together before
setting out to catch and these boats enter the sea in pairs. Such pairs are

* Large boats with high sides used in marine and river transport—Translator.

584

786

deployed at some distance from each other in a semicircle open on the
side of the anticipated approach of the herd of belugas. As the animals
enter this semicircle, each pair of boats begins to diverge in the opposite
direction, casting the sweep net into the water and joining it with the
successive pairs to ultimately form a circle with the sweep net. This circle
is gradually closed, its diameter reduced, and the boats enter the circle
of the sweep net. The belugas are killed from the boats by shooting or
battering with ice chisels. The killed animals are towed to the coast and
dressed. The production cost of hunting by this method is extremely high
(Klumov, 1936b, 1939).

Another, much older piece of equipment used in catching is the Nor-
wegian stationary net. Its wall height varies from 8 to 14 m and length
450 to 1,200 m. The net is set up only at places of regular arrivals of
belugas at an angle of 30 to 35° to the coastline. A brigade of catch-
ers consists of 8 to 12 persons with some boats. The men in the boats
approach the beluga herds from the sea side and attempt to chase the
animals into the net inlet by raising a racket, after which the inlet is
sealed by a net. The belugas trapped within the net are battered with ice
chisels and dragged to the coast (Heptner, 1930).

The fabric mesh or so-called “polovinka” [meaning half] is similarly
passé. Its length is 25-50 m, height 8-12 m, and mesh size 50-60 cm.
These nets are set up on anchors perpendicular to the coast in a check-
ered pattern at a distance of not less than 15 m from each other. The
lower seine rope should reach the bottom. A team of five persons can
operate 30 to 40 such “рооушкКаз”. The animal passes through the mesh
but its caudal fin becomes entangled and the net twists around it. With no
possibility of floating, the animal succumbs to asphyxiation. Best results
are obtained on dark autumn nights when the belugas cannot see the net
(Klumov, 1939). These traps have been modernized in recent years, with
plant fiber replaced by capron [nylon] or flexible galvanized steel cables,
which are less visible in the water. Hunting is done in the open sea from
a ship (in the region of Novaya Zemlya and in Yenisey Strait), which
chases a herd of belugas into the net. Such innovations have significantly
increased the catch in such traps (Butorin, 1957).

The development of beluga hunting in the Far East took a different
course. The prototype of the beluga net here was an ordinary casting
net used for catching salmon. Special coastal nets of this type, up to
1,000 m or more in length and made of thick strong material, were used
for catching belugas. The ends of the net, laid on a kungas,* were affixed
to the coast. As a herd approached, the boat with the kungas in tow

* Open fishing boat with a carrying capacity of 3-5 tons—General Editor.

787

surrounded it and drew the second end of the пе! to the coast. The
net was then hauled in manually or with a winch to the coast and the
captured belugas gradually removed, killed, and dressed (Fig. 336).

Later, this coastal net was elongated to 2,500 m and was gathered
on two kungases standing together with two boats on anchors more than
1 km from the coast. As a beluga approached, the kungases were towed
simultaneously in the opposite direction which hastened trapping of the
animal and enlarged the area of catch. Both ends of the net were drawn
to the coast and its further hauling in and removal of the animals done
as in the coastal method of catching.

In the course of further modernization (I.K. Nepomnyashchii), a
deep net was designed with a length of about 1,000 m and a wall height of
18 m. Using two boats and one kungas, a herd of belugas was surrounded
comparatively far from the coast, the net with the caught belugas was
towed to the coast, and the trapped animals were recovered from the
large net by the conventional method. This did not make for venturing
beyond 10 km from the coast, towing took several hours, and if the
weather turned foul, there was danger of losing not only the catch, but
the net itself. A new net 1,600 m in length with a wall height of 18 m
was subsequently designed. It was cast from two motor boats without
a kungas. On sighting a herd of belugas, these boats diverged in the

Fig. 336. Caught belugas laid out on the coast. Sakhalin, 1938 (photograph by
V. A. Arsen’ev).

585

788

opposite direction and surrounded the herd with the net. After this,
much of the net was gathered in so that a small circular section remained
under the water. The boats, traveling in the opposite direction, gradually
tautened and transformed this circular section into a line. The belugas
caught in the net died of asphyxiation, were dragged to the approaching
kungas, and hauled to the coast (Fig. 337). The team of catchers using
these nets consisted of 25 to 30 persons (Arsen’ev, 1940).

At places with a high variation in sea level during low and high tides,
the belugas trapped in the net were left until low tide set in and then
dragged to the coast for killing and dressing.

The coastal people sometimes shoot belugas with rifles but many
of the killed animals are lost. Sometimes the beluga is first harpooned
manually and then shot. This method was in vogue for sometime in the
Gulf of St. Lawrance and other regions of the Canadian Arctic. At some
places the belugas are chased into narrow straits and the exit sealed off
by a net. In most cases, these primitive methods of hunting result in a
low catch.

In dressing the killed belugas, the subcutaneous fat layer together
with the skin is removed first. The fat layer is later separated from the
skin and oil obtained from the blubber. The skin, now free of fat, is
salted and after a few days, the thick layer of epidermis recovered. The
skin is again salted and, after sometime, packed in barrels for dispatch

Fig. 337. Towing the trapped belugas. Sakhalin, 1938 (photograph by V.A. Arsen’ev).

586

587

586

789

to leather manufacturers. The remaining carcass is processed into edible
parts and those to be used in fertilizer meal. Where there are no process-
ing facilities, the carcass is used as feed for fur-bearing animals. The oil
melted from the blubber is used for commercial purposes. The oil from
the head (corpus adiposum [melon]) and from the hollow of the lower
jaw is collected separately. This oil possesses the property of remaining
fluid at low temperatures and is used a special lubricant. The skin of the
beluga can be used in making shoe soles and other footwear articles as
well as halters and reins. Further, the intestine of the beluga can be satis-
factorily used in the sausage industry; 30 m long, the intestine lengthens
three times over after removal of the muscular layer. Blood accounts for
roughly 5% of the overall weight of the animals. The valuable product,
black albumin, can be produced from the blood (Dorofeev and Arsen’ev,
1936; Druker and Gakichko, 1936).

The average weight of males and females of different color groups of
belugas are given in Table 55. Table 56 gives the weight of the body parts
of some animals. The ratio of the weight of different body parts of belugas
without classification into age (color) groups is shown in Table 57 (those
sections of the carcass used commercially were weighed).

The beluga reserves in all the regions of its extensive range are not
as large as often assumed and hence their catch should be based on a
knowledge of the biology and population of these animals. Beluga hunt-
ing has been restricted in recent years but could be enlarged without
affecting these reserves. This could be done first in the Far East seas
where not more than a thousand belugas have been caught annually
since the 1940s. In the northern seas the increase should be more cau-
tiously approached since hunting here is already vigorous although the
possibilities of commercial exploitation of the animal population have
not been fully realized. It should not be forgotten, however, that in Hud-
son Bay, for example, some restrictions have been imposed on hunting ©
since even the comparatively low catch over several years has reduced

Table 55. Average weight of whole animals from Sakhalin Bay, kg (Arsen’ev, 1935b)

Sex Color group
White Blue Gray Sucklings Total

No. Av.wt. No. Av.wt. №. Ау. м. №. Av.wt. №. Av. wt.

Males 120 — 880.1 3811 489.4: 21 310.4 2 150.0 181 726.2
Females 81 6566 31 506.1 Zi 28184 4 224 1432 53726
? 7 — 3 — 63 == 2 — 15 —

Total DOSE SOO een. 494.35 111 266.5 8 122.9, 399. 2577.4

586

587

790

Table 56. Weight of body parts of belugas from Sakhalin Bay, kg (Druker and
Gakichko, 1936)

Body part Color group of belugas
White Blue Gray
1 2 3 1 2 a ani
Skin:
Epidermis (‘“‘jacket’’) 34.5 aan 28.2 26.5 24.6 14.2
Dermis 18.5 57.2 27.3 13.9 11.8 9.1
Fat of:
Trunk 240.8 151.5 294.3 146.6 114.5 66.6
Неаа 4.1 TES) 10.9 4.1 2.8 1.4
Jaw 0.8 0.5 0.3 0.6 0.6 0.2
Flesh 242.2 184.4 217.4 208.6 155.7 58.6
Bones 68.2 ТР 86.0 65.3 37.3 25.3
Flippers 8.1 9.1 9:7 9.1 52 32
Caudal flukes 8.9 9.0 12.5 9.5 5.2 3.5
Viscera:
Liver, heart, lungs, 723 49.0 65.6 lel 35.5 15S)
kidneys, and stomach

Intestine 22.6 16.0 18.1 19.0 11.5 5.5

Table 57. Weight of carcass sections of belugas from Sakhalin Bay, kg (Dorofeev and
Arsen’ev, 1936)

Number Av. м. Wt. of skin Wt. of carcass Wt. of skin Wt. of skin

of of one with fat and without viscera = and epidermis without
belugas animal epidermis without fat epidermis
Ау. wt. Yof Av.wt. %of Av.wt. %of Av.wt. Hof
body body body body
wt. wt. wt. wt.

66 509.4 262.5 51.5 246.9 48.5 52.3 10.3 22.5 8.6

the beluga reserves. Investigations should continue for a more precise
determination of numbers in each population. (V.A.)

Genus of Narwhals (Narwhals or Unicorns)
Genus Monodon Linnaeus, 1758

1758. Monodon. Linnaeus. Syst. Nat., ed. X, I, p. 75. Monodon monoceros
Linnaeus, 1758.

1772. Ceratodon. Brunnich. Zoologiae fundamenta, р. 48. Substituted for
Monodon Linnaeus, 1758.

1804. Narvalus. Lacépéde. Hist. Nat. Cétacées, p. 37. Narwalus vulgaris
Lacépéde = Monodon monoceros Linnaeus, 1758. (V.H.)

588

791

Dimensions are the largest in the family, the body length reaching 610 cm.

Compared to the beluga, the neck is poorly developed. The head is
obtuse and spherical. The mouth section is small.

The body of young animals is monochromatic, dark (bluish-gray,
slaty, or slaty-blue) but in adults light-colored with numerous dark spots
of irregular shape not exceeding 5 cm in diameter on the head, dorsum,
and flanks. The spots on the upper parts of the head, neck, and caudal
stem sometimes merge to form an overall dark background.

The skull structure in general is the same as in the genus Delphi-
napterus but asymmetry is more sharply manifest. Teeth are lacking in
the lower jaw. The upper jaw has only one tooth in each half (in females,
they usually do not emerge). The left tooth in males is transformed into
a spiraled screwlike tusk reaching 3 m in length. The right tooth, how-
ever, is more often concealed within the gum. Sometimes two tusks grow
in males. From time to time, tusks are seen among females too. Verte-
brae 50 to 55; cervical vertebrae not fused. Phalangeal formula: I, _,,
II; _ 9, Ш4-в, [V>-4, and V,_-,. Compared to the beluga, the sternum of
narwhals is short and broad.

Narwhals seek food in the pelagic part of the sea. They feed mainly
on cephalopods (teuthophagous) but also consume fish, possibly crus-
taceans as well. The periods of mating and parturition are protracted.

Narwhals are distributed in the waters of the Arctic and North
Atlantic. There is no special hunting for narwhals.

The genus comprises a single species: the narwhal or unicorn, М.
monoceros Linnaeus, 1758. (V.S.)

NARWHAL OR UNICORN

Monodon monoceros Linnaeus, 1758

1758. Monodon monoceros. Linnaeus. Syst. Nat. ed. X, I, p. 75. Northern
seas of Europe and America.

1781. Monodon narhval. Borowski. Gemeinn. Naturgesch. d. Thierreichs,
2 Did:

1804. Narwalus vulgaris. Lacépéde. Hist. Nat. Cétacées, р. 37, 142, pl. 4.
Substituted for Monodon monoceros Linnaeus, 1758.

1811. Ceratodon monodon. Pallas. Zoogr. rosso-asiatica, I, p. 295. Seas
of the Russian arctic (“In mare arcticum Ruthenum imperium
alluente”). (У. H.)

Diagnosis

Only species of the genus.

589

792,

Description

Males can attain a body length of 610 cm and females 420 cm.

A rounded small head is set off from the trunk by a weakly dis-
cernible neck (Fig. 338). The spermaceti case slightly overhanging the
upper jaw gives the head its roundness. The upper lip projects slightly
forward above the fleshy lower lip. The slightly convex dorsum carries a
low fold of skin up to 5 cm thick and 75 cm long.

The asymmetry of the skull (Fig. 339) is manifest in the more intense
development of maxillae and premaxillae on the left side while the cra-
nium is better developed on the right side. The importance of the tusk
(tooth) to the narwhal is not clearly understood. It has been suggested
(Thompson, 1939) that the spiraled tusk serves to stabilize the animal
body against rotation along the axis during circular movements of the
caudal flukes during swimming. It is perhaps more correct to regard the
tusk as a secondary sex characteristic of males (Chapskii, 1941; Sleptsov,
1955) serving as a defense organ. It is possible that the animal uses this
tusk to pierce holes in the ice through which not only he himself, but
also other members of the herd, i.e., females and young animals without
tusks, can breathe (Tomilin, 1957).

Cervical vertebrae 7, thoracic 11-12, lumber 6-10, and caudal
26-27. пи

The main measurements of the skull (Tomilin, 1957), average of one
to four animals are (in cm): condylobasal length 61; zygomatic width 39;
length of rostrum 28; width of rostrum at base 21; length of lower jaw
50; and mandibular symphysis 6. (V.S.)

Geographic Distribution

Basin of the northern Arctic Ocean. This species lives mainly in the
arctic zone, less often seen close to the coasts.!?

Geographic Range in the USSR

Narwhals have been observed in the Barents Sea in Novaya Zemlya at
76° N lat. (Russkaya Gavan’), in Franz Josef Land archipelago, in the
midpart of the Kara Sea, around Dickson Island, and in the Chukchi
Sea. In 1938, during the drifting of ice breakers “Sedov,” “Sadko,” and
“Malygin,” narwhals were sighted no less than 15 times in the region
81° 21’-82° 15’ М lat. and 136° 15’ - 138° 15’ Е long. They were sighted

19 Distribution is equally known from the records of animal remains as from field
observations.

793

ут

А

РРР

РРР

ЕЕ

= РР
РИ,

22:57:

589

from the floating arctic stations in June, July, and August of 1950 north-
west of Wrangel Island, in 1955 north of De Long Islands, and in 1956
between Severnaya Zemlya and Franz Josef Land. Remains of narwhals
have been found in the estuaries of rivers Olenek, Anabar, Khatanga,
and Lena, and also on Lyakhovsk Islands, Cape Serdtse-Kamen’, and in

the Chukchi Sea. In the summer months narwhals are most common in
the waters from 80° to 85° N lat.

589

590

794

Ie =

7

Fig. 340. Skull of a narwhal with two tusks (figure by М.М. Kondakov).

Geographic Range outside the USSR

American Arctic, north of Labrador and Alaska and waters of Green-
land (Fig. 341); known in the Canadian archipelago in Hudson Bay,
Davis Strait, and in Baffin Bay right up to Smith Strait, in the waters-of
Spitsbergen, later in the waters of Cape Barrow and in the Beaufort Sea
close to the Colville estuary. Beached narwhals or their remains have
been found at some points along the coasts of Greenland, in the Beau-
fort Sea, near Cape Gal’skett, and in Kiwalik Bay in the Chukchi Sea at
60° N lat. and 162° E long. (Tomilin, 1957, 1962; Uspenskii, 1958; Hall
and Kelson, 1959; Gest, Buckley, and Manville, 1960). (V.A.)

Geographic Variation
Not established.
Biology

Population. One of the comparatively rare forms of cetaceans, this species
does not form large congregations. It is more abundant in the waters of
Greenland than in other parts of the range.

795

(лэмэзту “WA

) 504220и0щ иороиой! Чечилеи dy} JO a8ueI $10945 ‘1$ "ВЫ

065

59

—

796

Food. {Information on this aspect is very scant. Cephalopods evidently
serve as the main sustenance, followed by fish to a lesser extent. Among
cephalopods, the squids Gonatus fabricii and possibly Rossia, Bathipoly-
pus, Ommatostrephes, and some others have been detected. Fish food
includes cod, flounder, halibut, skate, salmon, and herring (Smirnov,
1935; Tomilin, 1957). These animals feed predominantly in the pelagic
region. The oral cavity is well adapted to feeding on cephalopods. Most
of the above fish represent deep-dwelling forms.

Behavior. There are quite a number of references in the literature
to narwhals being encountered in large herds numbering thousands of
animals. In our times, however, such large herds have not been reported.
Narwhals are most often encountered in small groups or singly; rela-
tively large collections are very rare. Herds consisting of a relatively large
number of animals are probably of mixed composition, with adult males,
females, and young animals. Small herds, in some cases, consist exclu-
sively of females with calves. Some twenty instances are known when
even a relatively small herd consisted exclusively of adult males.

The narwhal usually remains far from coasts and approaches only
sometimes in search of food. The animal can remain submerged under
water for quite a long time, after which it surfaces eight or nine times
successively at intervals of roughly 3 sec. During surfacing, the animal
makes a loud expiration, flexes its body arcuately, but does not usually
display the tusk above the water. Only when a group of narwhals dives in
a small air hole, do their tusks project above the water (Gorbunov, 1940).

In July, 1955, in the western part of Franz Josef Land archipelago
(Cambridge Bay), a group of narwhals, a hundred strong, was sighted
in a state of intense excitation. Sometimes the animals flexed arcuately,
leaped out of the water like dolphins or, while diving under the ice, cast
their head down such that the caudal flukes rose high above the water.
While emerging from under the ice they often lifted the entire anterior
half of the body with the tusk above the water surface. Meanwhile, at
some distance, some ten narwhals remained immobile on the surface.
The diving narwhals were probably feeding. At night, the animals were
quiet and stationary with tusks visible above the sea surface. Some of
the tusks crossed like swords (Sushkina, 1956). Once a narwhal was seen
following a ship. Instances are known of groups of narwhals abandoning
the ice and remaining a long time in small air holes, thereby falling
easy prey to hunters. Mixed herds of narwhals and belugas have also
been recorded. Narwhals produce various sounds, such as loud gurgles,
resonant groans, and howling hoots (Tomilin, 1962).

592

797

Seasonal migrations and transgressions. Observations оп these aspects
are very scant. In the waters of the Soviet Union, the zone of polar ice
edges probably serves as the southern boundary of distribution in winter.
In summer the animals ascend northward and are usually encountered
beyond the 80th parallel. In Baffin Bay they arrive with the ice in Smith
Strait in winter but again return south in the autumn, reaching Disko
Strait by December (70° N lat.). Only some stray animals penetrate south
of Sukkertoppen region (about 65° N lat.) while the main herd remains in
the north. Narwhals are common from Angmagsalik village to Scoresby
Sound (65 to 70° М lat.) on the east coast of Greenland in summer
months (May to August) but the animals may ascend northward up to
perhaps 80° N lat. or beyond.

Some distant migrations of narwhals into the south are known. Their
remains have been found in Mezensk Bay of the White Sea, along the
Murman coast close to the Pechora estuary, at Jan Mayen, Iceland, Faroe
Islands, and along the coasts of Norway, Scotland, and England. One
instance of the death of a female narwhal was recorded in the Elba
estuary in Holland. In the Pacific Ocean, transgressions are known in
Bering Strait and toward St. Lawrence Island. One dead narwhal was
found in the region of Port Moller in the central Alaskan peninsula. In
the last century, one narwhal was found on Bering Island (Commander
Islands) (N. Smirnov, 1935; Tomilin, 1957; Gest, Buckley, and Manville,
1960).

Reproduction. Mating and parturition may occur at almost any time
of the year (N. Smirnov, 1935). The birth of a narwhal calf was witnessed
from the floating station SP-5 in the waters between Franz Josef Land
and Severnaya Zemlya on July 30, 1956. It was 1.5 to 2 m long (Uspenskii,
1958). Very small as well as almost mature embryos were found at the
same time (Fig. 342). The duration of gestation is not known. Usually one
calf is born and twins are rare. The body length of a newborn calf hovers
around 150 cm. Suckling calves are bluish-gray or slate-colored. As the

ica

й мой Ds i A z В Е. .
be VELHO. pete pep Zz LG
ip CEG ETE

Fig. 342. Newborn narwhal (figure by N.N. Kondakov).

592

798 ©

animal grows, the ventral part of the adult gradually lightens (white or
yellowish) while the flanks and dorsum darken with innumerable random
dark brown spots spots of irregular shape. On the dorsum, especially the
upper part of the neck and head, the spots are darker and denser than
on the flanks. The periods of color variation as also the degree of their
consistency with the changing age of the animal have not been established
(Tomilin, 1957).

Enemies, diseases, mortality, parasites, and competitors. One of the
possible enemies of the narwhal is the white polar bear; the young are
threatened by Greenland sharks. Only bone diseases are known in the
narwhal.

Two species of lice have been found in the skin folds of the nar-
whal: Cyamus monodontis and C. nodosus. Among endoparasites, three
species of nematodes are known. Anisakis (Anisakis) simplex Rudolphi,
parasitizing the gullet, stomach, and intestine, has also been found in ten.
species of toothed whales, two species of baleen whales, and in Steller’s
sea lion. It has been detected in the North Sea and in the North and
South Pacific Ocean. Terranova (Terranova) decipiens Krabbe, common
in pinnipeds, seems doubtful in the narwhal. It definitely occurs in two
species of baleen whales, in toothed whales, and in common porpoises.
Torynurus alatus Leuckart is known only in the narwhal from the coast
of Greenland. It has been detected in the skull cavity, eustachian tube,
venous vessels, and lungs (Delamure, 1955).

Field characteristics. Spotted coloration of the dorsum and body
flanks is a characteristic feature of the species. Teeth one pair but usually
do not emerge among females; one tooth among males turns into a long
(up to 3 m) spiraled tusk projecting forward. A dorsal fin is absent. This
species inhabits predominantly the Arctic Zone and can often be seen
in air holes among highly compact ice. (V.A.)

Economic Importance

Only the local Greenland people regularly catch some narwhals and uti-
lize their products to meet personal needs. Elsewhere in the range these
cetaceans are caught merely incidentally in small numbers. Hence this
species has almost no economic importance.

The skin of the narwhal is quite tough and can be used for the same
purposes as the skin of the beluga. The carcass of the animal is useful
for preparing the same products as those from other marine mammals.
The large tusk of the male can evidently be carved or sold as a sou-
venir because of its rarity. Narwhal tusks were highly prized at one time
(Vinogradov, 1949; Tomilin, 1957). (V.A.)

593

799

SUPERFAMILY OF SPERM WHALES
Superfamily PHYSETEROIDEA Gill, 1872

Family of Sperm Whales
Family PHYSETERIDAE Gray, 1821

Dimensions range from medium to large.

The body form is diverse: the anterior half of the trunk of sperm
whales is relatively short but very thick; in dwarf sperm whales, it is
more elongated while the thickness is maximum between the flippers
and the dorsal fin. The head may be very large, up to one-third of the
body length, and obtusely flattened or slightly rounded anteriorly, or
markedly smaller and more proportionate and rounded. The dorsal fin
is low and humplike or quite high and falcate.

The rostrum of the. skull is broadened, flattened, and: has a large
depression on the dorsal surface. The lower part of the spermaceti sac
lies inside this depression; among males, this sac projects forward in
relation to the skull bones. The occipital bone forms a high crest. The
lachrymal is fused with the zygomatic. The petrosal is enlarged toward
the skull. The narrow lower jaw does not extend beyond the anterior
section of the head: the symphysis of both its halves usually forms not
less than one-third of its length.

The teeth, located in deep alveoli, are preserved in most animals
only in the lower jaw, being either rudimentary or altogether absent in
the upper jaw. Keratinized projections on the palate replace the teeth
functionally.

Sperm whales feed mainly on cephalopods of different sizes, includ-
ing very large ones (teuthophagous).

These whales are distributed throughout the World’s Ocean. The
northern limit of the range extends into the Pacific Ocean from Maina-
Pylgo village and Cape Navarin to the Pribilof Islands and in the Atlantic,
from Davis Strait to the region west of Jan Mayen and Spitsbergen and
eastward up to Kanin Nos Peninsula. The southern boundary of distri-
bution reaches the edges of drifting ice in the Antarctic. Sperm whales
are common and abundant while dwarf sperm whales are few and very
rarely encountered.

Diaphorocetus Ameghino, found in the Lower Miocene, already had
a skull with features characteristic of Physeteridae Linnaeus, though it
was not as large as the modern forms, and had teeth in the upper jaw.
Reduction of upper teeth is noticed among representatives of sperm
whales from the Middle Miocene (for example, in Aulophyseter Kellog).

The family is usually (Simpson, 1945) divided into three subfamilies,
of which one is extinct (Hoplocetinae Cabrera, six genera) and two

594

800

contemporary: Physeterinae Flower (sperm whales proper) and Kogiinae
Gill (dwarf sperm whales). The total number of genera is 19. Physeterinae
includes ten extinct genera and one contemporary; Kogiinae comprises
one extinct and one contemporary genus. Thus the contemporary gen-
era of the family are: sperm whales Physeter Linnaeus and dwarf sperm
whales Kogia Gray. In the context of systematics, the contemporary fam-
ily of sperm whales is not a complex group and its division into genera
raises no doubts. However, subspecies have not been well studied.

Only one subfamily of sperm whales proper, physeterinae, with one
genus of true sperm whales, Physeter Linnaeus, has been established for
certain in the fauna of the USSR. The presence of the subfamily of dwarf
sperm whales, Kogiinae, with the single genus Kogia Gray, has not been
established but is entirely possible.

Only sperm whales are of practical importance. They are hunted in
large numbers, especially in our Pacific Ocean waters. (V.S.)

Subfamily of Sperm Whales
Subfamily Physeterinae Flower, 1864

Genus of Sperm Whales
Genus Physeter Linnaeus, 1758

1758. Physeter. Linnaeus. Syst. Nat. ed. X, I, p. 76. Physeter catodon Lin-
naeus.

‚1761. Catodon. Linnaeus. Fauna suecica, р. 18. Catodon macrocephalus

Linnaeus = Physeter catodon Linnaeus. (V.H.)
Large whales with sharply manifest sexual dimorphism: males up to
18-20 m long and females up to 11-13 m.

The large head constitutes one-third of the total length of the body.
The dorsal fin is weakly developed, low, and usually rounded in the
form of a prominence. A few very low tubercles occur posterior to it.
The broad flippers have a rounded outer margin. The caudal flukes are
broad. The blowhole (nostril) is present in the anterior section of the
head to the left.

The length of the rostrum constitutes more than one-half the condy-
lobasal length of the skull. Its dorsal surface has a deep depression. The
skull bones are sharply-asymmetric. The right nasal bone fuses with the
right premaxilla. The pterygoids are adjacent for a significant distance.
Vertebrae usually 50. Teeth 257° 0°).

The narrow scapula has a large coracoid process. Ribs up to 11 pairs,
of which only 3 are joined to the sternum. The body color is dark and
the abdomen only slightly lighter than the upper part.

595

801

The main food object is cephalopods while fishes represent ап addi-
tional item. The ability to dive deep and remain long under water is a
characteristic feature of sperm whales. These are polygamous animals.
Various researchers have put the duration of gestation at 10 to 16.4
months. The periods of mating and parturition are protracted, Females
apparently bear offspring once in two years.

These whales are distributed predominantly in the warm and temper-
ate waters of the Pacific, Indian, and Atlantic oceans (for more details,
see “Description” of the species). Females, compared to males, are more
thermophilic. Significant migrations are a characteristic feature. |

In USSR waters, they are encountered in the Barents Sea and in the
seas of the Pacific Ocean.

Fossil remains have been found in the Upper Miocene and Pleis-
tocene of North America and in the Lower and Middle Miocene of
Europe.

These whales are of great economic importance. A large number is
caught every year.

The genus comprises a single species: the sperm whale, Physeter
catodon Linnaeus, 1758. (V.S.)

SPERM WHALE
Physeter catodon Linnaeus, 1758

1758. Physeter catodon. Linnaeus. Syst. Nat., ed. X, I, p. 76. Orkney

Islands.

1758. Physeter macrocephalus. Linnaeus. Syst. Nat., ed. X, I, p. 76. “Euro-
pean Ocean”.

1818. Physeterus sulcatus. Lacépéde. Mem. Mus. Hist., Nat., Paris, 4,
р. 474. Japan.

1822. Physeter australasianus. Desmoulins. Dict. class. Hist. Nat., 2,
p. 618. Moluccas and New Zealand.

1851. Catodon australis. Wall. Mem. Australian Mus., I, р. 1. Australia.!
(V.H.)

Diagnosis

Only species of the genus.

1 After Hershkovitz (1966); А.С. Tomilin (1957) attributes this name to MacLeay, 1851.

802

Description

The dimensions of this species are the largest of any toothed whale.”

The body is teardrop-shaped and considerably thickset in the anterior
half (Fig. 343). The anterior part of the huge head terminates bluntly
in a “forehead”. The dimensions of the head are mainly determined by
the massive spermaceti sac situated above the skull in the depressions
of the maxillae and projecting beyond the anterior tip of the jaw. The
зрегтасей sac consists of lower [melon or “Junk” and upper [spermatic
organ or “сазе”| sacs made up of connective tissue with a high content
of an only liquid, i.e., spermaceti (Fig. 344). The sacs are surrounded by
connective tissue and muscles. Liquid wax (cetin) is the main constituent
of spermaceti (96.99%). Adult sperm whales carry 1.0 to 5.7 tons of
spermaceti.

The head of males is relatively larger (up to one-third of body length)
than in females (slightly more than one-fourth body length). With age,
the head section of the sperm whale is relatively enlarged while the caudal
end is correspondingly shortened (Ivanova, 1955). The relative increase
in head size commences even in the fetal growth period (Ivanova, 1959).

The cross section of the head appears rounded from above and keel-
shaped from below (anterior to the jaw). The narrow lower jaw with the
mouth closed is held to the head by the connective tissue edges [lips] of
the upper jaw. The unpaired blowhole is S-shaped and located, unlike

596 in Other cetaceans, in the anterior part of the head left of midline and is

595

Fig. 343. Sperm whale, Physeter catodon, male and female (figure by N.N. Kondakov).

Data on the weight of the animals and of the various parts of their body are given
later under “Economic Importance” and Table 62 (р. 838).

595

596

803

Fig. 344. Section of the head of the sperm whale, Physeter catodon (figure by

N.N. Kondakov): 1—blowhole; 2—left nostril; 3—right nostril; 4—upper sper-

тасей sac [spermaceti organ or “сазе”]; 5—lower зрегтасей sac [“melon” or
“Junk”; 6—sinew-tendon layer.

20 to 60 cm long. The eyes are roughly at the center of the head at the
level of the posterior corners of the mouth. The section of the orbital
aperture is 6-12 cm long and the diameter of the orb 15-17 cm. Small
(up to 1 cm long) crescent-shaped ear openings are located posterior to
the eyes and slightly below them.

The head is set off from the trunk by a faintly perceptible neck. The
trunk attains maximum thickness in the zone of the flippers but thins
out gradually thereafter. The flippers are short and broad, with rounded
outer edges. The large lobes of the caudal flukes are separated by a
deep notch. Two to six very low tubercles are located posterior to the
humplike dorsal fin. A rather low leathery keel runs along the lower side
of the caudal stem.

The integument of the sperm whale is quite thick (Table 58).

In some sperm whales the thickness of the abdominal skin [epidermis
plus dermis] may go up to 50 cm. The thickest epidermis and the densest

7 14227
O/ VLE
2 OM as
4 7. РА

й

Fig. 345. Sperm whale, Physeter catodon (ventral view) (figure by М.М. Kondakov).

596

597

804

Fig. 346. Head of a male sperm whale, Physeter catodon (figure by М.М. Kondakov).

Table 58. Skin [epidermis plus dermis] thickness in different sections of the trunk of
the sperm whale (in cm) (V.E. Sokolov)

Sex Date of length, Abdomen Dorsum

birth m

Body Lateral
ee ee ee ee АЗ 08 рате
Under Around Before Above Anterior around of head

flippers navel vulva _ flippers to flippers
dorsal
fin
Female Aug. 7 10.2 15 10 = 21 — 6.5 6
Female Aug. 7 9.8 18 14.3 22 20 25 95 8.5
Female Aug. 7 9.3 16 11.5 15.5 12 16 10 7.5
Мае Sept.13 148 31 30 = 38 33 22 8

collagen fiber fascicles in the dermal layer occur on the anterior surface
of the head. This is explained not only by the fact that this part of the
head faces maximum resistance against water while swimming, but also
by the fact that males in the period of mating butt each other with their
“foreheads” just as rutting rams do (Sleptsov, 1955). The epidermis of
the sperm whale is characterized by prominent projections of epidermal
barriers extending into the dermal papillae. These projections, in many
cases, have thickened distal and thin proximal sections. On the dorsum
and flanks the skin surface is corrugated. In the depressions between the

597

598

805

tubercles, the epidermis forms projections into the dermal layer. On the
throat 10 to 40 longitudinal furrows occur, up to 1 cm in depth and 5 to
50 cm in length (Tomilin, 1957). It is probably because of these that the
throat can expand slightly while swallowing a large quarry (Beddard, 1900).

A typical leathery callosity on the dorsal fin is regarded as a sec-
ondary sex-related feature of females (Kasuya and Ohsumi, 1966).

The body color is monochromatic dark (black, blackish-brown,
blackish-cinnamon) with a much lighter-colored abdomen bearing an
irregular white patch. The outline of this patch varies sharply in different
animals. The sperm whales encountered in the waters of Japan can be
classified into four color groups (Omura, 1950b): (1) with continuous
dark gray color all over the body; (2) with a slightly lighter coloration on
the lower side of the head and the lower jaw; [(3) with a light coloration
on the lower side of the head and the lower jaw—this is an addition to
Omura’s classification]; (3) with a light coloration all over the head; and
(4) with a very light coloration all over the trunk. Animals of the third
group are the rarest. Sperm whales caught off the Kuril Islands are mainly
of two colors, dark and light, and form no less than eight variations of the
white pattern (Ivanova, 1959*). With age, the color of the animals turns
somewhat lighter. Many white and gray patches, bands, scratches, etc.,
formed as a result of skin infections caused by ectoparasites, lampreys,
and suckers and hooks [jaws resembling the parrot’s beak] of squids are
seen On the skin surface. Such patches are more among females than
males (Omura, 1950b).

In the sharply asymmetric skull the upper lateral sections of the
maxillae form crests laterally surrounding the base of the spermaceti
sac. The horizontally disposed palatines are covered from the rear by the
pterygoids. The halves of the lower jaw form a symphysis constituting
up to 50% (or slightly more) of the entire jaw length (Fig. 349). The
relative length of the rostrum and mandibular symphysis increases with
age (Tomilin, 1957).

Cervical vertebrae 7, thoracic 11, lumbar 8, and caudal 24. Infor-
mation on the fusion of the cervical vertebrae is contradictory. Some
scientists assume that the cervical vertebrae of the sperm whale fuse
into two independent groups of 1 to 3 and 4 to 7 vertebrae (Sleptsov,
1955). According to other authors, the sperm whale represents a unique
case among cetaceans with its atlas remaining free while all the other
vertebrae are fused (Tomilin, 1957). The level of the transverse processes
of the lumbar vertebrae is much lower than that of the thoracic.

In the five-digit limb (Fig. 348) the humerus is sometimes fused
with the ulna and the radius. Phalangeal formula: I,, I, HI;, 1V4, and
V3 (Beddard, 1900).

&06

597 Fig. 347. Head of a female sperm whale, Kuril Island, 1962 (photograph by
A.V. Yablokov).
Ge
S
:
598

Fig. 348. Flipper of the sperm whale, Physeter catodon (figure by N.N. Kondakov).

The peg-shaped homodont teeth of the sperm whale, often slightly
curved midlength, can reach large proportions, up to 27 cm in length
(Yablokov, 1958a), with the largest occurring in the middle of the jaw.
The teeth are larger among males than females. Large teeth are encoun-
tered only in the lower jaw. The teeth on the upper jaw are small, not
strong, or may be altogether absent. The number of upper teeth among |

599

807

Fig. 349. Skull of the sperm whale, Physeter catodon (figure by М.М. Kondakov).

males varies from nil to 19 and among females from nil to 10 (Sleptsov,
1955). The tooth crowns in embryos may have three cusps (Nishiwaki,
Hibiya, and Ohsumi, 1958). The teeth of calves usually emerge by the
end of the lactation period. With advancing age, the size of the teeth
increases considerably.* Mandibular teeth begin to wear out a few years
after birth (the anterior and middle ones in the first instance), some-
times right up to the base, although the upper and lower teeth may not
come into contact with each other. The gaps between the teeth vary from
3.5 to 15 cm. The teeth are firmly set in the alveoli and rise above the
gum level only to one-sixth to one-fifth of their length; intense growth
of cement is a characteristic feature. The volume of the pulp cavity can
measure up to one-half that of the tooth. The teeth of adult animals
have no enamel layer (Yablokov, 1958a).

The total length of the largest male caught was 20.7 m, of the female
15.8 m. The average length of 2,510 males caught in the Northern hemi-
sphere was 14.6 m and of 15,526 males caught in the Southern hemi-
sphere 15.9 т; the average length of 1,190 females (waters of Japan) was
10.64 m (Tomilin, 1957).

ЗА method was suggested for determining.:the age of sperm whales by counting the
number of layers in a longitudinal slice of tooth (Nishiwaki, Hibiya, and Ohsumi, 1958).

‘599

808

The average main body measurements as percentage of body length
among males and females caught off the Kuril Islands (Tomilin, 1957)
are: distance from tip of snout to blowhole in males 2.30 (two mea-
surements) and females 1.50 (2); distance from anal opening to notch
between caudal flukes 30.30 (94) and 32.20 (46); height of dorsal fin 2.40
(147) and 2.60 (58); length of dorsal fin 8.00 (145) and 8.10 (62); length
of flippers 9.6 (147) and 9.7 (74); maximum width of flippers 4.90 (26)
and 4.90 (20); anterior-to-posterior width of caudal flukes 9.40 (192) and
9.70 (88).

The main skull measurements (see Fig. 349) of three male sperm
whales (Tomilin, 1957; Omura et al., 1962) are (in cm): condylobasal
length 510, 450 and 360; interorbital width 220, 170; length of rostrum
370, 320, and 250; width of rostrum at base 150, 170, and 120; and
length of lower jaw 450, 300 [The authors gave the skull measurements
in decimeters instead of centimeters. I have added a zero to them to
convert them to centimeters—Ed.] (V.S.)

Geographic Distribution

Sperm whales inhabit the warm and temperate zones of the World Ocean
and are encountered almost everywhere. The summer ranges of males
and females are diverse: the former are encountered at very high lati-
tudes, reaching the arctic zone in the Southern hemisphere; the latter
do not emerge beyond the temperate waters throughout the year.

Geographic Range in the USSR

Constitutes a small section of the total range of the species. These ani-
mals are encountered in the basin of the Atlantic Ocean in the southern
part of the Barents Sea: along the Murman coast, possibly up to Kanin
Peninsula in the east (Fig. 350). In the Pacific Ocean they are distributed
in the Sea of Japan where they are sometimes sighted in Peter the Great
Gulf and even transgress into Zolotoi Rog Bay. They are encountered
in the waters of the Kuril Islands, predominantly from the Pacific Ocean
side. Large males inhabit the Sea of Okhotsk, mainly its northeastern
section. Farther northward, these animals are known in Kronitskiy and
Kamchatka gulfs off the southeastern coast of Kamchatka and around
the Commander Islands. They have been sighted off Cape Navarin in the
southern part of the Gulf of Anadyr.

Geographic Range outside the USSR

Distributed almost everywhere in the North Atlantic Ocean (Fig. 351).
Throughout the year, sperm whales are sighted off the Cape Verde,

809

‘(aa,uasIV “W'A) USSN ЭЦ: Ш! ‘Uopowws sajaskyg ‘этеца шло@з ay] Jo э8иеы “OSE ‘9

009

“(лэ.иэзлу У А) иоро1р> dajaskyg ‘aye шлэ4$ ay) Jo э8иел $910э4$ ‘16$ “314 109

SSS SSS
ШУ 000$ 000% 0006 0002 000: 0 0001

602

811

Canary, Azores, and Madeira islands and along the coasts of Spain and
Portugal, off the Antilles and Bermuda islands, and along the coasts of
Florida, i.e., up to 520- 15° N lat. They are quite common in the Mediter-
ranean Sea. They are not infrequent in the waters of Great Britain, off
the Orkney and Shetland islands, widely distributed along the coasts of
Norway, in the waters of Iceland, and off the Faroe Islands. Data on
their distribution along the American mainland between Florida and
Newfoundland are altogether lacking although some random observa-
tions suggest that these animals are encountered here. In the North
Atlantic, sperm whales reach Newfoundland, Labrador, Davis Strait, and
the coasts of southern Greenland. They inhabit the waters of Jan Mayen
(70° N lat.) and even Spitsbergen (around 80° N lat.).

Information on the distribution of sperm whales in the equatorial
section of the Pacific Ocean is highly approximate. They have been
assumed to be quite common close to Kalimantan, Celebes, Mariana,
Marshall, and Bonin islands, and are possibly encountered in the Sulu
and Banda seas, i.e., not only reach the equator, but sometimes even
the equatorial waters of the Southern hemisphere. The contemporary
regions of wintering of northern sperm whales are possibly bounded by
the line: Bonin Islands - Hawaii- California coast (Berzin and Rovnin,
1966), 1.е., within 15 to 20° М lat. Sperm whales are common along the
southern and eastern coasts of Japan but rare in the East China Sea
and in the southern part of the Sea of Japan. In the northeastern half
of the Pacific Ocean, sperm whales are distributed from California to
the Gulf of Alaska and the Alaskan Peninsula and along the islands
of the Aleutian range. In the Bering Sea, they are common in Bristol
Bay and in the waters of the Pribil of Islands, occupy the eastern part
of the sea bound by the shallow-water line running from Bristol Bay
to Cape Navarin on the Asian coast and probably up to St. Lawrence
Island.

In the Southern hemisphere, sperm whales occupy the entire zone
of tropical and temperate waters. In the Atlantic Ocean, they are known
from the coasts of Brazil to the Falkland Islands and Tierra del Fuego in
the south, and sighted in the waters around Tristan da Cunha, Ascension,
and St. Helena islands. In the eastern part of the Atlantic, they are com-
mon from the equatorial waters of the Gulf of Guinea to the southern
tip of Africa. In the Indian Ocean, the largest number probably inhab-
its the waters of southeastern Africa and Madagascar; they are known
off the Mascarene, Seychelles, Cocos, and Christmas islands, and along
the coasts of Australia. Penetrating northward, the distribution crosses
the equatorial zone and this animal is known in the Arabian Sea and
transgresses into the Red Sea. In the southern half of the Pacific Ocean,

603

812

sperm whales are abundant along the coasts of Peru and common in
the waters of Chile. They are encountered close to numerous islands,
including Fiji, Samoa, etc., and are common in the waters of Australia
and New Zealand. In the summer months male sperm whales are widely
distributed in the Antarctic waters and reach the southern Arctic zone
(Kirpichnikov, 1950b; Tomilin, 1957; Berzin and Rovnin, 1966; Berzin,
1971).

In spite of the fact that the two ranges of the species—northern and
southern—are distinct, there is no real demarcation in the distribution
of sperm whales. They are encountered in the equatorial zone of the
Northern and Southern hemispheres throughout the year and hence it is
very difficult to draw the boundary of distribution in either hemisphere.
While a demarcation is possible in the Atlantic Ocean (though highly
approximate), it is difficult to do so in the Pacific Ocean. Apparently
the equatorial zone serves as the area of contact between the northern
and southern populations of the sperm whale but information is not
sufficient to resolve this problem. (V.A.)

Geographic Variation

A comparative morphological study of sperm whales from different
parts of the World Ocean revealed no significant differences in the
body proportions (Ivanova, 1955, 1961). However, some difference in
the overall dimensions does exist between the northern and southern
sperm whale populations (Nishiwaki, 1955). Based on this and the time
differential in the main biological rhythms of the whales of the North-
ern and Southern hemispheres, two subspecies of sperm whales are
recognized (Tomilin, 1957). Only one of these subspecies is found in
the USSR.

Northern sperm whale, P. c. catodon Linnaeus, 1758 (syn. macro-
cephalus).

Animals of relatively smaller dimensions, with the males measuring
an overall length of 14.6 m (North Pacific Ocean).

They are encountered in the Barents, Japan, Okhotsk, and Bering
seas, and the Pacific Ocean.

Outside the USSR, they are found in the North Pacific and Atlantic
oceans.

Systematic relationships between the sperm whales of the North
Pacific and Atlantic oceans have not been studied. Evidently these pop-
ulations are highly isolated (amphiboreal distribution) and possibly may
not be identical. The name sulcatus Lacépéde could be applied to the
Pacific population.

813

Outside the USSR, some scientists recognize the form Р. с. aus-
tralasianus Demoulins, 1827,* characterized by very large dimensions
(average length of males 15.9 m).

These animals are encountered in the warm and temperate waters of
the Atlantic and Pacific oceans in the Southern hemisphere, the Indian
Ocean, and the Antarctic.

The interrelations between the two subspecies of sperm whales in the
equatorial zone, especially in the Pacific Ocean, have not been studied.
It is possible that many local herds of sperm whales exist in different
parts of the species range; sometimes such herds can be localized in a
relatively small water body (Klumov, 1955) (see “Seasonal Migrations
and Transgressions”). (V.A. and V.H.)

Biology

Population. The sperm whale can be regarded as one of the more abun-
dant species of large whales. The population is perhaps most abundant
in the Pacific Ocean. The probable population of the sperm whale in the
North Pacific Ocean has been put at roughly 150,000 (Nishiwaki, 1966).
Up to 4,000 - 5,000 sperm whales were caught in some years off the coasts
of Chile and Peru, indicating a large population of these whales in the
South Pacific Ocean; they are also abundant in the other waters of the
Southern hemisphere. Thus, in some years in the Antarctic alone, up
to 6,000-7,000 male sperm whales were caught. Insofar as the North
Atlantic is concerned, even approximate population data are not avail-
able. It is probable that the population here is relatively small, which
is Supported by the comparatively low level of hunting. The maximum
number of sperm whales is caught in the waters of the Azores where the
total catch sometimes does not exceed 1,000 per year.

Food. Two groups of animals, cephalopods and fish, constitute the
main food of sperm whales in both the Northern and Southern hemi-
spheres, with cephalopods predominating. The food of sperm whales has
been studied in great detail in the North Pacific Ocean. Data for the other
regions of the World Ocean are fragmentary and incomplete (Table 59).

Three species of squids predominate in the food of the sperm
whale from the Bering Sea: Gonatopsis borealis, Gonatus magister, and
Moroteuthis robusta; Meleagroteuthis separata is of lesser importance
(Table 59). Among the fishes, redfish occupies first position of
importance as a food item. It is followed by the smooth lumpsucker

1 А.С. Tomilin (1957) designated this form as P. с. australis MacLeay, 1851, which is
hardly correct (see synonyms and note on p. 801).

604

814

Bering Sea (Berzin, 1959)

Cephalopods

Gonotopsis borealis
Gonatus magister
Gonatus fabricti
Moroteuthis robusta
Meleagroteuthis separata
Galiteuthis armata
Onychoteuthis banksti
Chiroteuthis veranyi

Fish

Smooth lumpsucker,
Aptocyclus ventricosus
Perches, Sebastodes
(few species)

Rat-tails, Coryphaenoides sp.

Lancetfish, Alepisaurus
aesculapius
Skates, Raja (two sp.)

Pacific lamprey, Entosphenus

tridentatus

Sculpins, Myoxocephalus sp.

Table 59.

British Columbia (Pike, 1950b) Kuril Islands (Betesheva and

Cephalopods

Gonatus fabricii
Moroteuthis robusta

Fish
Sea perch:

Sebastodes ruberrinus
Sebastodes sp.

Ragfish, [costeus aenigmaticus

Skate, Raja rbina

Arctic lamprey, Lampetra

Japonica

Salmon, Salmonidae

Akimushkin, 1955; Akimushkin,
1957; Betesheva, 1960,
1961; Tarasevich, 1963)

Cephalopods

Gonatus magister
Gonatus fabricii

Taonius pavo
Moroteuthis robusta
Onychoteuthis banksii
Architeuthis japonica
Meleagroteuthis separata
Chrioteuthis veranyi
Galliteuthis armata
Stigmototeuthis dofleini
Octopodoteuthis longiptera
Crystoloteuthis berhingiana
Paroctopus conispadiceus
Japatella heathi
Grimpoteuthis albatrossi
Alloposus mollis

Octopus gilbertia
Cirroteuthis sp.
Amphitretus sp.

Octopus sp.

Octopodidae sp.

Fish

Black rat-tail:

Coryphaenoides acrolepis
Albatross rat-tail, C. pectoralis
Piked dogfish, Squalus acanthias
Lancetfish, Alepisaurus aesculapius
Pacific saury, Cololabis saira
Pacific cod, Gadus morhua
macrocephalus

Alaska pollock, Theragra
chalcogramma

Smooth lumpsucker, Aptocyclus
ventricosus

Humpback salmon, Oncorhynchus
gorbuscha

Navaga, Eleginus navaga gracilis
Podonema, Podonema longipes
Sharks, Somniosus sp.
Anglerfish, Oneirodes sp.
Sculpins, Cottidae gen. sp.
Salmon, Oncorhynchus sp.
Greenlings, Pleurogrammus sp.
Skates, Raja sp.

815

Food objects of sperm whales

Japan (Mizue, 1951b) Azores Islands (Clarke, 1956) Antarctic (Korabel’nikov, 1959)

Cephalopods
Squids
Fish

Pacific cod, Gadus
macrocephalus

Alaska pollock, Theragra
chalcogramma

Scorpionfishes:

Sebastodes flammeus
Sebastodes iracudus

Pacific saury, Cololabis saira

Pacific sardine, Sardinella
melanosticta

Round herring, Etrumeus
micropus

Japanese anchovy, Engraulis
Japonica

Pacific mackerel,
Pneumotophorus japonicus
Spotted mackerel, Scomber
tapeinocephalus

Sharks, Selachii

Cephalopods

Histioteuthis bonnelliana
Cucioteuthis unguiculatus
Tetronychoteuthis dussumierii
Lepidoteuthis physeteris
Ancistrocheirus lesneuri

Loligo forbesi

Architeuthis sp.

Fish

Long-finned tuna (albacore),
Thunnus alalunga

Anglerfish, Ceretias holbolli
Atlantic footballfish,

Himantolophus greenlandicus

Barracuda, Sphyraena sp.

Cephalopods

Onychoteuthis banksii

Architeuthis sp.

Squid (genus not identified)
Fish

Anglerfish, Ceratias holbolli

Patagonian toothfish,
Dissostichus eleginoides

Southern putassu,
Micromesistius australis

Skate, Raja griseocaudata

604

816

which, however, is encountered in the stomach of sperm whales more
often than any other fish. The ratio of the various groups of animals in.
the food of the sperm whale is as follows: of the 107 stomachs with food,
69 (64.4%) contained only squids, 28 (26.2%) squids and fish, and 10
(9.4%) only fish. The importance of cephalopods and fish in the food of
sperm whales undergoes variation relative to the region of habitation.
In the waters of the Commander Islands, 74.4% of the stomachs anal-
ysed contained squids exclusively, 23.9% squids together with fish, and only
1.4% fish exclusively; for the region of the western islands of the Aleutian
range, these values were respectively: 54.0, 26.9, and 19.1%, and in Olyu-
torskiy Gulf and adjoining waters 35.7, 35.7, and 28.6% (Berzin, 1959).
The stomach of sperm whales from the waters of the Kuril Islands
(360 stomachs) contained 28 varieties of cephalopods (21 squids and 7
octopuses). Cephalopods constitute about 95% (by weight) of the food
intake of the sperm whale and fish less than 5%, although the latter were

_ detected in almost one-third of the sperm whales investigated. Squids are

605

606

of basic importance among cephalopods; octopuses constitute no more
than 4% of the intake. But only seven species of squids are of prac-
tical importance as food: Gonatus magister, G. fabricii, G. simile, Tao-
nius pavo, Galliteuthis armata, Chiroteuthis уетапу, and Maleagroteuthis
separata constitute up to 80% of the cephalopods consumed, with 60%
accounted for by the three species of the Gonatidae family alone. Cuttle-
fish, abundant in the Far Eastern seas, have not been found in the stom-
ach of the sperm whale (Betesheva and Akimushkin, 1955; Akimushkin,
1957; Betesheva, 1960, 1961).

In the waters of Japan an analysis of 1,627 stomachs showed that squids
represent the main food here too, although octopuses were found quite
often but in small quantities. All the other food objects were comparatively
rare. The male sperm whale feeds on a more diverse diet than the female
(Mizue, 1951). The species of cephalopods were not indicated in this report.

In the waters of British Columbia, where only 50 stomachs were
investigated, squids were similarly found to be of primary importance.

Frequency of Encounter (Number of Stomachs) of
Various Objects in the Food of Sperm Whales from
the Waters of British Columbia (Pike, 1950b)

Squids 35
Ragfishes (Icosteidae) 16
Sea perch 16
Skates Э
Salmon (?) 3

Lamprey 1

606

817

Squids serve as the main food of Antarctic sperm whales too: fish
was found in only 6 (5.2%) of the 129 stomachs investigated (Kora-
bel’nikov, 1959). According to the latest data, the large Antarctic tooth-
fish is often found in the stomach of sperm whales from various regions
of the Antarctic (V.L. Yukhov).

Cephalopods have been recorded as the main food of sperm whales
from many other regions of the World Ocean: the North Atlantic, the
Mediterranean Sea, tropical zone, and along the coasts of Australia
(Tomilin, 1957) (see Table 59).

Sperm whales consume mainly comparatively small squids, 30-40 to
100 cm and less often 150 cm long, but sometimes the stomachs have
contained giant animals. Thus a sperm whale 14.3 m long killed in the
Azores in 1955, contained a mollusk (with tentacles) 1,049 cm long,
weighing 184 kg (Clarke, 1956) (Table 60).

In the Azores the main food of males as well as females consists of
comparatively small squids of nearly identical size even though the male
is considerably larger than the female. Sponges, crabs, crustaceans, and
even bits of the skin of pinnipeds, sometimes found in the stomach of
sperm whales, are merely items swallowed incidentally.

Stomachs with a large volume of food are relatively rarely seen. Of
the 264 stomachs (Kuril Islands) analyzed, 13 (4.9%) were well filled,
28 (10.6%) were moderately full, 125 (47.4%) contained little food, and
98 (37.1%) were empty (Tarasevich, 1963). The weight of the stomach
contents of 120 sperm whales showed 33% with 4 to 10 kg of food, in
some cases going up to 200 [sic] kg (Betesheva and Akimushkin, 1955).
Most often, the stomachs revealed only the “beaks” of squids, sometimes

Table 60. Sizes of squids consumed by sperm whales (Clarke, 1956)
Squids found
Mantle length Male sperm whales Female sperm whales
of squids, cm wl es el УРНЫ
Number % Number %
60-90 41 59 28 67
90-150 26 37 13 31
150-180 2 3 = =
180-240 1 1 1 2
Total 70 100 42 100
Average length 95 92

of squids, cm

607

818

more than 10,000. The maximum number of beaks found was 28,000,
corresponding to the consumption of 14,000 squids. This, however, does.
not represent a one-time intake since beaks are digested very slowly (or
not at all) and probably accumulate over a rather long period.

It is interesting that the squid Ommatostrephes sloanei-pacificus,
inhabiting the upper horizons of the sea and most abundant in the Far
Eastern waters, is of almost no importance in the food of the sperm
whale and that the abundant cuttlefish, living close to the surface, is
not touched at all. The main food consists of bathypelagic species of
cephalopods and fish, most of which do not rise to a depth of less than
500 m and some of which live at depths of 1,000 m or less (Akimushkin,
1954b). Thus, sperm whales catch their quarry at depths of not less than
300 to 400 m, where these whales have almost no food competitors with
the possible exception of beaked whales (Ziphiidae). In search of food,
sperm whales can descand right to the ocean floor since their stomachs
sometimes contained bottom-dwelling deepwater animals.

In the 1960s, with the increasing geographic spread of sperm whale
hunting, more information has become available about the food of these
whales in regions earlier not studied. In the open waters of the northeast-
ern part of the Pacific Ocean, the predominant contents of the stomach
of sperm whales were the remains of squids, mainly Taonius pavo, Chi-
roteuthis veranyi, Meleagroteuthis separata, Galiteuthis armata, and Gona-
tus magister. Among fish, the following were encountered: Alepisaurus
aeusculapius, Pseudopentaceres richardsoni, Sebastodes alutus, and Aptocy-
clus ventricosus. The stomach of sperm whales from the waters of central
California revealed the squids Moroteuthis robusta and Gonatus borealis,
and among fish, the sharks Apristurus brunneus and Squatina californica,
and sablefish, greenling, and lanternfish (Tarasevich, 1968; Berzin, 1971).
In the waters of New Zealand and the nearby islands, squids were found
in 84% of the stomachs studied. Remains of scorpionfish, skates, sharks,
some invertebrates (fire salpians, shrimps) and even brown algae were
detected. In the southern part of the Indian Ocean the stomach of sperm
whales predominantly contained squids, including large ones (up to 9 m
[sic] in length). Barracuda and sometimes porcupinefish were among the
fishes frequently encountered in the stomach contents.

On the whole, the main food of the sperm whale every where is
cephalopods (squids up to 80% of intake and octopuses), including
roughly 40 species and constituting not less than 95% of the total weight
of the food intake. The stomach of sperm whales also contained the rem-
nants of over 50 species of fish but, in spite of this diversity, fish account
for no more than 5% of the total intake. Most of the animals serving as
food objects of the sperm whale are deepwater species (Berzin, 1971).

608

608

819

Relatively often, the stomach of sperm whales revealed altogether
extraneous matter (Fig. 352): rubber boots, wire reels, glass fishing buoys,
rubber gloves, plastic toys (motor cars, pistols, dolls, pails, etc.), jugs,
plastic bags, coconuts, vinyl chloride bags, fishing tackle with hooks,
empty bottles, apples [sic], and many others (Berzin, 1971).

Daily activity and behavior. Sperm whales are polygamous animals
with sexual dimorphism distinctly manifest in their dimensions. In warm
waters they form groups consisting usually of 10 to 15 females and a large
male. But groups of hundreds of animals are not uncommon. Adult males
that are not members of these groups (harems) remain aloof, often indi-
vidually; they gather into relatively large groups only sometimes at food
sources. The latest observations have shown that the males included in
harems are not the largest and oldest but much younger males of 13 to
14 m length with high sexual prowess. They are the leaders of harems
and drive away the old males, who subsequently remain aloof. All of
this calls for a detailed review of the contemporary concepts of harems,
their composition, behavior, interrelationships between individual age
and sex groups in small and large collections, and other aspects of fam-
ily and herd behavior of the sperm whale (Berzin, 1971; V.A. Zemskii,
D.D. Tormosov, Yu.A. Mikhalev).

Fig. 352. Foreign objects recovered from the stomach of sperm whales (photo-
graph by A.A. Berzin).

609

820

In the regions of summer habitation, depending on age and size of
animals, male sperm whales generally form groups of definite composi-
tion. The groups most often consist of comparatively same-sized animals
rather than animals of different sizes. Of the 23 groups analyzed in the
Gulf of Alaska, 18 (78.3%) consisted of whales of nearly similar size
and only five (21.7%) of animals of different sizes. It has been suggested
that the reason for animals of the same size grouping together lies in
their identical abilities for getting at food available at different depths.
Large whales can dive deeper and feed at places where much smaller
animals cannot. The same-sized groups of very small whales usually con-
sist of nearly same-aged animals (with a difference of one or two years).
In groups consisting of much larger whales, the age differences of the
members are greater (sometimes 10 years). This factor probably deter-
mines the high individual variations in the dimensions of adult whales
whose growth has ceased (Tarasevich, 1967a).

The nature of diving of the sperm whale is typical they can remain
submerged under water for up to an hour and sometimes even longer
(two hours is considered maximum). Before prolonged submergence, the
sperm whale dives sharply, its body is steeply curved or humped and the
bent caudal stem is visible on the sea surface. The animal dives almost
vertically and, in most cases, the caudal flukes are displayed above the
water. The appearance of this “butterfly form” above the sea surface
serves as a positive indication that the whale has submerged for a long
time and may surface very far from the site of submergence.

- The maximum depth of submergence of the sperm whale has not
been established. Nineteen instances are known of damage to underwater
cables caused by sperm whales entangled in them, exclusively in tropical
and temperate waters (between 46° S lat. and 46° N lat.). The depth at
which dead sperm whales were detected ranged from 118 to 1,116 m.
Having become entangled in the cable, the sperm whale grips it with
the teeth, tears the insulation, and thereby damages it. In 1951, a cable
connecting Lisbon and Malaga revealed this type of damage at a depth
of 2,200 m, so far the maximum known depth. In most cases, the depth
of cable damage exceeded 500 m, sometimes 1,000 m. Most often, the
lower jaw and caudal fin of the sperm whales were entangled in the
cable. It has been assumed that the whale held the cable in its mouth
while chasing for food and then began to whip it when the slackened
cable wound around it. It has been suggested that sperm whales become
entangled at low depths and then gradually slip down along the slope of
the sea bed. However, the nature of the entanglements revealed that they
occurred at the places where the animals were actually found (Khizen,

609

821

1957; Yablokov, 1962). Sperm whales perhaps are actually capable of
descending to a depth of 1,000 m or more.

After prolonged residence under water, the sperm whale rests a long
time on the surface (Fig. 353). At this time, it remains almost stationary,
only slightly moving forward; in a horizontal position, it rhythmically
submerges and blows every 10 to 15 sec. The number of blows varies and
probably depends largely on the time spent under water. Usually there
are 20 to 40 blows but up to 75 in some cases. Compared to the older
animals, the young ones produce fewer blows. The former evidently dive
deeper and for a longer duration. During this characteristic rest period,
the sperm whale remains very quiet and unconcerned, lying on the surface
for 10 min or longer; in most cases, a whaling boat can approach such
a whale within shooting range. During hunting around Antarctica there
have been instances of a whaling boat literally approaching a sperm whale
so close as to hit the animal with its stempost (V.A. Arsen’ev).

Sometimes a sperm whale rising from the deep emerges almost ver-
tically from the water, thrusting about half of its trunk clear. In other
cases, the animal leaps high at an angle to the sea surface and falls on
the water with a loud splash.

The speed of the sperm whale is comparatively slow. The so-called
moving or migrating whales (traveling from one region to another)
usually travel straight at 5 to 7 miles/hr. They swim close to the water
surface, often make shallow dives, and produce many blows. Feeding

Fig. 353. Sperm whales at sea (figure by NN. Kondakov).

610

612

822

animals move more slowly, probably at not more than 3 miles/hr. When
searching for food, they remain submerged a long time, changing course
under water regularly and very sharply. A frightened or wounded sperm
whale moves at maximum speed, quite often along a straight course, and
produces blows frequently. The maximum speed of a large sperm whale
hardly exceeds 10 miles/hr. In any case, ships covering 12-12.5 miles/hr
invariably overtake an escaping sperm whale after 40-50 min of pursuit.

A sperm whale wounded by a harpoon sharply dives deep but soon
reemerges on the surface, rising almost vertically, and displaying its huge
obtuse head. If the movement of the whale boat is not arrested in time,
the surfacing whale can appear underneath it and damage the bottom
severely by colliding against it with its head. Instances are known of such
situations wherein the whale damaged the whaling boat by distorting the
propeller blades or even the crest of the shaft. These cases formed the
basis for the widespread belief that a wounded sperm whale often actively
attacks a ship. Such attacks are no doubt unintentional and inflicted at
random by the surfacing sperm whales (V.A. Arsen’ev).

Instances of sperm whales being beached, either singly or in groups,
are not infrequent along the coasts of Denmark, France, Florida, Cal-
ifornia, Tasmania, and New Zealand. Eight cases have been registered
of groups of beached whales ranging from 16 to 36 animals. Sometimes

Fig. 354. Typical blows of the sperm whale. Pacific Ocean (photograph by
M.M. Sleptsov).

613

823

such groups consisted exclusively of females but often of lone males.
Twenty-two males cast up on January 16, 1954 on the coast of California
were roughly identical in size (Gilmore, 1957; Tomilin, 1962) (Fig. 355).

Seasonal migrations and transgressions. The nature and courses of
migration of sperm whales have not been thoroughly studied. The scheme
of migrations is generally as follows: in the winter months, most of the
sperm whales gather in warm and tropical waters and move in sum-
mer into temperate waters, although some males even depart into the
cold waters of the Northern and Southern hemispheres. The main fac-
tor determining the courses of migrations and the distance of travel of
sperm whales in cold waters is the effect of such currents as the Kuroshio
in the Pacific Ocean and the Gulf Stream in the Atlantic. The zone of
influence of these currents provides favorable conditions for the survival
of the main food objects of the sperm whale, 1.е., cephalopods, whose
entry into the north in summer months is determined by the intensity
and direction of the currents. Sperm whales move into relatively high
latitudes in pursuit of these cephalopods.

Early in spring, in the North Pacific Ocean, sperm whales begin to
migrate from the warm waters northward. Around the Bonin Islands
the first females are sighted February end and by March their catch is
maximum. Most of the whales migrate along the eastern coast of Japan.
Some whales migrate northwest and are caught in small numbers on the
western coast of Kyushu Island. They are sometimes encountered in the
East China Sea. Sperm whales are not caught off the Korean coasts but

Fig. 355. A group of beached sperm whales. California (from R.M. Gilmore, 1956).

611

824

some numbers enter the Sea of Japan from the south (Omura, 1950).
Some lone sperm whales, quite rare in fact, are noticed in Peter the
Great Gulf and sometimes transgress even into the Bay of Zoloti Rog
(Vladivostok). In the waters of southern Japan, sperm whales are seen
early in spring, up to April; the earliest to arrive here are young, already
mature males. In June-July, most of the sperm whale herds*concentrate
in the waters of Japan; at this time, males, females, and calves live here.
In July, they move northward but the groups break up in the waters north
of 45° М lat. Males separate from the general groups and go northward
while females with calves inhabit the much warmer waters. The hunting
of sperm whales in the waters of Japan is most fruitful from June through
November (Nishiwaki, 1966).

In the central part of the Kuril range, the first sperm whales, excep-
tionally large single males, are sighted early in April and groups begin
arriving by May. By mid-May, herds of sperm whales do not actually
transgress north of the 48th parallel.

In the northern part of the Kuril range, lone male sperm whales
are seen at April end to early May and by May end the so-called harem
whales begin to arrive. By mid-June, stable collections, called herds, of
sperm whales of different sexes and ages are formed here. Large males by
this time move farther northward in large numbers (Tarasevich, 1965). In
the first half of June, mixed groups of sperm whales are fairly uniformly
distributed from the Pacific Ocean side of the middle and northern sec-
tions of the Kuril range and their herds depart farther from the coasts.
Adult lone males by this time are seen in the Sea of Okhotsk, which is
not at all visited by females or young animals. In July, the number of
sperm whales around the Kuril Islands is far smaller and only minimal
groups are seen throughout the region. It can be assumed that herds of
sperm whales often travel at this time toward southeastern Kamchatka
and are partly scattered over this immense water body.

At the end of July in the central part of the Kuril range (Bussol
Strait-Simushir Island), new herds of sperm whales of medium size, con-
sisting mainly of young males, are seen approaching from the south.
At the very end of July, new herds are again seen in the region of
Urup—Simushir. These herds contain many females and some large
males. These are clearly the returning harems. During the first 20 days
of August sperm whales remain in this region, even slightly extending
southward (up to Friza Strait), but do not advance northward at all. At
this time herds consist of very small animals but large whales begin to be
sighted again by the end of August. They probably now begin to move
out of the Sea of Okhotsk southward.

612

825

This pattern of arrivals confirms the assumption that during summer
two narrowly localized sperm whale herds, i.e., southern Kuril and north-
ern Kuril populations, inhabit the Kuril Islands. Whales of the southern
Kuril herd inhabit the northern parts of the Japanese Islands and the
southern parts of the Kuril range, reaching its middle in the north. The
northern Kuril herd inhabits the waters of the northern Kuril Islands
and is confined to the coasts of southeastern Kamchatka but may reach
the waters of the Commander Islands (S.K. Klumov).

In September, the maximum number of sperm whales is seen in the
Kuril waters since the whales returning from the northern waters now
begin to arrive. October is the month of en masse departure of sperm
whales southward and their herds around the Kuril Islands gradually thin
out. Individual animais or even small groups may, in favorable years,
remain for wintering close to the Kuril range (Sleptsov, 1955).

The northern boundary of distribution of adult females and young
sperm whales of both sexes is regarded as 51°N lat. (Omura, 1950)
although they are encountered even farther north, up to 52-54° N lat.
in Kronotskly and Kamchatka straits and around the Commander Islands
(Kirpichnikov, 1950b; Sleptsov, 1950; Tomilin, 1957). Hence, in favorable
years, females can reach even up to 53-54° N lat. The usual boundary of
distribution of adult males is 61 -62° М lat. (Cape Navarin) where whales
from both the western and eastern coasts of the Pacific Ocean live. It
is more probable that the region of Olyutorskiy Gulf (60° N lat.) serves
as the northern boundary of distribution for males of the Asian herd
although some stray whales have reached 65°30’ М lat. under favorable
conditions (Omura, 1950).

Migrations of the sperm whales of the American herd are as fol-
lows. From the wintering regions, disposed along a tentative line from
the Hawaiian Islands to the coast of California, sperm whales begin to
migrate northward at March end to early April. The period of migra-
tions is considerably prloonged and hence they are sighted even in May
in the southern parts of the range. The large extent of the wintering
range along the longitude makes for several courses of migrations. One
group moves along the coasts of the American mainland, another moves
within 145 - 150° W long., and yet another between 162 and 167° W long.
In April or early May, all these groups reach roughly 50° N lat. where
the comparatively linear course takes a complex route, At these latitudes
the males from the mixed groups finally separate out, with some ani-
mals moving into the middle and eastern sections of the Gulf of Alaska
and the Pacific waters of the Aleutian range, and others occupying the
northernmost part of the Gulf of Alaska, moving far westward right up
to the Commander Islands, entering-the Aleutian Straits into the Bering

613

826

Sea (Bristol Bay and Pribilof Islands) and, spreading along the east-
ern Shallow-water section of the sea, reach Cape Navarin and the Gulf
of Anadyr. These are exclusively males of various dimensions and ages
which gradually isolate from the mixed herds and gather in the regions
of food concentration, at times forming significant herds. It has been
assumed that the much younger, although sexually mature males are the
first to separate from the common.herds, followed by the larger older
animals. This is possibly associated with the concluding phase of repro-
duction. The northern boundary of distribution of groups of females
and young animals in all probability is 50-51° М lat. They are confined
predominantly to the Gulf of Alaska but are not infrequent in these
same latitudes around the Aleutian Islands, sometimes reach the west-
ern section of the range and, more rarely, even the Commander Islands.
Under favorable conditions, in some years small groups penetrate even
farther north and are seen in the Bering Sea (Berzin and Rovnin, 1966;
Tarasevich, 1967a). i

The return of groups of sperm whales to the wintering regions com-
mences in September. The sequence and periods of migrations of indi-
vidual age and sex groups have not been unraveled. Migrations proba-
bly occur by the same routes as when moving northward. In November,
the sperm whales practically abandon the northern regions of the range
although some stray animals or small groups may overwinter there. Such
instances are known even in the waters of the Commander Islands. Sperm
whales reside almost everywhere around the central and southern Kuril
Islands, and sometimes comparatively large herds are seen here in winter.
Wintering sperm whales have been recorded even in the Gulf of Alaska
(Zenkovich, 1936b, 1952; Sleptsov, 1955; Tomilin, 1957).

The periods of migrations of sperm whales in the North Atlantic
are roughly the same as in the Pacific Ocean. After overwintering in the
southern regions of the range, the whales begin to move north along the
western and eastern coasts of the ocean in early spring. Mixed groups
of sperm whales probably do not proceed north of 50°N lat. but mature
males of different ages reach Iceland, Greenland, Davis Strait, and Jan
Mayen and Spitsbergen islands. This is somewhat more northward than
in the Pacific Ocean due to the influence of powerful Gulf Stream cur-
rents. Some males, depending on hydrological and trophic conditions,
may sometimes overwinter in the north but most of the whales return to
the southern waters in autumn.

Regular seasonal migrations are also known among populations of
the Southern hemisphere but their extent differs for mixed groups and
adult males. Mixed herds of sperm whales apparently do not travel
beyond the south 40° latitudes since only one case has been recorded of a

614

827

female sperm whale being caught off South Georgia Island (54° 30’ $ lat.).
In the summer months, however, males are widely distributed in the
waters of Antarctica and reach the ice there—which is not the case in
the Northern hemisphere. In autumn males abandon the Antarctic Zone
and return to their points of wintering. Sperm whales have not been
observed at any time of the year in the Gulf of Guinea (Kirpichnikov,
1949b, 1950b).

Observations over several years in the various zones of the South-
ern hemisphere revealed that groups of small sperm whales, consisting of
immature males and females as well as adult females, remained mainly in
tropical waters. Large males were quite rare among them. The approxi-
mate body length of the small sperm whales was 7 to 10 m.

In the subtropical zone small groups of sperm whales, consisting of
males that had attained sexual maturity but were still not participating
in reproduction (approximate body length 12.0-12.5 m), were found.
Groups of males about to attain maturity (body length 11.0-11.5 m)
were encountered separately. The large males here were larger than in
the tropical region but nevertheless constituted an insignificant minority.

In the zone of temperate waters (40° latitudes) there were no imma-
ture males or small whales measuring 7-8 m in body length. Mature
males 12-13 m long predominated while large males constituted up to
20% of the total strength.

In the Antarctic waters (south of the 50th parallel), only large males
14-15 m long or more were encountered. Females and young sperm
whales of either sex were totally absent (Tormosov, 1970).

Reproduction. Female sperm whales usually do not emerge beyond
the warm and temperate waters and hence the seasons of mating and
parturition among them are not as sharply manifest as among those
whales whose females perform regular migrations into the cold waters
of both hemispheres. Births occur among sperm whales throughout the
year but the maximum number occurs in a relatively brief period. For
the Northern hemisphere, this evidently is the early autumn months. In
the North Atlantic, more births have been recorded from May through
November with the maximum number taking place from July through
September (Clarke, 1956).

On the eve of parturition, females concentrate in quiet zones where
conditions are more favorable for the newborn. Such zones in the Pacific
Ocean are the waters of the Marshall and Bonin islands, eastern coast of
Japan, and to a lesser extent the waters of the southern Kuril Islands and
the Galapagos Islands; in the Atlantic Ocean these are the waters of the
Azores and Bermuda islands and the coasts of the African provinces of
Natal and Madagascar. Sperm whales gather in regions with clean deep

615

828

water оп the leeward side of islands or reefs (Sleptsov, 1955; Tomilin,
1957).

In the Southern hemisphere, births occur from December through
April with a maximum in February. They take place in regions of quiet
and relatively warm waters, where sharks, killer whales, and other ene-
mies of the newborn sperm whales are few. The optimum water surface
temperature would seem to be 15-17°C. For the Atlantic and Indian
oceans, such waters are found between 35 and 39° S lat. and it is here
that the majority of female sperm whales undergo parturition.

In April, 1962, close to Tristan da Cunha Islands, the process of par-
turition was observed from a helicopter (F. Khomchik). Among several
groups of sperm whales, numbering 25-30 animals each, one group of
six was obviously isolated. These whales dived continuously together in
a row, churning the water, which soon became blood-stained and a new-
born whale was shortly sighted on the water surface. The newborn imme-
diately began to swim alongside its mother. The two were accompanied
by four other sperm whales, probably also females.

Observers have noted that during birth the female assumes a vertical
posture with almost one-quarter of the body length projecting above the
water. The caudal flukes in the newborn remain coiled for sometime in
the form of a small tube. Other adult females dive under the newborn
calf and help it to remain afloat on the sea surface until the caudal flukes
unfold (Tormosov, 1970).

Gestation in the sperm whale is far more prolonged than in most
of the other species of cetaceans. It has been assumed to extend for
16-18 months (Matthews, 1938b; Nishiwaki, Hibiya, and Ohsumi, 1958;
Ohsumi, 1965). The ratio of male to female embryos is close to 1:1. Of
the 1,068 embryos studied in the northern part of the Pacific Ocean,
males accounted for 48.1%. In the waters of Chile and Peru, of 1,118
embryos, males were 56.2% and in the waters of Africa (491 embryos)
45.6%. Of the total of 2,677 embryos studied, males accounted for 51.0%
(Ohsumi, 1965). Females usually deliver a single calf; twins are very rare
(0.66%, Clarke, 1956). The periodicity of reproduction has been put at
three (Clarke, 1956) or even three to four years (Ohsumi, 1965).

Growth and development. Embryonic growth of the sperm whale con-
tinues for up to 16 (probably even 18) months. Thus the growth of
embryos proceeds more slowly than among other species of whales, which
are characterized by a very short period of embryonic growth (Fig. 356).
The growth pattern of sperm whale embryos is depicted in Table 61.

The length of the newborn calves varies from 350 to 500 cm (Tomilin,
1957). The average size of the newborn from the North Atlantic is
392 cm (Clarke, 1956) and from the Southern hemisphere 415 cm (Lous,

829

ИРИ, ( 2
РСС

р 94472

р 2222

2222222
777 =

7
Е
a Се 77
4 Fe POO PA GLE

tt AZZ eg

Paar

7

614 Fig. 356. Embryonic growth of the sperm whale, Physeter catodon (figure by
N.N. Kondakov).

615 Table 61. Growth of sperm whale embryos (in cm) (Ohsumi, 1965; Berzin, 1971)

Month of growth Average length, according to
Ohsumi Berzin
151 6 10
2nd 24 20
3rd 52 30
4th 76 50
Sth 103 70
6th 127 90
7th 155 5
8th 183 140
9th 210 170
10th 234 215
11th 262 260
12th 289 290
13th 314 320
14th 341 350
1518 369 380
16th 393 400

1959). The average size of calves, based on data for both hemispheres,
is 405 cm (Ohsumi, 1965). The largest embryo from the North Pacific
Ocean was 460 cm long (Berzin, 1964a). A newborn calf can weigh up
to 1,000 kg.

The duration of milk suckling, on analogy with the other species of
large whales, was initially considered to be 6-7 months (Vinogradov,

616

830

1949; Sleptsov, 1955), later 13 months (Clarke, 1956), but according to
the latest data, 24-25 months (Ohsumi, 1965).

The age of the sperm whale has been determined by the number of
layers in the dentine formed as a result of seasonal metabolic variations
associated with changing conditions of habitation. Right up to the fill-
ing up of the pulp cavity, two layers of dentine are laid every year: one
very broad, light-colored, and intensely calcined, the other very narrow,
dark-colored, and less calcined (Fig. 357). These two layers form a sin-
gle annual ring whose number reflects the age of the whale (Nishiwaki,
Hibiya, and Ohsumi, 1958; Berzin, 1961). The question of the number
of layers laid in the course of a single year has not yet produced a com-
monly accepted opinion. It is possible that the annual rings may be two,
or a single annual ring may consist of different numbers of intermediate
rings. This aspect continues to be studied.

By the end of the first year, sperm whales of the North Pacific Ocean
reach a length of 6 m or slightly more, 1.е., the increment in body length
at the end of the first year is almost 2 m (all figures are average values).
The growth tempo subsequently slows down somewhat and by the third
year the animal reaches a length of 8 m, adding another 2 m in these two
years. By age 3.5 -4 years, the female sperm whales have attained sexual
maturity and their growth tempo subsequently is markedly slower than
that of the males, with the annual increment not exceeding 50 cm. The
onset of physical maturity and growth cessation among females occurs
at the age of 15 years at a body length of 11 m. Females 11.7 m long
are over 30 years of age and have already lost their middle pairs of

Fig. 357. Tooth of the sperm whale, Physeter catodon. A—external view;
B—longitudinal section (figure by N.N. Kondakov).

616

831

teeth. It may tentatively be assumed that at maximum age females have
attained a length of 12 m or slightly more. Males attain sexual maturity
at 5 years of age and a body length of 9.5 m but their growth tempo
subsequent to reaching sexual maturity does not decrease initially. For
the next few years the annual increment averages 80 cm, then gradually
decreases thereafter. Growth cessation is noticed at the age of 23-25
years at a body length of about 16 m (Berzin, 1961, 1964a). The body
length of male antarctic sperm whales at the time of attaining sexual
maturity is 11.5-12.5 m because of the generally larger dimensions of
whales of the Southern hemisphere (Matthews, 19386; Nishiwaki, 1955;
see under “Geographic Variation’’).

Under conditions of intense contemporary hunting practices, the
maximum length of male sperm whales very rarely reaches 18 m and
whales larger than this size are not found at all. Age determination
revealed the oldest females to be 30 years (body length 10.7 m), over
30 years (body length from 10.9 to 11.7 m), and over 35 years (11.1 and
12.4 m); males were 27 years (15.6 and 14.8 m), 29 years (16.2 m), and
over 30 years (14.8 m) or even 32 years. It has been assumed that age
determination of animals older than 35 - 40 years is not possible since the
pulp cavity is totally sealed. Be that as it may, this age is not a maximum
for the sperm whale (Berzin, 1971).

Enemies, diseases, parasites, mortality, and competitors. There is only
one known case of a piece of the rostrum of a swordfish being found in
the trunk of a beached sperm whale (Tomilin, 1957). Diseases have not
been studied. Only dental caries, distortion of the lower jaw, and some

other diseases are known. .

A significant film of diatomaceous algae with Cockoneis ceticola
predominating, has sometimes been observed on the body of the
Antarctic sperm whales. In tropical waters minute ulcers have been
detected on the skin of sperm whales, which form scars during the
residence of the animals in the cold seas: the nature and origin of these
ulcers are not understood. Crustaceans of Penella sp., common in rorqual,
infect individual sperm whales occasionally. Barnacles are also rare and
found in small numbers. They comprise three species of the genus
Conchoderma (С. auritum, С. virgatum, and С. cuvieri). Coronula sp. has
been encountered as a rare exception. The whale lice Cyamus physeteris
and C. catodontis are common parasites of most sperm whales. They
infest more often the urogenital and anal openings, less so the wrinkled
flanks, and very rarely the dorsum (Clarke, 1956; Tomilin, 1957).

Thirty-one species of helminths are known in sperm whales. These
include trematodes 1, cestodes 12, nematodes 10, and acanthocephalans
8 species. The single species of trematode, found in the Kuril waters,

832

Zalophotrema curilensis Gubanov, parasitizes the liver of only the sperm
whale.

Of the 12 species of cestodes, 5 are found exclusively in sperm whales.
Tetrabothrium curilensis Gubanov and Hexagonoporus physeteris Gubanov
parasitize the small intestine and Tetragonoporus calyptocephalus A. Skr-
jabin localizes in the bile ducts of the liver; all three have been found
in Kuril waters. Multithuctus physeteris Clarke parasitizes the bile ducts
of the liver and Polygonoporus giganticus A. Skrjabin the intestine in
the Antarctic. The cestode Tetrabothrius affinis Lonnberg, found in the
waters of Norway, Sourh Africa, New Zealand, and Antarctica, para-
sitizes the intestine of the sperm whale and three species of baleen
whales. Priapocephalus grandis Nybelin was found in the intestine of the
sperm whale and four species of baleen whales in the Pacific Ocean. Phyl-
lobothrium delphini Bosk, parasitizing the skin and found at many places
in the Atlantic Ocean, the Mediterranean Sea, waters of the Commander
Islands, Australia, and Antarctica, has been detected in the sperm whale,
six Other species of toothed whales, Weddell seal, and the Arctic whale.
Further, some cestodes not identified to species level, have been detected
in the sperm whale. These are: species of Tetraphyllidae (larvae) in the
subcutaneous adipose tissue, Diplogonoporus sp. in the bile ducts of the
liver, Trigonocotyle sp. in the small intestine of the sperm whale and
Baird’s beaked whale, and Trypanorhyncha sp. (larvae) in the stomach of
the sperm whale, sei whale, Minke whale, and Steller’s sea lion.

Of the 10 species of nematodes, 5 are found exclusively in the sperm
whale. The stomach parasite, Anisakis (Anisakis) catodontis Baylis, has
been detected in the waters of South Africa; Anisakis (Skrjabinisakis)
physeteris Baylis in the North Pacific Ocean, waters of South Africa, and
Antarctica; and Anisakis (Anisakis) ivanizkii Mosgovoy around the Com-
mander Islands. Placentonema gigatissima Gubanov was found in the pla-
centa (Kuril Islands). Anisakis (Skrjabinisakis) skrjabinit Mosgovoy раг-
asitizes the stomach and small intestine of the sperm whale and spotted
dolphins (Commander Islands, Sea of Okhotsk, and Antarctica). Anisakis
(Anisakis) dussumierit Beneden, parasite of the stomach and large intes-
tine of the sperm whale and spotted dolphins, was detected in the waters
of Japan and the Commander Islands. Anisakis (Anisakis) simplex Rudol-
phi, found in the sperm whale and widely distributed in marine mam-
mals (North Sea, eastern Kamchatka, Japan, and New Zealand) para-
sitizes the gullet, stomach, and intestine. In the northwestern part of
the Pacific Ocean, Anisakis (Anisakis) pacificus A. Skrjabin was detected
in the stomach of the sperm whale, killer whale, and fin whale. More-
over, Anisakidae g. sp. and Tetrarhynchidae m. sp. (not identified to the
species level) also parasitize the sperm whale.

618

833

Two species of acanthocephalans parasitize the intestine only of
the sperm whale: Corynosoma curilensis Gubanov was found off the
Kuril Islands and Corynosoma mirabilia A. Skrjabin in Antarctic waters.
Corynosoma strumosum Rudolphi, parasite of the intestine, is known
only in belugas, common porpoises, 11 species of pinnipeds, and land
mammals and birds. Bolbosoma physeteris Gubanov was detected in the
intestine of the sperm whale and the killer whale in the waters of the
Kuril Islands. Bolbosoma capitatum Linstow parasitizes the intestine of
the sperm whale and two more species of toothed whales from the
Atlantic Ocean and the Mediterranean Sea. Bolbosoma brevicolle Malm,
detected in the large intestine and rectum of the sperm whale, is also
known in five species of baleen whales; it has been found in the North
Atlantic, in the waters of South Africa, and near South Georgia Island.
Bolbosoma turbinella Diesing parasitizes the intestine of the sperm whale,
beaked whales, and five species of baleen whales from the Atlantic and
Pacific oceans of the Northern and Southern hemispheres. Finally, Bolbo-
soma tuberculata A. Skrjabin, detected in the South Atlantic and Indian
oceans, parasitizes the intestine of the sperm whale, sei whales, and
Bryde whales (Margolis, 1954; Delamure, 1955; Margolis and Pika, 1955;
A. Skrjabin, 1958, 1959, 1960, 1961, 1970; Berzin, 1971).

Natural mortality of the sperm whale has not been studied nor have
the reasons for the death of besched whales been ascertained.

The sperm whale has almost no trophic competitors with the possible
exception of members of the family of beaked whales.

Population dynamics. Population variations in sperm whales due to
natural factors are not known but due to hunting are significant. Hunting
of sperm whales using manual harpoons and sailboats commenced in the
first half of the eighteenth century and continued over the next one hun-
dred years on an extensive scale. By the mid-nineteenth century, sperm
whale reserves had shrunk markedly in all the seas; hunting became
unprofitable and almost ceased. This promoted a significant restoration
of the reserves of the sperm whale.

The second stage in sperm whale hunting commenced with mod-
ern techniques using whaling fleets and reached a high magnitude. As
a result, by the 1960s there was a sharp reduction in the population of
sperm whales in some regions, accompanied by a reduction in the aver-
age size of the animals caught, increase in the relative number of much
younger animals in the catch, and some other features of diminishing
reserves. This situation was mainly noticed in the waters of Japan and
the Kuril Islands and to a lesser extent in the other regions, including the
Antarctic where only adult male sperm whales that have not participated
in reproduction are known to gather.

834

Field characteristics. The huge angular head occupies over one-third
of the body length. The blowhole in the sperm whale is at the end of
the snout on the left side and not on the upper part of the head, as in
all other whales; hence the blow of this animal is not vertical but set
forward to the left at an angle of roughly 45° to horizontal. The blow is
broad, “bushy,” with a height of not more than 3-4 m. The sperm whale
can be recognized with certainty from a distance by the nature of the
blow. The body color is monochromatic, usually dark. While diving to a
depth it often exhibits the caudal flukes. After prolonged submergence,
it lies on the water surface almost without movement and allows the very
close approach of ships. (V.A.)

Economic Importance

The development of sperm whale hunting intensified mideighteenth cen-
tury following the rapid decline of the catch of right whales whose’
herds had been destroyed. Hundreds of sailboats, working mainly in
the tropical waters of all the three oceans, 1.е., predominantly in the
reproduction sites of sperm whales, were engaged in hunting them. This
hunting proved highly profitable since the demand for spermaceti was
inexhaustible and its price very high. Spermaceti was used mainly in the
candle-making industry: candles made from it burn with a bright flame
and without soot. By the mid-nineteenth century, kerosens was being
used for illumination and the demand for spermaceti fell sharply. By
this time a perceptible reduction in number of sperm whales had also
made hunting unprofitable and it almost ceased circa 1860. The let up
in hunting of the sperm whale continued for about half a century.

Early in the twentieth century, the hunting of sperm whales (mech-
anized by now) was resumed. It was particularly well developed in the
North Pacific Ocean, along the coasts of South America, and later even
in the Antarctic. Following the reduction in the population of the main
commercial species of baleen whales, the importance of the sperm whale
rose gradually in all the regions of the World Ocean although not to the
same extent everywhere.

In the North Atlantic Ocean the sperm whale is of utmost commer-
cial importance in the southern part of the range. Thus in the Azores it
represents a unique commercial species of whale. From the end of the
1930s to the end of the 1950s, 500 to 700 sperm whales were caught annu-
ally. From the beginning of the 1960s, the volume of hunting decreased
somewhat—from 400 initially to 300 whales later to 145 in 1969. Around
Madeira Island, too, the sperm whale was caught exclusively but the mag-
nitude of hunting was relatively small. Commencing from the early 1940s,

619

835

the catch did not exceed 200 a year and after 1967 did not reach even 100.
This species is of some commercial importance in the whaling industry
of Spain, Portugal, and the northern part of western Africa where sperm
whales have accounted in some years for 30 to 80-90% of the 200 to
300 whales caught annually. In the northern waters these whales are of
some importance in Norway and Iceland although the number caught is
small. On the Norwegiian coasts no more than a hundred sperm whales
are caught per year, or 15 to 40-50% of the total catch; they numbered
120-170 animals per year or 25 -45% of the total catch in the waters of
Iceland from the end of the 1950s to the mid-1960s. At the end of the

1960s, the annual catch was 100 sperm whales.

The more abundant sperm whale population of the North Pacific
Ocean ensures a larger catch. The catch is particularly high in the
coastal stations of Japan and from large floating whaling stations. Up to
1950, less than 1,000 sperm whales were caught off the coasts of Japan,
which represented about hair the total catch of whales of all species.
From 1950, hunting activity rose considerably and the number of sperm
whales caught ranged between 1,200 and 2,600 animals per year. In 1968,
the sperm whales caught here numbered 3,747 and in 1969—3,668. In
some years sperm whales accounted for 60-70% or even more of the
total catch. Until the mid-1950s, when the “pelagic” fleet was not large,
the catch of sperm whales usually did not exceed 500-700 animals per
annum. But from the mid-1950s, with the introduction of new Soviet and
Japanese whaling fleets, it began to rise rapidly: initially up to 2,000 and
then 4,000, reaching almost 8,000 sperm whales a year by the early 1960s.
Between 1965 and 1969, the number of sperm whales caught totaled
10,500 to 12,500 per year or 50-70% of the total catch of the fleet.
Sperm whales caught from the coastal bases on the Kuril Islands between
1950 and 1962 totaled 1,400 to 2,000 (70-80% of the total catch). The
whaling industry of the Kuril Islands came to a halt in 1964. Large-scale
hunting of sperm whales in the northeastern part of the Pacific Ocean
also ceased. From the coastal bases of British Columbia, from the end or
the 1940s to the early 1960s, 120 to 320 sperm whales were caught per
year but whale hunting ceased in 1967. Off the coasts of California, the
catch never exceeded a few tens (sometimes up to a hundred) of sperm
whales per year.

The sperm whale catch in the Southern hemisphere has been large
with several areas participating in the hunt: waters of Chile, Peru, South
Africa, pelagic hunting in the Antarctic, and, in recent years, in the more
northern waters. In the African waters (Natal and Cape Province) whale
hunting was resumed in 1948 and 400 to 1,000 sperm whales of a total
of 1,200 to 2,700 whales were caught every year until 1956. From 1957,

620

836

the catch of sperm whales increased considerably and 1,200, 2,500, and
even 3,600 sperm whales were caught every year (50-70% of the total
catch). Off the coasts of Chile and Peru, sperm whales represented the
main species of whales hunted and accounted for 70.9 and even 100% of
the catch. The maximum number of these whales caught from the mid-
1950s to the mid-1960s ranged from 2,500 to 5,000 animals per year.
At the end of the 1960s, hunting fell slightly to less than 2,000 animals.
After the Second World War, the “pelagic” whaling fleet of the Antarctic
developed rapidly and caught a large number of sperm whales (excep-
tionally large males). But, of the 2,000-4,000 to 6,000-7,000 animals
caught, sperm whales in a season constituted only 10-15% and later
20-22% of the total catch of the whaling fleet. From 1967 to 1969, the
catch of sperm whales in the Antarctic fell to 2,500 to 2,600 animals per
year but the Antarctic whaling fleet began catching these whales north of
40° S lat. along the course into Antarctica and back. This catch accounted
for 2,000 or more sperm whales in a season. In Australia hunting of the
sperm whale commenced only in the mid-1950s, following a reduction in
the population and subsequent total ban of hunting of humpback whales
(these were the lone target of hunting in these waters). From the mid-
1960s, the number of sperm whales caught in Australian waters totaled
500 to 600 per year.

Modern hunting of sperm whales, like that of other large whales, is
carried out in special steel ships equipped with harpoon guns. The whale
is killed with a harpoon with a grenade screwed to its tip and primed
with gun powder. The grenade fires 4 sec after shooting and by this time

the harpoon has penetrated the whale body (Fig. 358). The firing can be

fatal or otherwise depending on the resistance of that part of the trunk
struck by the harpoon. The wounded whale is dragged toward the boat
by a rope tied to the harpoon, using a special winch, and the animal
finally killed by shooting a second (or third) time. Instances of a whale
requiring up to six shots are known.

The behavioral aspects of the sperm whale make for some typi-
cal features in hunting them. After prolonged submergence, the sperm
whale usually remains a long time on the sea surface. This facilitates the
approach of the ship within shooting range while the animal’s attempt
to escapeton a straight course enables the chasing whale boat to catch
up with it quickly. Thus most of the sperm whales sighted from the ship
can be killed. But if the marked sperm whale succeeds in diving before
the ship can approach, it is futile to wait for it to surface again.

Air is pumped into the body cavity of the killed sperm whale so that
the carcass floats and can be towed to the floating base or the coastal
station for dressing. First the blubber is removed in layers and then the

620

621

837

Fig. 358. Harpooned sperm whale (photograph by V.A. Arsen’ev).

head is severed from the trunk. Using a steam or electric hoist, the large
stock of fat in the spermaceti sac is recovered, sliced into large lumps,
then charged into a boiler for melting the spermaceti. Then the viscera
are removed from the carcass and the flesh from the bones. The bones
of the skeleton and skull are cut by steam-powered saws into bits that
can be fed into fat-melting boilers to ensure more complete conversion
into oil.

The main product obtained from the sperm whale (Table 62) is
the oil from the blubber, bones, and viscera. The physical and -chemi-
cal properties of the oil of the sperm whale differ from those of baleen
whales (low specific gravity, low saponification number, low iodine num-
ber, etc.). The oil is used for commerical purposes by various industries.
A large male (length 14-16 m), on average, yields about 10 tons of fat.
Spermaceti has a characteristic chemical composition and is collected
and processed separately from the blubber. Spermaceti is used by per-
fumeries in preparing cosmetic media (creams and pomades) and to a
lesser extent in other industries as well. The liver of the sperm whale
contains the maximum amount of vitamin A compared to the liver of
all other species of whales: about 6,000 units, on average, versus 1,500
units in a female fin whale (Mrochkov, 1953). The mechanical method
of pressing the subcutaneous fat layer yields more oil than digesting in a
boiler (Dormenko, 1952; Zaikin, 1953). In the former method the upper

838

layers of the integument can be used in making leather goods. For this
purpose two layers of 5-6 mm thickness each are taken from the carcass
of the whale and a third additional layer up to 8 mm thick from its head.
Much deeper layers can also be used in the leather industry but only
after special compaction using special fillers.

Table 62. Weight of the body parts of the sperm whale, kg (Zenkovich, 1937b;

M.V. Ivashin)
Body part Far East (Kronotsk Bay) Antarctica
Male 13.5 m Male 18.0 m Male 16.1 m
Subcutaneous fat 4,955 12,663 11,450
Spermaceti 620 5,757 4,400
Spermaceti sac 3,981 10,343 8,665
Flesh 5,640 7,697 7,850
Skull 1,800 6,087 4,100
Lower jaw 320 880 1,050!
Vertebral column 1,986 3,387 3,4552
Caudal flukes 314 DS
Ribs 965 2,147 3,2903
Flippers 485 978
Pelvic bones 108 200
Sternum 209 329
Viscera 1,282 2,199 3,340
Total 22,665 53,365 47,600
Liver — — 620
Liver and kidney 331 368 ==
Heart 113 211 —
Lungs 161 317 —
Lungs, heart, neck, and gullet — — 1,300
Stomach and intestine — — 700
Intestine 254 465 —
Testes 10.5 16.5 —
Penis 88 106
Remaining parts 323 716 720

1Lower jaw together with tongue.
Vertebral column together with caudal flukes.
3Ribs together with flippers.

Length of whale, m Area of raw skin, т?

8.9 19.2
9.5 25.5

10.5 28.4

11.0 33.0

14.5 55.5

15.3 71.0

622

839

Soles made from the skin of the sperm whale are not inferior (some-
times even superior) in wearing properties to those made from cattle
hide. However, the skin of the sperm whale always suffers from defects,
such as flaws of 5-10 mm in diameter and scars and scabs sometimes up
to 50 cm? or more in area. As these damages are randomly scattered all
over the skin, cutting is difficult and much wastage results (Bulgakov et
al. 1954). Hence the use of sperm whale skin in making leather goods has
not developed. The lower layers of the skin, after mechanical degreasing,
can be used in making gelatin and glue (Kolchev, 1954). The teeth of
the sperm whale are excellent material for carved articles (chess pieces,
handles, etc.) and almost indistinguishable from those made of ivory.

Sperm whales carry ambergris in the form of a hard waxlike substance
present in large lumps of indeterminate form. Crude ambergris varies
from gray to black and smells like wet soil. It has a granular and layered
structure. Its specific gravity varies from 0.730 to 0.780; it softens from
the warmth of the hand, melts at 60°C without bubbling, and volatilizes
at 100°C. It dissolves well in hot alcohol and burns with a light blue
flame and slight resinous smell.

Although ambergris is known from ancient times, its origin has yet
to be ascertained. It is sometimes found floating on the water or cast on
seashores. In the remote past it was regarded either as refuse of birds
or as an exudation of plants and roots of trees. Later, the remains of
chitioous “beaks” of cephalopods consumed by the sperm whale were
detected in the ambergris, which suggested its origin from the intestinal
tract of the whale. Its formation is thought to be a reaction to infection
by parasites inhabiting the intestine or a product of the normal secre-
tion of the rectal glands. The third point of view would appear to be
more correct, according to which ambergris is formed as a result of the
retention of undigested remnants of “beaks” of cephalopods, which are
enveloped in special secretions in the digestive tract of the sperm whale.

Depending on its quality, ambergris costs 150-200 rubles per kg. The
production of synthetic ambergris has somewhat affected the cost of the
natural product but the quality of the synthetic is inferior to the natural.
In olden times ambergris was used as an antispasmodic and in Chinese
medicine as a stimulant or antiseptic. The chief value of ambergris lies in
its exclusive property of absorbing odor and exceptional retention of it.
Therefore, natural ambergris is used in producing perfumes of superior
quality.

Ambergris is found in the intestinal tract of sperm whales in lumps
weighing several hundreds of grams to tens of kilograms. Remarkably
large lumps of ambergris have also been found: one lump weighing 392 kg
was recovered in the Azores and another weighing 420 kg off the coast of

623

840

Australia. In our country a lump of ambergris weighing 351 kg was recov-
ered in 1958. In subsequent years, ambergris production has exceeded 3
tons (Ivashin, 1966). Ambergris as a raw material requires sometime for
maturation and hence all varieties of ambergris do not command the
same price and some are even totally unsuitable for use in perfumeries.

Following the conclusion of the International Whaling Convention,
1946, some restrictions were imposed on the hunting of sperm whales.
According to the international hunting rules, killing of sperm whales less
than 10.7 m long was prohibited from the coastal whaling stations and
less than 11.6 m long from the floating fleets, even though many sperm
whales of these sizes are mature animals. In spite of these restrictions,
highly intense sperm whale hunting continued and enormous catches
led to a steady depletion in numbers, especially in the better exploited
populations. Now a restriction has been imposed on killing sperm whales
by prescribing annual quotas in all parts of the World Ocean, taking
into consideration the strength of individual populations. In most cases,
it would be extremely useful to restrict the killing exclusively to male
sperm whales which, given the prevailing polygamous mode of life of
these whales, could not adversely affect the normal replenishment of the
various populations. (V.A.)

Subfamily of Dwarf Sperm Whales
Subfamily KOGIINAE Gill, 1871

Genus of Dwarf Sperm Whales
Genus Kogia Gray, 1846

1846. Kogia. Gray. Zoology of the Voyage of H.M.S. Erebus and Terror,
I, p. 22. Physeter breviceps Blainville. :
1876. Cogia. Wallace. Geogr. Distr. Anim., 2, p. 208. Correction of Kogia
Gray. Nom ргаеосс. (V.H.)
Dimensions the smallest in the family, with a total length up to 3.4 m.
The body form is dolphin-shaped. The head is small, proportional,
constitutes 1/8th to 1/6th of the body length, and is rounded anteriorly.
The spermaceti зас” is well developed on the head but much smaller than
in the sperm whale. The blowhole is situated on the parietal section of
the head, almost at its center. The blowhole opening is crescent-shaped.
The end of the lower jaw falls far short of the anterior end of the head.
The fairly high dorsal fin lies roughly midbody. The flippers are relatively

5 See description under sperm whale, р. 799.

625

841

narrow and short. The body is dark-colored dorsally and light-colored
ventrally.

The rostrum is broad at the base, constricted sharply in the anterior
part, and is shorter than the cranium. The depression on its dorsal sur-
face is small and has a longitudinal crest formed by the premaxillae and
maxillae. The nasal bones are not fused with the premaxillae. The max-
illae have very large preorbital processes. The zygomatic bone does not
articulate with the squamosal. The symphysis of the lower jaw is shorter
than half its length.

Teeth: 8-16 pairs on the lower jaw and 1-3 pairs in the anterior
section of the upper jaw (but sometimes totally absent in the latter). They
are thin, curved, and the sharp tip set posteriorly. Enamel is absent.

Vertebrae 52-57: cervical 7, thoracic 12-14, lumbar 9-12, and cau-
dal 21-27. The cervical vertebrae are fused. The flattened scapula has
a small acromion and large coracoid processes. Ribs 13-14 pairs. The
sternum is short. The phalangeal formula in the forelimbs is subject to
individual variation: I,, Пб-з, II,-3, [У4 -з, and V>-7.

The population of the dwarf sperm whale is small. Biology has not
been well studied. It lives singly or in small groups and apparently feeds
mainly on cephalopods. The periods of mating and parturition are pro-
tracted. Gestation continues for about nine months. Females evidently
reproduce annually. They live in the warm waters of the Indian, Atlantic,
and Pacific oceans (Fig. 359).

There is no special hunting of dwarf sperm whales in view of their
rarity.

The genus comprises two species (Handley, 1966): 1) dwarf sperm
whale, Kogia breviceps Blainville, 1838; and 2) Owen’s dwarf sperm whale,
Kogia simus Owen, 1866.

Dwarf sperm whales are not observed in USSR waters but summer
transgressions are possible (especially of Owen’s dwarf sperm whale) into
the Sea of Japan and toward the southern part of the Kuril Islands. (V.S.)

DWARF SPERM WHALE
Kogia breviceps Blainville, 1838

1838. Physeter breviceps. Blainville. Ann. Franc. étrag. d’Anatomie
Physiol., 2, p. 337, pl. 10. Cape of Good Hope. (V.H.)

Diagnosis

Dimensions largest in the genus, with the overall body length varying
from 2.7 to 3.4 m. The relatively low dorsal fin lies slightly posterior to

“(лэчэзту УЛ) 21801 ‘зэеЧм шлэ4$ улемр Jo зпиэ8 ay) Jo э8иез ‘66$ “314

051 06

тот ost Е ,

з A! ae
= ON =

see =
= `. а т
>... O}
Е BST.
в == хх =
г. z ee, =>
3 7 у =

79

625

843

midbody. Teeth are absent in the upper jaw. Mandibular teeth 12-16
pairs (less frequently 10-11 pairs). (V.S.)

Description

The body dorsally, including the caudal flukes and flippers is dark, almost

black. The flanks are light gray and the abdomen light-colored. Poste-
rior to the eyes and anterior to the flippers, one or two light-colored
projections run bottom upward into the dark-colored region (Fig. 360).
Light-colored spots are visible on the ventral surface of the caudal flukes.

The body dimensions of adult male and female dwarf sperm whales
from New Jersey and Texas states (USA) (Tomilin, 1957) are respectively
(in cm): body length 299 and 274; distance from tip of snout to center
of blowhole 32 and 35, up to anterior margin of dorsal fin 154 and 150;
distance from anal opening to notch between caudal flukes 97; length
of flippers 41 and 43; height of dorsal fin 9 and 13; length of dorsal fin
14 and 41; and width of caudal flukes (between apices) 50. The animals
range in weight from 318 to 408 kg.

The condylobasal length of the skull (Fig. 361) is 391-469 mm.
The mandibular symphysis is long (8.6- 12 cm), with a keel on the ven-
tral surface. The alae of the pterygoids and basioccipitals are elongated.
The foramen magnum is situated roughly midpoint of the skull height.
(V.S.)

Geographic Distribution and Biology

Inhabits predominantly the warm belt of the Pacific Atlantic, and Indian
oceans. Distribution has been studied mainly from the finds of animals
cast on coasts or shoals. More than 15 finds of beached dwarf sperm
whales are known on the coasts of South Africa, Australia, and New
Zealand, and over 50 on the Atlantic coast of the USA. Finds in the
Pacific Ocean are not rare near Japan, the East China and South China
seas, Gulf of Siam, along the coast of North America (Washington and

Fig. 360. Dwarf sperm whale, Kogia breviceps (figure by N.N. Kondakov).

625

844

Fig. 361. Skull of the dwarf sperm whale, Кода breviceps (figure by М.М. Kondakov).

626 California states), and along the coasts of Mexico, Peru, Chile, Fiji

Islands, Australia, Tasmania, and New Zealand. In the Indian Ocean,
it is distributed from southern Australia to South Africa. In the Atlantic
Ocean, beached animals are known on the coasts of North America from
Nova Scotia to Florida and Texas, and on the coasts of South Africa,
France, and Holland. In the Atlantic Ocean, these whales probably move
into the open sea even in latitudes more northern than Holland (Dell,
1960; Tomilin, 1962; Gaskin, 1966; Hershkovitz, 1966; Handley, 1966).
The residence of these animals in the waters of the Soviet Union has
not been established. Transgressions are possible in the summer months
into the Sea of Japan and the southern part of the Kuril range.
Geographic variation of the species has not been studied.
Information on the biology of dwarf sperm whales is very
fragmentary. Remains of cephalopods (Sepia officinalis and others) have
been detected in their stomach. Cephalopods apparently serve as their
main food. Bones and otoliths of fish (Тисйо4оп sp. and others),

845

crabs (Carcinides maenas), shrimps (Paciphola pacifica, Pandalus sp.,
Pandalopsis sp., Penaeus sp., Hymenodora sp.) have been found less often.

The period of mating is greatly prolonged and newborns have been
encountered even in November and February; however, the majority of
these whales probably mate in summer. Gestation is thought to extend
for 9-10 months. The length of a newborn calf is around 110 cm; whales
153-171 cm long were still suckling and a female with a body length of
188 cm was immature. Females attain sexual maturity at a body length
of 218-219 cm. A female about 3 m long, beached on a Dutch coast in
December, 1925, contained an embryo 20 cm long. Females give birth to
a single calf. An instance is known of a female, accompanied by a small
calf, who was simultaneously pregnant and lactating.

Dwarf sperm whales live singly, in pairs, or in small groups of up to
five animals. Their movements are slow.

Penela sp. was found among the ectoparasites. Larvae of the cestodes
Monorygma grimaldii Monier and Phyllobothrium delphini Bosk and the
nematodes Crassicauda magna Jonston and Mawson and Pseudoterra-
nova mogiae Jonston and Mawson, have been detected in the abdominal
cavity and subcutaneous tissue.

A female 277 cm long weighed 369 kg while large males НЫ up
to 500 kg. There is no special hunting for these whales. The Japanese
industry while catching various species of dolphins occasionally nets
dwarf sperm whales also. Although their flesh is edible, these whales
are not of economic importance (Delamure, 1955; Tomilin, 1957, 1962;
Nishiwaki, 1965; Handley, 1966). (V.A.)

OWEN’S DWARF SPERM WHALE
Kogia simus Owen, 1866

1866. Physeter (Euphyseter) simus. Owen. Trans. Zool. Soc. London, 6
(Г), р. 30, pls. 10-14. (V.A.)

Diagnosis

Dimensions the smallest in the genus, with overall length ranging from
2.1 to 2.7 m. The relatively high dorsal fin is roughly midbody. The upper
jaw usually has 1-3 pairs of teeth and the lower 8-11 pairs (less often,
13 pairs). (У.5.)

Description

The body is dorsally dark gray and ventrally white.

627

846

The body measurements of a male and two females from Japan
(Yamada, 1954) are: overall length 2.22 m; distance from tip of snout to
blowhole 16.3, 20, and 20 cm, from tip of snout to apex of flippers 78.5,
87, and 86 cm; length of base of dorsal fin 34, 33, and 42 cm; height of
dorsal fin 13, 11, and 17.5 cm; and distance from caudal notch to anal
opening 68.5, 82, and 82 cm.

The weight of the animals varies from 136 to 272 kg.

The condylobasal length of the skull is 262-302 mm. The mandibular
symphysis is short, 37-46 mm, and smooth on the ventral surface. The
alae of the pterygoids and basioccipitals are short. The foramen magnum
lies considerably below midskull height.

The skull measurements of a male and two females from Japan
(Yamada, 1954) are (in mm): condylobasal length 271, 284, and 297;
length of rostrum 140, 165, and 160; width of rostrum at base 128, 136,
and 140; interorbital width 218, 240, and 259; and length of lower jaw
224, 253, and 255. (V.S.)

This species is encountered in the coastal waters of South Africa,
southern Australia, India, Sri Lanka, Hawaiian Islands, Japan, and off
the east coast of the USA. It is known predominantly from beached
animals. The maximum number of beached animals has been found on
the coasts of Japan and on the eastern coasts of the USA (Handley,
1966).

Geographic variation has not been established.

Biology has not been studied. Differences between the dwarf sperm
whale and Owen’s dwarf sperm whale are not widely recognized and
hence some data on the biology ascribed to Kogia breviceps may actually
pertain to Kogia simus. (V.A.)

Family of Beaked Whales
Family ZIPHIIDAE Gray, 1865°

Body dimensions medium and large. Males are larger than females or
vice versa.

The body is more or less spindle-shaped. Anteriorly, the snout forms
a narrow “beak” sharply demarcated from the frontal adipose body
among bottlenose and beaked whales or gradually rising above the head
in the rest of the members of the family. The blowhole is crescent-shaped
and its bulge may face the head or the tail. The small dorsal fin is situated
at the level of the anal opening or slightly anterior to it. There is no notch

6 Hyperoodontidae, according to some authors.

629

847

between the caudal flukes ог it is very small. The flippers are low. The
body is usually monochromatic brown or gray but somewhat lighter on
the ventral side.

The rostrum is long and narrow, longer than the cranium, with an
open mesorostral groove (closed in old animals). The maxillae, premax-
illae, and frontal bones bear crests. The palatine bones form part of the
anterior wall of the nasal passage. The petrosal bones are fused with
the skull. The lachrymal bones are large and not fused with the zygo-
matic bone. The pterygoids are large and separated by some distance.
The lower jaw is wider than the upper and projects forward slightly.

Numerous teeth are seen in the embryos, which reduce in number
subsequently. The adults of most of these species have just a few teeth
only in the lower jaw. The number of functional teeth varies from г in
Tasmacetus to § in Berardius and $ in the rest of the genera.

The number of vertebrae does not exceed 50 and from 2-7 cervical
vertebrae are fused. Ribs do not exceed 10 pairs and their sternal sections
are not ossified.

The system of air sacs is simple, without preorbital and postorbital
lobes. In adult animals, rudiments of the olfactory nerves are preserved.
The stomach comprises 4-14 chambers. The members of this family feed
mainly on cephalopods (teuthophagous) but also consume fish. They
are capable of diving deep and remaining submerged for a long time.
Some perform annual migrations. They live singly or in herds, usually
comprising 10-15 animals.

These animals are distributed in the warm, temperate, and cold
waters of the World Ocean (Fig. 362).

The oldest of the Ziphiidae is evidently Notocetus Moreno from the
Lower Miocene of Patagonia, resembling in many features the Miocene
Squalodon Grateloup (family Squalodontidae Brandt) and having, in par-
ticular, numerous teeth. Reduction of teeth among the primitive Ziphi-
idae occurred in the Miocene and typical members of contemporary
beaked whales appeared at the end of this period.

The family consists of 17 genera, of which 5 are extant: Tas-
macetus Oliver (Tasmanian beaked whales), Berardius Duvernoy
(Pacific beaked whales), Mesoplodon Gervais (“sword-tooth dolphins’),
Ziphius G. Cuvier (Cuvier’s beaked whales), and Hyperoodon Lacépéde
(bottlenose whales).

The generic division of the family has given rise to no doubts among
researchers. However, the systematics of the species of beaked whales
requires further development (especially of genus Mesoplodon). About
15 species are usually recognized in the family.

Beaked whales are of little economic importance.

|
ues
ЩИ
ИЗ
‘НИЦ

АИ
|
I

x |
ИН и

at pa
ты ©

"| is Е
| MTA A ий
|} | | | [ИН Ш | in) AAT |
UPL La
MUAH) ПИ Mil iil!

9.
«ар. 1
| НИНА О
ИИ

(4
yi Ш

ЩИ

ИИЦ

1000 0 1000 2000 3000 4000 5000 Кт
и Н |

Fig. 362. Range of the family of beaked whales, Ziphiidae (V.A. Arsen’ev).

628

849

All of the genera, except Tasmacetus, are encountered or could be
encountered in USSR waters. Our hunting of these animals is in small
numbers. (V.S.)

Genus of Pacific Beaked Whales

Genus Berardius Duvernoy, 1851

1851. Berardius. Duvernoy. Ann. Sc. Nat., Zoologie, 15, p. 52. Berardius
атоихи Duvernoy.
Dimensions large, the largest in the family.

The frontal projection is quite high. The “beak” is highly elongated
and somewhat flattened. The low dorsal fin is situated above the anal
opening and is not flexed along the posterior margin [falcate]. The flippers
are relatively short. The lower jaw is slightly longer than the upper one.

The color is dark brown, the under varying from grey to whitish.

Asymmetry of the skull bones is less manifest than among the other
members of the family Ziphiidae. The rostrum is narrow, its length more
than double that of the cranium, and it is slightly flattened dorsoventrally.
The maxillary crests are low. The premaxillae are symmetrical and their
width almost equal throughout their length. The suture between the
_ ргетахШае and maxillae does not extend beyond 20% of its length on the
dorsolateral surface of the rostrum. The frontal bones have broad orbital
processes. The large, massive, rounded, and almost identically sized nasal |
bones project markedly into the frontals and occupy the highest position
on the skull. The mesethmoid bone is only partly ossified.

The anterior section of the lower jaw has two pairs of teeth, of
which the anterior larger ones are located at the tip of the lower jaw
and remain exposed even when the mouth is closed. The posterior pair
of teeth emerge 10-20 cm behind the anterior ones. A few more teeth
may remain concealed in the gums.

Cervical vertebrae 7, thoracic 10-11, lumbar 12, and caudal 17-19;
total 46 - 49. The three anterior cervical vertebrae are fused. The sternum
consists of five sections.

Biology has not been well studied. These animals remain confined
in groups of roughly 20. They feed mainly on cephalopods. Mating and
parturition are protracted. Gestation continues for 10 months.

These animals are encountered in the Pacific and Indian oceans and
in the South Atlantic.

The genus consists of two species: Arnoux’s beaked whale (southern
beaked whale), Berardius arnouxi Duvernoy, 1851, and Baird’s beaked
whale (northern beaked whale), Berardius bairdi Stejneger, 1883.

630

850

Only Baird’s beaked whale is encountered in USSR waters.’
There is no special hunting for Baird’s beaked whale; it is caught
only incidentally. (У.5.)

BAIRD’S BEAKED WHALE®

Berardius bairdii Stejneger, 1883

1883. Berardius bairdi. Stejneger. Proc. Ц. 5. Nat. Mus., 6, р. 77. Staraya
Gavan’, east coast of Bering Island.

1883. Berardius vegae. Malm. Bihang Svenska Vet. Acad. Handl. 8 (4),
p. 109. Commander Islands.

1947. Rostrifer nestorésmirnovi. Zenkovitsh (Zenkovich). Zh. “Rybnoe
Khozyaistvo,” No. 10, p. 15. Nomen nudum.

1955. Berardius vegana. Bourdelle and Grassé. Traité de Zool., 17, р. 429.
Erroneously for vegae Malm. (V.H.)

Diagnosis
Only species of the genus found in waters of the USSR.

Description

|
The head is relatively small (Fig. 363). The “beak” is flattened dorsoven-

trally. The “forehead” of males rises steeply upward and is set off from
the “beak” by a distinct transverse groove (Fig. 364). In females the fore-
head rises more gently and the transverse groove is absent. The blowhole
is crescent-shaped and its bulge faces forward. Two (less often, three or
four) deep longitudinal grooves occur on the throat (distinctly visible
even in embryos; Fig. 364). The shape of the dorsal fin is variable—from
low humplike to relatively high and well-proportioned. It is two or three
times longer than its height. The flippers are placed close to the ventral
surface of the body. They are quite short and broad (length only 2.5 -3.2
times their width). The caudal flukes have a small notch.

The body color turns dark with age. In young animals the upper
portion is gray with a brown tinge and the abdomen and flanks somewhat

7 Вана’; beaked whale, caught by our whalers in the Far East, was long erroneously
classified as a bottlenose whale (Hyperoodon). Only after 1950 (Tomilin, 1952, 1957) was
it demonstrated that these animals were actually Baird’s beaked whales. Thus all the infor-
mation about bottlenose whales in the Pacific Ocean reported until 1956 pertains to this
species.

8 Sometimes, especially in the earlier literature, and also in foreign publications (Her-
shkowitz, 1966), also referred to as the “Pacific beaked whale”. (V.H.)

630

631

631

851

777.7

ae

72
LZ РРР 2227 Е

3 22.25...
ИИ АР
LLY SEE 7777 —_
Я LP =
LE LZ РРР ЕЕ 2
И: РРР
= и Be 5

CALLE

iy ico
и р

Fig. 363. Baird’s beaked whale, Berardius Бата (figure by М.М. Kondakov).

Fig. 364. Head of Baird’s beaked whale, Berardius bairdi, front view. Pacific Ocean
(photograph by M.M. Sleptsov).

lighter. In adult animals the dorsum and flanks are dark brown and the
abdomen slightly liger in color (Sometimes may be white). White spots
are seen on the skin surface of adult whales in the navel region between
the flippers and under the throat. It is possible that these spots reflect an
age-related variation of skin pigmentation (Tomilin, 1957). White scars
(probably caused by bites of males) are more numerous on the dorsum
than on the abdomen. Both the pairs of teeth occur in the region of
the mandibular symphysis. The anterior teeth are set on the very tip
of the jaw and may rise to a height of 79-89 mm and the posterior
ones to 53-67 mm (Omura et al., 1955; Tomilin, 1957). Rear teeth are
usually absent in females. In the newborn and suckling calves the teeth
are not erupted. In young animals the anterior teeth are conical and
may be longer than in adults. With advancing age, the teeth wear down
somewhat and become more massive and flattened laterally, especially
that part in the alveolus. Barnacles of the genus Conchoderma sometimes
colonize on the anterior teeth.

852

631 Fig. 365. Head of Baird’s beaked whale, Berardius Бат, ventral view. Pacific
Ocean (photograph by M.M. Sleptsov).

632 Fig. 366. Baird’s beaked whale on a deck. Pacific Ocean (photograph by М.М. Sleptsov).

The main measurements of four adult females (Tomilin, 1957) are
(in cm): body length 1,225, 1,080, 1,095, and 1,090; distance from tip of
upper jaw to blowhole 132, 120, —, and —; distance from anal opening

853

to posterior edge of caudal flukes 355, —, 323, and 324; length of flippers
152, 108, 118, and 125; height of dorsal fin 30, 30, 31, and 33; and width
of caudal flukes (between apices of lobes) 310, —, 270, and 268. The
dimensions of males are less than those of females. The maximum known
length of a female is 12.2 m, of a male 11.9 m.
The measurements of the skull (Fig. 367) of two adult females and
a young male Baird’s beaked whale (Omura et al., 1955 and Tomilin,
1957) are respectively (in cm): condylobasal length 152, 142, and 106
(tip of rostrum broken in the male); zygomatic width 75 (measurement
not taken in the second female and male); length of rostrum 96, 92,
and 52; width of rostrum at base 47, 43, and 31; length of lower jaw
133, 130, and 90; and length of mandibular symphysis 29, 27, and 17.

(V.S.)

AS

ey ed

SSS ==
SSS

=>
SSSSSS

SSSR

SSeS SSS
~

7
ИИА
"И
1 AAW NY NYS

BAW LY SRS
ГИЯ! Ss
ЧИ!

0;

632 Fig. 367. Skull of Вана’; beaked whale, Berardius Бата! (figure by М.М. Kondakov).

632

633

854

Geographic Distribution?
These animals are distributed in the North Pacific Ocean.

Geographic Range in the USSR (Fig. 368)

In the Sea of Japan, it is distributed in Peter the Great Gulf and along the
southern coasts of Sakhalin; in the Sea of Okhotsk, along the islands of
the Kuril range and the coasts of western Kamchatka, close to the eastern
coasts of Sakhalin, in the Gulf of Sakhalin in waters of the Shantarsk
Islands and [ona Island, in the central part of the Sea of Okhotsk; in
the Pacific Ocean, in waters of the Kuril Islands, along the southeastern
coast of Kamchatka, in Avachinsk Bay and Kronotskiy Gulf, and around
the Commander Islands; and in the Bering Sea, around Karagin Island
and in Olyutorskiy Gulf. It apparently does not penetrate farther north
than Cape Navarin (about 62°N lat.).

Geographic Range outside the USSR (Fig. 369)

The southern boundary of distribution probably traverses at the latitude
of southern Japan. It is known in the Sea of Japan as well as the Pacific
Ocean side of the islands of Japan but is more common and abundant
in the Pacific Ocean than in the Sea of Japan. On the eastern side of
the North Pacific Ocean, it lives from the coasts of California along the
entire North American continent to the coasts of the Alaskan peninsula
and Aleutian Islands. In the Bering Sea, it is distributed from Bristol
Bay and the Pribilof Islands along the boundary of the eastern Bering
Sea shallow-water zone to northwest of Cape Navarin on the Asian coast
and the southern part of the Gulf of Anadyr (Zenkovich, 1939; Sleptsov,
1955; Tomilin, 1957; Chapskii, 1963). (V.A.)

Geographic Variation

Geographic variation has not been established. It is possible that Baird’s
and Arnoux’s beaked whales (bairdi and arnouxi) represent only subspe-
cific forms of the same species (Hershkowitz, 1966). Further, there is
justification to assume that the species forms only some local herds or
populations and not subspecies. For example, the Pacific Ocean coasts of
Honshu and the Sea of Okhotsk’s coast of Hokkaido are evidently inhab-
ited by various populations of Pacific beaked whales (Omura, Fujino, and
Kimura, 1955; see below). (V.A.)

9 [See footnotes on page 854][sic; pp. 843 -844—Gen. Ed.]

635

855

Biology

Population. Only a few hundred beaked whales are caught every year
in the North Pacific Ocean. It may be concluded from this that the
population of these whales is comparatively small. However, in most
parts of the range the beaked whale is caught only incidentally while
hunting for other whales. This no doubt has a bearing on their overall
catch.

Food. Data on the food of beaked whales are very meagre. Evidently
cephalopods and to a lesser extent fish represent the main food.

Food Objects of Baird’s Beaked Whale
(Betesheva, 1960, 1961)

Cephalopods Fish

Gonatus magister Alaska pollock, Theragra

Gonatus fabricii chalcogramma

Gonatus borealis Podonema, Podonema longipes

Ommatostrephes sloani- Navaga, Eleginus navaga

pacifius gracilis
Albatross rat-tail, Coryphaenoides
pectoralis

Black rat-tail, Hemimacrurus
acrolepis = [Coryphaenoides
acrolepis|

There are references to finds of redfish (ocean perch), skates and
their eggs in the stomach of beaked whales (Zenkovich, 1939; Tomilin,
1957).

Daily activity and behavior. Pacific beaked whales are usually confined
to groups of 5-7 each or sometimes up to 20 animals (Zenkovich, 1939).
They dive for 8-20 min (less frequently up to one hour) after which
they “rest” on the sea surface for 3-4 min producing 10-15 blows in
this period. The blow of the beaked whale is low, 1.0-1.5 m in height
(Tomilin, 1957). On surfacing after prolonged submergence, the forehead
of the whale is seen first on the surface followed immediately by a small
blow. The anterior part of the dorsum and the dorsal fin are seen next.
During intermediate submergences, only the upper end of the dorsal fin is
exposed since the whale floats right on the water surface almost without
flexing the body. Before undertaking a fresh prolonged submergence, at
the moment of appearance of the dorsal fin, the body flexes and the
head is concealed under water. Next the anterior part of the trunk is
gradually flexed more sharply and assumes an almost vertical posture.
The dorsal fin and the caudal stem adjoining it then gradually rise above
the water and, when they are highest above the surface, the entire body

856

25
140 150

те J | затык
у
2
160 170

633 Fig. 368. Range of Вана’; beaked whale, Berardius bairdi, in the USSR (V.A. Arsen’ev).

begins to descend slowly and that part of the trunk visible above the
water gradually submerges into the sea, the last to disappear being the
caudal flukes. However, the caudal flukes, unlike in sperm whales, do
not rise out of the water.

On surfacing, the beaked whales form a typical chain, sometimes
in twos or even threes, one behind the other (Fig. 370). The animals
leap from the water at an angle to the sea surface roughly equal to one-
third the length of the trunk, then fall on the water with a loud thud
(Tomilin, 1957). Sometimes they completely breach the water and dive
with the head down like a dolphin without landing on the flanks or on
the abdomen, as generally happens in the case of large whales.

The herds usually differ in composition. Adult males and females
with calves remain singly, far away from the coasts, forming independent
herds, while young animals gather in separate groups close to the coasts
and transgress even into bays and gulfs (Zenkovich, 1939). Mixed herds
are also noticed, with males, females, and young animals living together

857

180

|
|
|
! в
| Е:
И, т к
: Е ИНН И
ТО Wace
Kiss eit HL! Hi, !
: AAS
@ a \ j “eit Ри! | |] м i
р сх Ц ИЯ.

Fig. 369. Species range of Ваша’ beaked whale, Berardius bairdi (V.A. Arsen’ev).

634

858

2 Ри... -/ р PN ane a
И LZ 4 ZI AE BE eee
= 22 AZ А, att. Le, ee ie SOO DES
LA GI Е LZ РРР =
Digs ЕЕ Ct a ee eZ
oe ene za LEAF AEG gee ae’. РОЗИ РР
р р

АС

635 Fig. 370. Ваша’; beaked whale, Berardius bairdi, at sea (figure by М.М. Kondakov).

(Tomilin, 1962). Many cases of beached Baird’s beaked whales have been
recorded: near Centerville and Santa Cruz on the Californian coast, in
Tokyo Bay in Japan, and on the Pribilof and Commander Islands.

Seasonal migrations and transgressions. Beaked whales perform reg-
ular seasonal migrations but their periods and courses have not been
established. It is known that they appear in spring in the northern part
of the range, in the Sea of Okhotsk and the Bering Sea, and leave for
the south in autumn for the waters of California and southern Japan

636 where they winter. Since more beaked whales are caught along the Pacific
Ocean coast of Honshu than in the Sea of Japan, it has been assumed
that most of the herd wintering in southern Japan migrates north along
the eastern side of the islands of Japan. Here they are seen in early May
with the maximum number sighted in the waters of Japan in July and
August, fewer in September, and hunting ceases altogether in November.
Apparently a part of this herd (population) spends the summer months
in the waters of Japan and another part farther north since, in May, June,
and July, they are noticed along the eastern coast of Kamchatka and in
August near the Commander Islands, Karagin Island, and in Olyutorskiy
Gulf (Zenkovich, 1939).

In the Sea of Japan, beaked whales are seen earlier than along the
coasts of the more southward Honshu Island. In March they are sighted
in Peter the Great Gulf and in April, along the northeastern coasts of
Hokkaido in the Sea of Okhotsk. In May the hunting of beaked whales in
Hokkaido increases considerably, lessens in June, and ceases almost alto-
gether by July and August. In September and October the second arrival
of beaked whales occurs off the Hokkaido coasts and-their catch in these
months increases again. It is possible that an independent population of

859

beaked whales lives here, migrating in summer into the Sea of Okhotsk
and again appearing in autumn during autumn migrations to the winter-
ing sites.

A preponderance of males has been recorded in the annual catch of
beaked whales from the coastal stations in the waters of Japan. In five
years of hunting, they totaled 68.3% of the total catch. It would seem that
whales of different sex (and probably of age) groups migrate at different
periods by different routes (Omura, Fujino, and Kimura, 1955).

Reproduction. Mating and parturition occur in winter months in
the wintering areas, but have not been established very accurately. The
majority mate in February with most births recorded in December. Both
these periods are perhaps protracted by a few months. On August 5,
1957, a female 1,076 cm long held an embryo 12 cm long. Apparently
this female had mated in summer. On the other hand, on August 21, 1927,
a female caught in the same region was found with an embryo 242 cm
long. This female had perhaps mated in winter. Gestation extends for
about 10 months (Omura, Fujino, and Kimura, 1955; Omura, 1958).

Growth and development. Adults and embryos differ in body propor-
tions (Omura, Fujino, and Kimura, 1955). Among embryos: (1) the head
is relatively large; (2) flippers are shifted closer to the tail; (3) flippers
are relatively large; (4) navel caudally situated although position of the
anal opening is the same as in adults; (5) dorsal fin is relatively very
high although its relative length along the base is the same as in adults;
and (6) caudal flukes at the point of their articulation with the caudal
stem are relatively broader. The average size of newborn calves is 4.6 m.
Calves 580 cm long were still suckling.

The animals attain sexual maturity at three years of age or slightly
later at an average body length of 10.1- 10.4 m (females) and 9.7- 10.1 m
(males) (Table 63).

The smallest pregnant female was 9.8 m long and the largest imma-
ture female 10.4 m (Tomilin, 1962).

Enemies, diseases, mortality, parasites, and competitors. It would
appear that Pacific beaked whales have no enemies. Their diseases
have not been studied. From among the ectoparasites, whale lice of
the genus Platicyamus have been found. Some instances of barnacles
(Conchoderma) adhering to the teeth have also been recorded.
Overgrowth of diatomaceous algae Cockoneis sp. and Navicula sp.
and adhesions caused by suction of the Pacific lamprey (Entosphenus
tridentatus) have also been observed.

Nine species of endoparasites are known: trematodes one, cestodes
two, nematodes four, and acanthocephalans two. The trematode
Oschmarinella sobolevi Skrjabin has been detected in the bile ducts of the

860

637 Table 63. Dimensions of Pacific beaked whales caught in the waters of Japan (Omura,

637

Fujino, and Kimura, 1955)

Length of whale, m Caught
Males Females Total
5.5 — 1 1
5.8 — 1 Л
6.1 — 1 1
7.0 7 3 5
73 1 4 5
7.6 1 1 2
7.9 1 — 1
8.2 6 7 8
8.5 4 4 8
8.8 8 6 14
9.1 25 17 42
9.4 19 11 30
9.7 38 17 55
10.1 76 18 94
10.4 149 38 187
10.7 158 50 208
11.0 91 61 152
11.3 43 33 76
11.6 9 20 29
11.9 —- 4
12:2 — it 1

liver in beaked whales only from Kuril waters. Neither of the cestodes
Trigonocotyle sp. and Tetrarhynchidae gen. sp. has been identified at the
species level. Of the four species of nematodes found in beaked whales,
Anisakis (Anisakis) simplex Rudolphi, parasitizing the gullet, stomach,
and intestine, is very widely distributed among marine mammals. It is
known among many species of toothed and baleen whales and Steller’s
sea lion among Pinnipedia (detected in the North Sea, off the coasts of
Kamchatka, Japan, and New Zealand). The nematode Anisakis skrjabini
Mosgovoy, found in beaked whales and sperm whales in the waters of the
Commander Islands, the Sea of Okhotsk, and in the Antarctica, localizes
in the stomach and small intestine. Crassicauda giliakiana Skrjabin and
Andreeva has been found in the kidneys and ureter of beaked whales and
belugas of the Sea of Okhotsk and Delamurella hyperoodoni Gubanov in
the lungs (tracheae) of only the beaked whale of the Sea of Okhotsk.
The acanthocephalan Bolbosoma nipponicum Yamaguti detected in the
intestine of beaked whales from Kuril waters is also known among three
species of baleen whales and two species of pinnipeds of the Kuril Islands
and the seas of Japan and Okhotsk. Echinorhynchus gadi, detected in the

638

861

stomach of beaked whales, is а known parasite of fish (Delamure, 1955;
A. Skrjabin, 1959, 1960).

The extent of mortality among beaked whales is not known. These
whales face some competition for food from sperm whales.

Field characteristics. The body length is around 10 m (for adults)
and the color monochromatic—dark brown on top and lighter below.
The “beak” is flattened. The anterior mandibular section has two pairs of
teeth. The much larger first pair projects outward with the mouth closed.
The dorsal fin is high and well-proportioned and situated above above the
anal opening. These animals live in small groups, dive simultaneously,
and never display the caudal flukes. The blow is small, at a height of
1.0-1.5 m, resembles a flash, and is bushy. (V.A.)

Economic Importance

Minimal but regular hunting of Baird’s beaked whale is carried out only
in the waters of Japan. At other places this whale has no economic
importance (Table 64).

The Japanese and Soviet whaling fleets catch some ten beaked whales
every year while hunting for large whales; the coastal stations of British
Columbia and California on the North American coast also catch a few
of these whales.

As these whales are of small dimensions, the quantum of products
obtained from them is also comparatively small (Table 65).

The quantity of fat obtained from one average size whale is, on
average, 2,700 kg (Khar’kov, 1940).

Japanese whalers catch beaked whales using small boats equipped
with small-bore harpoon guns in the same manner as in hunting large

Table 64. Catch of beaked whales in the waters of Japan (international whaling

statistics)
Year Whales caught Year Whales caught
1948 73 1959 186
1949 92 1960 147
1950 186 1961 133
1951 252 1962 145
1952 321 1963 160
1953 262 1964 189
1954 — 1965 172
1955 258 1966 171
1956 297 1967 107
1957 186 1968 117

1958 229

638

862

whales using large whale boats. The animals caught are brought to the
coastal stations.

The fat of these whales is used only for commercial purposes. The
fat from the jaws (from the mandibular hollow space) and around the
jaws serves as raw material for producing a high-quality oil useful for
lubricating fine mechanisms. The flesh is generally not used as food but,
after cooking, can be used for feeding fur-bearing animals. In some cases,
however, the flesh of these whales is processed for human consumption
also.

Based on their population, the hunting of beaked whales could be
somewhat expanded. Rules for controlling hunting have not been formu-
lated and, at the present rate of their exploitation, there is yet no need
for such. (V.A.)

Genus of Beaked Whales [Sword-tooth Dolphins]
Genus Mesoplodon Gervais, 1850

1850. Mesoplodon. Gervais. Ann. Sc. Nat., Zoologie, 14, p. 16. Delphinus
sowerbiensis Blainville = Physeter bidens Sowerby. (V.H.)
Medium body dimensions, with a length up to 6.7 m.

The dorsal fin is low and its apex flexed posteriorly. The flippers
are pointed at the tips. The bulge of the crescent-shaped blowhole faces
backward. The sloping forehead transits gradually into a fairly long beak.
The body color is dark (from black to gray) and the abdomen light-
colored.

The long and thin rostrum is narrower on top than in other mem-
bers of the family of beaked whales and exhibits mesorostral ossification,

Table 65. Weight of body parts of Baird’s beaked whales (Zenkovich, 1937b; Tomilin,
1951; Sleptsov, 1961)
Body part Body length and sex
Female Female ?
10.0 m 10.8 т 11.1 м
Subcutaneous fat 3,337 2,258 5,616
Flesh 2,400 2,838 1,9741
Skeleton 1,953 1,256 3,050!
Viscera 958 1,148 399
Total 8,648 7,500 11,039

Note: The heart, lungs, and liver of another female, 11.1 m long (pregnant), weighed 54,
281, and 64.5 kg respectively (Khar’kov, 1940).
['These figures are possibly reversed.—Ed.]

863

formed through ossification and fusion with the surrounding bones of the
vomer. The supraoccipital and interparietal bones form a high crest. The
frontal bones have small orbital processes. The mandibular symphysis is
considerable in size (one-fifth to one-third the jaw length).

One pair of teeth, strongly compressed laterally, is located at the
level of the anterior third of the lower jaw (at the end of the jaw only in
М. muirus) and projects outside the mouth when it is closed, while cov-
ering the upper jaw from the sides. The roots of the teeth run obliquely
to the longitudinal axis of the jaw. Among some species of these beaked
whales, the teeth are sharply enlarged; they are larger among males than
females. Numerous minute teeth are concealed in the gums of the upper
and lower jaws, particularly among young animals (they may be resorbed
with advancing age).

Cervical vertebrae 7, thoracic 9-11, lumbar 9-12, and caudal 18-21;
total 46-48. The atlas and axis are fused; sometimes a third cervical
vertebra is attached to them. The spinous vertebral processes are very
large. The sternum has four (less often five) sections. Of the five digits
on the forelimb, the second and third are the longest.

Biology very poorly studied. These are pelagic whales, rarely encoun-
tered along coasts. They remain singly, in small groups, or much larger
herds. They apparently feed mainly on cephalopods. The periods of mat-
ing and parturition are very prlonged.

The distribution is extremely wide: in the eastern part of the Atlantic
Ocean, from Norway and the British Isles to Madeira, the Mediterranean
Sea, and coasts of South Africa; in the western Atlantic, from Newfound-
land and Canada; and in the Caribbean Sea, from Trinidad and the coast
of Argentina (42° S lat.) to the Falkland Islands. In the Pacific Ocean, in
the east from the Bering Sea to La Jolla in California in the north and
along the coasts of Chile in the south, and in the west from the Bering
Sea to Japan and from Australia to New Zealand; and the Indian Ocean
(Hershkowitz, 1966). Abundant everywhere (Fig. 371).

Fossil remains have been found in the Upper Miocene of North
America and in the Upper Miocene and Middle Pliocene of Europe.

The genus comprises 10 (Nishiwaki and Kamiya, 1958; Hershkowitz,
1966) or 11 (Moore, 1968) species: M. bidens Sowerby, 1804—North
Atlantic from Norway, the Baltic Sea, and Great Britain in the east to
the Mediterranean Sea (inclusive) and in the west from Newfoundland
to Massachusetts state (USA); M. europaeus Gervais, 1848 - 1852—North
Atlantic (English Channel and from New York to Florida, Gulf of Mex-
ico, and also the Caribbean Sea from Cuba to Trinidad); M. mirus True,
1913—North Atlantic (British Isles and coasts of France and from Cape
Breton and Nova Scotia peninsula to Florida), South Atlantic (southern

Ц

И in ИНЫХ
Uystglystetadalateletate
|

Е
a
$
3
8
8
8

Fig. 371. Range of the genus of beaked whales [sword-tooth dolphins], Mesoplodon (V.A. Arsen’ev).

640

641

865

coasts of South Africa); М. grayi Haast, 1874—Indian Ocean (coasts of
South Africa and Australia), South Pacific Ocean (from New Zealand
to Chilean coasts), South Atlantic from the coasts of Argentina and the
Falkland Islands to South Africa, North Atlantic (Netherlands coast);
M. ginkgodens Nishiwaki and Kamiya, 1958—North Pacific Ocean from
Japan in the west to California in the east and coasts of Sri Lanka
in the Indian Ocean; М. layardi Gray, 1865—South Pacific Ocean and
Indian Ocean from Australia and New Zealand to the South Atlantic
between the Falkland Islands and South Africa; М. densirostris Blainville,
1817—eastern part of the Atlantic Ocean (coastal waters of Madeira and
South Africa), western Atlantic (Nova Scotia peninsula, Canada, and

' Bahama Islands), western part of the Pacific Ocean (Lord Howe Island

and Queensland in Australia), North Pacific Ocean (Midway Island), and
Indian Ocean (from South Africa and the Seychelles to apparently west-
ern Australia); М. stejnegeri True, 1885—Pacific Ocean from the Bering
Sea to Oregon state (USA) in the east and Japan in the west; M. bow-
doini Andrews, 1908—New Zealand coast; M. carlhubbsi Moore, 1946
[1963]—North Pacific Ocean (east coast of Japan and west coast of the
USA between 32° and 47° N lat.); and M. hectori Gray, 1871—Southern
hemisphere, temperate waters.

The systematics of the genus of beaked whales has not been suffi-
ciently developed. Many species have been described on the basis of just
one or two specimens and the differences between some of them repre-
sent only individual variations. Thus, it is possible that M. stejnegeri, M.
bowdoini, M. ginkgodens, and M. carlhubbsi, inhabiting the Pacific Ocean,
belong to the same species, as conceded by some scientists who regard
the differences between these species as inconsequential. The main dif-
ferences are: the lateral thickness of the tooth crown is greater than
its front-to-back width—in M. stejnegeri by seven times, M. ginkgodens
six times, and М. bowdoini three or four times; and the ргетахШагу
foramena lie at the same level as the maxillary foramena or posterior
to it in M. stejnegeri and M. bowdoini, but markedly more anteriorly in
M. ginkgodens. There are other differences also. The collection of more
factual data should clarify the systematics of the genus.

On the other hand, the differences between some species (groups
of species) are quite significant. Thus, M. layardi Gray is isolated in a
special subgenus, Dolichodon Gray, M. densirostris Blainville in subgenus
Dioplodon Gervais, and all the rest of the species placed in the subgenus
Mesoplodon Gervais (Moore, 1968).

Only one species, Stejneger’s beaked whale, M. stejnegeri, has been
reported in USSR waters. Some other species may also be encoun-
tered: in the southern part of the Baltic Sea—Sowerby’s beaked whale,

642

866

М. bidens; in the seas of the Far East—M. carlhubbsi and М. ginkgodens.
The latter two species could, however, be regarded as belonging to the
species M. stejnegeri and hence are not described here.

There is no special hunting of these beaked whales. (V.S.)

STEJNEGER’S BEAKED WHALE
Mesoplodon (Mesoplodon) stejnegeri True, 1885

1885. Mesoplodon stejnegeri. True. Proc. U.S. Nat. Mus., 8, p. 585. Bering
Island and Commander Islands. (V.H.)

Diagnosis

Total body length up to 6 m.

Color black, more light-colored ventrally. The antorbital notches in
the skull are developed relatively poorly. The foramena of pair V of the
nerves in the maxilla lies anterior to that in the premaxilla or at the same
level. The palatine bones are not adjacent. The ratio of length of tooth
crown to its width is more than in the other species of these beaked
whales (6.4:1 or even slightly more). (V.S.)

Description

The lower jaw is longer than the upper by 15 mm. The flippers are
relatively small. The caudal flukes are lighter in color than the trunk
and may be white on the underside. Quite a large number of small white
patches (traces of scars or parasitic infections) are present on the body
surface (Fig. 372). The teeth are strongly built in the males (Fig. 373).
but barely emerge from the gums in the females.

The following are the main body measurements of an adult male
Stejneger’s beaked whale (Tomilin, 1957) (in cm): body length 509; max-
imum girth of body 254; distance from tip of lower jaw to base of flip-
pers 112; distance from tip of upper jaw to anterior margin of blowhole
59; distance from anal opening to notch between the caudal flukes 141;

Fig. 372. Stejneger’s beaked whale, Mesoplodon stejnegeri (figure by N.N. Kondakov).

867

(9)

Fig. 373. Head of Stejneger’s beaked whale, Mesoplodon stejnegeri, lateral view
(figure by N.N. Kondakov).

length of flippers along the anterior (lower) margin 55; maximum width
of flippers 16; height of dorsal fin 21; length of dorsal fin along the base
34; and spread of caudal flukes 128.

The main measurements of the skull (Fig. 374) of male Stejneger’s
beaked whales caught off the coasts of Oregon and California (two speci-
mens), and Japan (one specimen) (Tomilin, 1957; Nishiwaki and Kamiya,
1959) are respectively (in cm): condylobasal length 81, 82, and 80; zygo-
matic width 39, 38, and 38; length of rostrum 49, 51, and 49; width of
rostrum at base 23, 22, and 16; length of lower jaw 69, 71, and 68; and
length of mandibular symphysis 18,—, and 18. (V.S.)

Fig. 374. Skull of Stejneger’s beaked whale, Mesoplodon stejnegeri (figure by
N.N. Kondakov).

643

868

Geographic Distribution and Biology

Pacific Ocean from the Bering Sea to the Californian coast in the east —
and coasts of Japan in the west (information on distribution is based
exclusively on the sites of animals stranded on coasts; field observations
are almost nil).

Geographic Range in the USSR (Fig. 375)

Known only from the waters of the Commander Islands where three
instances of beached beaked whales have been recorded on the shores of
the Bering Sea. Their presence is possible in the waters of Kamchatka
and the Kuril Islands.

Fig. 375. Range of Stejneger’s beaked whale, Mesoplodon stejneger1, in the USSR
(V.A. Arsen’ev).

643

645

869
Geographic Range outside the USSR (Fig. 376)

Aleutian Islands (one record), Alaskan peninsula (one record), Pacific
Ocean coast of North America (five records off the coasts of Washington,
Oregon, and California states), and Japanese coasts (two records in the
region of Ayukawa) (Tomilin, 1962; Hershkowitz, 1966).

Geographic variation has not been established.

Biology virtually not studied since the species is very rare through-
out its range. Cephalopods apparently serve as the main food although
some individual whales have been sighted at the sites of congregations
of salmon. The very small number of observations indicate that they live
singly or in very small groups (up to three animals). In life style, they are
probably similar, if not identical, to Sowerby’s beaked whale (M. bidens
Sowerby, 1804).

Adult animals are about 6 m long and black with a light-colored
underside. Teeth one pair, strongly compressed laterally, project from
the sides, and cover the upper jaw when the mouth is closed.

These whales are of almost no economic importance. (V.A.)

SOWERBY’S BEAKED WHALE
Mesoplodon (Mesoplodon) bidens Sowerby, 1804

1804. Physeter bidens. Sowerby. Trans. Linn. Soc. London, 7, p. 310.
Elginshire, Scotland.

1817. Delphinus sowerbensis. Blainville. Nouv. Dict. Hist. Nat. 9, p. 177.
Substituted for Physeter bidens Sowerby. (V.H.)

Diagnosis

Maximum overall body length up to 5.5 m.

Body dorsally black or bluish-black and usually grayish or whitish
ventrally. The antorbital notches are well developed in the skull. The
foramena of pair V of the nerves in the maxilla is located posterior to
the foramena in the premaxilla. The palatine bones are adjacent. The
rostrum is broad at the base. The teeth lie posterior to the mandibu-
lar symphysis. The ratio of length of tooth crown to its width is 3.5:1.
(V.S.)

Description

The head is slightly compressed laterally. The “beak” is slightly
flattened dorsoventrally (Figs. 377 and 378). The lower jaw is only

п’еу).

ge of Stejneger’s beaked whale, Mesoplodon stejnegeri (V.A. Arse

Fig. 376. Species ran

644

645

645

871

Fig. 377. Sowerby’s beaked whale, Mesoplodon bidens (figure by М.М. Kondakov).

V2)

Fig. 378. Head of Sowerby’s beaked whale, Mesoplodon bidens (figure by
N.N. Kondakov).

insignificantly longer than the upper (roughly by 13 mm). The body
color is variable—from black to gray with abdomen light, sometimes
white. The anterior margin of the caudal flukes and the upper and lower
jaws may be light-colored. Numerous whitish spots and bands—remnants
of scars—cover the body. The teeth of males are much larger than in
females; in the latter, the teeth in most cases do not emerge from the
gums.

The basic body measurements of two female Sowerby’s beaked
whales are as follows (Tomilin, 1957) (in cm): body length 345 and 431;
distance from tip of snout (in second animal from tip of lower jaw) to
flippers 91 and 109.8, up to blowhole 44 and —; length of flippers 30
and 39.2, their width 12 and 13; distance from tip of snout to dorsal fin
204 and 275.8; height of dorsal fin 27 and 20.9, its length 49 and 36.6;
and spread of caudal flukes 68 and 99.4.

The basic skull measurements (Fig. 379) of three males caught off the
Shetland Islands, Norway, and Sweden (Tomilin, 1957) are respectively
(in cm): condylobasal length 74, 73, and 74; zygomatic width 29, 30, and
27; length of rostrum —, 48, and 50; width of rostrum at base 18, 19,
and 17; length of lower jaw 70, 64, and 64; and length of mandibular
symphysis —, 21, and 22. (V.S.)

872

Geographic Distribution

North Atlantic Ocean from the coasts of Norway, the British Isles, and
the Baltic Sea up to the Mediterranean Sea in the east and from New-
foundland to Massachusetts state along the American coast (the distri-
bution of the species was established exclusively from information on
beached animals).

Geographic Range in the USSR

Stray transgressions are possible into our waters in the Baltic Sea.

Geographic Range outside the USSR (Fig. 380)

Finds of beached animals are known in Newfoundland and Massachusetts
state on the North American coast. In the eastern part of the North
Atlantic, they have been registered on the coasts of Norway, Sweden,
England, Poland, Holland, the FRG, the GDR, Belgium, and France.
This whale is encountered in the Mediterranean Sea (Sergeant and
Fisher, 1957). (V.A.) ;

645 Fig. 379. Skullof Sowerby’s beaked whale, Mesoplodon bidens (figure by
N.N. Kondakov).

646

646

Fig. 380. Species range of Sowerby’s beaked whale, Mesoplodon bidens (V.A. Arsen’ev).

Geographic Variation
Not established.
Biology

Very poorly studied. Absolutely nothing is known about the population
of Sowerby’s beaked whales. Information on their food is not available.

Behavior. These whales are seen more Often singly or in pairs. In the
shallow-water bays of Khertvik on Gassen Island (Norway), the “tech-
nique” of two of these whales stranded on the coast was observed on
April 18, 1957. Initially, they swam along the coast to a shallow point
almost touching the bottom. For a quarter of an hour they attempted to
return to the sea but a low tide occurred and the exit from the bay became
very Shallow. Then, the larger of the two whales turned directly toward
the coast and soon found itself on a sandy beach. It quivered its tail vigor-
ously and within minutes a large pit had formed in the sand. Meanwhile
the smaller whale swam along the coast but having covered a distance
of only 20 m slammed against some rocks and remained motionless; it
resisted only slightly when its tail was looped. From the first sighting
of these animals along the coast to their death, no more than 30 min
elapsed. One was an adult female 505 cm in body length and the other
a large female calf 315 cm long (Jonsgard and Hoidal, 1957).

Seasonal migrations and transgressions. In La Mancha Strait and on
the coasts of Great Britain, beached Sowerby’s beaked whales have been
found in March, April, September, and December; in the Baltic Sea,

647

874

such finds have been recorded in February, June, August, and September.
These animals are believed to inhabit these waters throughout the year.
Their migrations have not been traced.

Reproduction, growth, and development. Mating and parturition occur
at the end of winter and spring. The period of mating is quite prolonged.
Gestation extends for about a year. The body length of a calf at birth
varies from 152 to 213-244 ( 183) cm. In the period of lactation, the
calf adds 90-120 cm to its length. Suckling ceases at a body length of
about 300 cm (Jonsgard and Hoidal, 1957).

Enemies, diseases, mortality, parasites, and competitors. Enemies of
Sowerby’s beaked whale are not known. The diseases have not been
studied.

An endoparasite (Conchoderma sp.) was found on the teeth of
Sowerby’s beaked whale. Six species of helminths are known. Cysts of the
trematode Monostomum delphini Diesing were found in the blubber of a
whale from Gavra region but the highly superficial description does not
permit positive identification of the species affinity of this parasite. The
cestode Strobilocephalus triangularis Diesing, detected in the intestine of
Sowerby’s beaked whale, is also known in the bottlenose whale and two
or three species of dolphins. It has been found off the coasts of Portugel
and Brazil. The cestode Tetrabothrium forsteri Krefft, parasitizing the
intestine of Sowerby’s beaked whale, was also detected in two species
of dolphins from the Mediterranean Sea and New Zealand waters. The
widely distributed cestode Phyllobothrium delphini Bosk, parasitizing the
skin of Sowerby’s beaked whale, is also known in seven species of toothed
whales, the Arctic whale, and Weddell’s seal. It has been detected near
the Azores, in the Mediterranean Sea, off the Commander Islands, along
the coasts of Australia, and in the Antarctic.

The nematode Anisakis (Anisakis) simplex Rudolphi is very widely
distributed and, in addition to Sowerby’s beaked whale, parasitizes the
gullet, stomach, and intestine of ten other species of toothed whales, two
species of baleen whales, and Steller’s sea lion. It has been detected in
the North Sea and off the coasts of Kamchatka, Japan, and New Zealand.
The only species of acanthocephalans, Bolbosoma vasculosum Rudolphi,
was found in the intestine of Sowerby’s beaked whale and the common
dolphin from the Atlantic Ocean and the Mediterranean Sea (Delamure,
1955).

Field characteristics. The maximum body length is 5.5 m. The body
is almost black dorsally and slightly lighter ventrally. The lower jaw is
almost not longer than the upper. The teeth almost do not emerge among
females. With the mouth closed, a pair of large flat teeth projects upward
along the sides of the upper jaw in males.

875

Sowerby’s beaked whale has по economic importance and there is
no hunting of this animal. (V.A.)

Genus of Goose-beak [Cuvier’s Beaked] Whales
Genus Ziphius G. Cuvier, 1823

1823. Ziphius. G. Cuvier. Rech. ossem. foss., 5, р. 350. Z. cavirosiris
G. Cuvier.

1846. Xiphius. Agassiz. Nomencl. Zool., p. 389. Correction for Ziphius
G. Cuvier. (V.H.)

Medium size, with body length up to 8 m.

The “beak” is short and transitions smoothly into the low frontal pro-
jection of the corpus adiposum. The jaw line is short. The dorsal fin is low
and highly variable in shape in different animals, from crescent-shaped
to triangular. The flippers are relatively long and narrow. The lower jaw
projects forward more than the upper. The jaw line extends posteriorly to
less than half the distance between the tip of the snout and the eye.

The color is usually dark and subject to considerable individual vari-
ation. The dorsum may be lighter in color.

The skull is asymmetric. Among the members of the family, the ros-
trum of the skull is the widest and shortest (only slightly longer than
the cranium) in this genus. The vomer forms a stable mesorostral ossifi-
cation. Longitudinal maxillary crests are very poorly developed. A large
depression, housing the spermaceti sac, occurs on the dorsal side of the
skull between the outer edges of the maxillae. The mesethmoid bone is
ossified. Posterior to the nostrils, the maxillary, premaxillary, and nasal
bones form a large projection in the form of an arch, which overhangs
the nostrils. The nasal bones attain maximum width anteriorly.

Teeth one pair, located on the anterior tip of the lower jaw, and
projecting anterior to the upper jaw when the mouth is closed.

The sternum consists of five sections. Cervical vertebrae 7, thoracic
9-10, lumbar 10-11, and caudal 19-22; total 46-49. The anterior three
_ or four cervical vertebrae are usually fused. Phalangeal formula: [,, Из,
III;_¢, [У4, and У, ->. Ribs 9 pairs.

Biology not well studied. These animals represent forms of the open
sea and feed apparently on mollusks. The periods of mating and partu-
rition are prolonged.

These whales are distributed in the World Ocean but are few in
number everywhere.

The genus consists of a single species: Cuvier’s beaked whale, Ziphius
cavirostris G. Cuvier, 1823.

There is no special hunting of these whales. (У.5.)

648

876

CUVIER’S BEAKED WHALE

Ziphius cavirostris G. Cuvier, 1823

1823. Ziphius cavirostris. G. Cuvier. Rech. Oss. Foss. ed. 2, 5 (1), p. 350,
pl. 27. French coast in the department of Rhone delta (Bouches-
du-Rhone), Mediterranean Sea.

1826. Delphinus desmaresti. Risso. Hist. Nat. Eur. Mérid., 3, p. 23.
Mediterranean Sea.

1883. Ziphius grebnitzkii. Stejneger. Proc. Ц. $. Nat. Mus., 6, р. 77. Com-
mander Islands. (V.H.)

Diagnosis
Only species of the genus.

Description

The frontal projection is much lower than in bottlenose whales and
Pacific beaked whales (Figs. 381 and 382). The characteristic small oral
cavity is smaller than in other species of the family of beaked whales.
Two longitudinal grooves traverse the throat. The bulge of the crescent-
shaped blowhole faces backward.

648

Fig. 381. Cuvier’s beaked whale, Ziphius cavirostris (figure by N.N. Kondakov).

648

649

877

(9)

Fig. 382. Head of Cuvier’s beaked whale, Ziphius cavirostris (figure by М.М. Kondakov).

The body color is subject to great individual variation. Monochro-
matic black, gray, or bluish-gray animals are encountered. In others, the
ventral side may be lighter in color than the dorsal but the opposite,
albeit less often, has also been recorded. Other color variations are
likewise possible. Many depigmented patches of various shapes and sizes
are seen on the skin. The head and dorsum (up to the dorsal fin) are
sometimes very light-colored in old animals.

The rostrum of the sharply asymmetric skull is poorly set off from
the cranium. Adult males are characterized by a mesorostral ossification
that is more prominent than in females and the young. This ossification
lies in a very deep longitudinal depression of the rostrum between the
premaxillae.

The teeth of males are larger and more massive than in females
(tooth diameter in males 25-29 mm, in females 10-14 mm). Young
animals have 28-30 vestigial teeth in the gums of the upper and lower
jaws. These teeth reduce in number as the animal ages. Cuvier’s beaked
whale has 7 cervical, 9 thoracic, 11 lumbar, and 20 caudal vertebrae.
The proportions of these sections to the length of the entire vertebral
column are respectively 3, 17, 36, and 44% (Slijper, 1936a). The stomach
of Cuvier’s beaked whale has five sections; the first and last are large
and the three intermediate ones small; all of them are smooth-walled
(Kenyon, 1961).

The main body measurements of three females and one male Cuvier’s
beaked whale (Tomilin, 1957; Kenyon, 1961) are respectively (in cm):
body length 579, 584, 658, and 543; distance from tip of snout to posterior
edge of dorsal fin 389, 391, 425, and 350, to the axillae 115, 142, 132,
and 131, to the blowhole 60, —, 66, —; length of flippers 46, —, 56, and
42; maximum width of flippers 16, 17, 17, and —; height of dorsal fin 30,
19, 25, and 21; and width of caudal flukes from tip to tip 162, 182, 175,
and 133. The largest of the animals caught had a body length of 793 cm.

652

649

878

Females are larger than males. Of 51 males and 34 females caught in
Japan, the average length of the males was 550 cm (maximum 671 cm),
of the females 580 (701) cm (Table 66).

The following are the basic skull measurements (Fig. 383) of two
adult males and a female Cuvier’s beaked whale (Tomilin, 1957) (in
cm): condylobasal length 85, 81, and 88; zygomatic width 54, 52, and 53;
length of rostrum 48, 46, and 47; width of rostrum at base 33, 32, and 48;
length of left half of lower jaw 75, 74, and —; and length of mandibular
symphysis 18, 16, and 18. (V.S.)

Geographic Distribution
Warm, temperate, and partly temperate-cold waters of the World Ocean.

Geographic Range in the USSR

Constitutes a very small part of the entire range of the species. Cuvier’s
beaked whale has not been sighted with certitude in our waters of the
basin of the Atlantic Ocean but may be encountered on out coasts of the
Baltic Sea since it is known from Kattegat and waters of southern Sweden
(transgressions): in the Far East, waters of southeastern Kamchatka and
the Commander Islands where beached animals are a regular annual fea-
ture. According to less reliable information, the animal has been sighted
near the Kuril Islands and off the southern coast of Sakhalin (Sleptsov,
1961).

Table 66. Dimensions of Cuvier’s beaked whales caught in Japan (Omura, Fujino, and
Kimura, 1955)

Body length, cm Number of whales
Males Females Total
305 1 — 1
335 a — =
366 — 2 2
396 3 2 5
427 2 1 3
457 4 — 4
488 2 4 6
518 4 7 6
549 9 5 14
579 8 6 14
610 7 5 12
640 7 5 12
671 4 1 5
— 1

879

y =. а /
ON РК)
А
РК м yr

649 Fig. 383. Skull of Cuvier’s beaked whale, Ziphius cavirostris (figure by М.М. Kondakov).

Geographic Range outside the USSR (Fig. 384)

Encountered in the North Atlantic Ocean along the coasts of the British
Isles and southern Sweden, close to the coasts of Spain, Italy, France,
and Corsica (Ligurian Sea), northern Africa, in the Mediterranean Sea,
and in the Atlantic waters of North America. It is known in the Pacific
Ocean around the Hawaiian Islands, in the East China Sea, along the
coasts of Japan, and the Pacific Ocean coast of North America from Cali-
fornia to the Pribilof Islands in the north. In the Southern hemisphere, it
has been observed in the waters of Brazil, Argentina, South Africa, close
to Sri Lanka, Polynesian Islands, around Australia, Tasmania, and New
Zealand. Beached animals have been recorded at the following points:
on the British Isles and Shetland Islands, in Kattegat Strait and on the
coasts of Boguslan (Sweden), in the Baltic Sea, Bay of Biscay, Mediter-
ranean Sea (near Nice, in the Rhone delta, at Malaga near Valencia, on
Corsica, on the coast of Italy at Liguria, Messina Bay, and on the coast
of Algeria), on the coast of North America in the states of New Jersey,
Massachusetts, South Carolina, Rhode Island, California, and Alaska,
and in British Columbia (Fig. 385). A large number of beached animals
is known even in the Southern hemisphere (Tomilin, 1957; Clarke, 1958;
Chapskii, 1963). Comparatively uncommon throughout its range. (V.A.)

*(лэ.цэзгу “W'A) 5мколар smiydiz ‘этеца рэхеэа з.лэтлпо до э8ие1 $э1024$ ‘pee “314 059

i

"7

651 р. 385. Above: Cuvier’s beaked whale near Cape Chernyi, Mednyi Island,
July, 1962. Below: Cuvier’s beaked whale stranded on the shore of Bering Island,
: May, 1960 (photographs by S.V. Marakov).

653

882

Geographic Variation

Not studied.
Biology

Population. Comparatively common in our waters along the eastern coast
of Kamchatka, especially around the Commander Islands; very rare or
sighted occasionally elsewhere.

Food. Data on this aspect are negligible. It can only be assumed that
cephalopods serve as the main food of Cuvier’s beaked whale. Of the
three stomachs dissected on the Commander Islands, one was empty and
two contained the beaks of squids and crystalline lenses of cephalopods
(Tomilin, 1957). The stomach of a female found on Amchitka Island
(Aleutians) held the remains of 1,300 cephalopod mollusks—squids
Gonatus sp.—in the form of beaks, lenses, etc. (Kenyon, 1961).

Behavior. These whales live in small groups, remain on the sea sur-
face for about 10 min, dive together, and spend 10 to 30 min or more
under water.

Seasonal migrations and transgressions. Along the Commander
Islands (in the northern region of distribution), these animals are sighted
in March and continue to be seen until autumn. Along the coasts of
Japan, these whales are caught year-round although the best hunting
season is from May through October, peaking in August. Hunting
is carried out from coastal stations located exclusively on the Pacific
Ocean coast of Honshu and on the northeastern coast of Hokkaido.
In the Sea of Okhotsk and along the coast of the Sea of Japan, where
Baird’s beaked whales are hunted, Cuvier’s beaked whales are not caught
(Omura, Fujino, and Kimura, 1955). Apparently Cuvier’s beaked whales
are also encountered along the coast of California almost year-round, as
supported by the following records of whales stranded at roughly 41° N
lat.: a large female on February 17, 1963; an adult female on September
22, 1959; an adult male on March 2, 1957; and an immature small male
on November 24, 1957 (Mitchell and Houck, 1967).

Reproduction, growth, and development. The periods of mating and
parturition have not been established but in all probability both are
protracted. In 1951, 1952, and 1953, embryos 30, 97, and 213 cm long
were detected in August; embryos 170 cm long in September; and others
43 cm long in October, 1952 (Omura, Fujino, and Kimura, 1955). An
embryo 267 cm long, found in a dead female on Bering Island (Comman-
der Islands), was fully developed (Tomilin, 1957). Probably the size of
newborn Cuvier’s beaked whales is proximate to the above range. Males
attain sexual maturity at a body length of 5.5 to 5.9 m; their testes weigh

883

3.5 to 4.2 kg; females become mature at a body length of about 5.5 т. А
female 658 cm long, cast on Amchitka Island (Aleutians), contained at
least two traces of corpora lutea of pregnancy. This female had 24-28
layers in her tooth dentine and thus her age was estimated at 12-14
years. A male with a body length of 544 cm, found on the same island,
had 13 layers in the tooth dentine or was six-seven years of age (Kenyon,
1961).

Enemies, diseases, parasites, mortality, and competitors. Two species
of nematodes and one of cestodes are known in Cuvier’s beaked whale.
The nematode Crassicauda crassicauda Creplin parasitizes, in addition
to Cuvier’s beaked whale, the urogenital system of another species of
toothed whales and six species of baleen whales; Crassicauda boopis
Baylis, parasitzing the urogenital system of Cuvier’s beaked whale, the
humpback whale, and the fin whale, was found in the Atlantic Ocean in
the Northern and Southern hemispheres. A cestode, Phyllobothrium sp..,
was detected in the blubber of Cuvier’s beaked whale (Delamure, 1955;
Tomilin, 1962).

Field characteristics. The body length of adult whales averages 5-7 m.
The “beak” is broad and short and the forehead low and sloped. The
blowhole is crescent-shaped and its horns turned anteriorly. Teeth one
pair, located in the anteriormost part of the lower jaw, and not con-
cealed by the upper jaw, projecting anterior to its edge when the mouth
is closed. These animals live in small groups. While diving, the group
disappears under water in unison. (V.A.).

Economic Importance

Only a few tens of Cuvier’s beaked whales are caught annually and as
such this species has no economic importance whatsoever. It is caught
along with other species of small whales from the coastal stations along
the Pacific Ocean coast of Japan. From 1948 through 1955, 3, 10, 27, 35,
36, and 40 animals were caught annually (Omura, Fujino, and Kimura,
1955).

Sexually mature males ranging from 5.5 to 6.5 m in length constituted
the major part of the catch.

Weight of the Body Parts of Male Cuvier’s Beaked
Whale, Body Length 658 cm (kg) (Kenyon, 1961)

Flesh, blubber, bones (including head) О)
Heart 15.4
Lungs 39.4
Liver 25.9

Kidneys 16.4

884

Reproductive organs 20.7
Stomach:

Empty 173)

Its contents 18.6
Intestine with contents 41.3
Spleen 0.1
Aorta, bronchi, and connective tissue 31.6
Blood (some inevitably lost) 8.1
Total 2.9837

Hunting is carried out mainly from small boats and using small-bore
654 harpoon guns. A good number of animals fall into nets. The products
are used like those of other small whales. On the Commander Islands
the flesh and blubber of beached Cuvier’s beaked whales are fed to dogs
and foxes.
There is no hunting of this whale in our range. (V.A.)

Genus of Bottlenose Whales

Genus Hyperoodon Lacépéde, 1804

1804. Hyperoodon. Lacépéde. Hist. Nat. des Cétacées, рр. XLIV, 319.
Hyperoodon butskops Lacépéde = Balaena ampullata Forster.

1804. Anarnak. Lacépéde. Ibid., pp. XXVIII, 164. Anarnak groenlandicus
Lacépéde = Balaena ampullata Forster. (V.H.)

Medium-size whales.

The “beak” is prominently displayed, more sharply than among other
species of the family. The frontal projection of the prominent corpus
adiposum [forehead] descends rather steeply from the base of the beak.
The corpus adiposum is better developed in males than in females. Two
to four longitudinal grooves occur on the throat. The dorsal fin is flexed
along the posterior edge.

The body is blackish-gray but light gray or yellowish-gray on the
abdomen.

In the skull the maxillae dorsally form at the base of the rostrum
two (right and left) lateral crests which are particularly well developed in
adult males. The rostrum is long. The nasal bones are large and concave
at the top and the mesethmoid bone is incompletely ossified. Together
with the premaxillae, the former bones rise posterior to the bony nares
vertically upward and even overhang them. The maxillae have high crests
that extend along the edges of the rostrum. One pair, less often two pairs
of teeth develop at the tip of the lower jaw. The teeth are larger in males

655

885

than in females. Cervical vertebrae 7, thoracic 9 (8), lumbar 9-11, and
caudal 18-20; total 43 - 46. The cervical vertebrae are fused. The sternum
consists of three sections. Phalangeal formula: I, _5, Пб-в, Ш, [V4-5,
and V,_3.

These animals feed mainly on cephalopods. The periods of mating
and parturition are prolonged. The duration of gestation is about a year.

They are distributed in the North and South Atlantic Ocean, and in
the Indian and Pacific oceans. Seasonal migrations take place.

The genus comprises two species: northern bottlenose whale,
Hyperoodon ampullatus Forster, 1770, and southern flat-faced bottlenose
whale, Hyperoodon planifrons Flower, 1882.

Mesoplodon planifrons [Mesoplodon pacificus|, described by Long-
man in 1926, is sometimes treated as a synonym of Hyperoodon plani-
frons (Hershkovitz, 1966), or as a subspecies of Mesoplodon mirus Moore,
1960 (Moore, 1960), or as a lone species of a different genus, Indopace-
tus Moore, 1968 (Moore, 1968). Additional data are required before an
accurate conclusion can be drawn (descriptions are based on just two
skulls).

Only the northern bottlenose whale is encountered in the seas of the
USSR. There is no special hunting of this species in our waters. (V.S.)

NORTHERN BOTTLENOSE WHALE
Hyperoodon ampullatus Forster, 1770

1770. Balaena ampullata. Forster. Kalm’s Travel into N. America, I, p. 18.
Maldon, Essex, England.

1776. Balaena rostrata. Muller. Zool. Danicae. Prodr., p. 7. Waters of
Denmark and Norway.

1789. Delphinus butskopf. Bonnaterre. Tabl. Encycl. Méth. des Trois Reg-
nes de la Nature. Cectologie, р. 25. Honfleur, France.

1822. Delphinus hyperoodon. Desmarest. Encycl. Méth. Mamm., 2, p. 520.
Thames, England. (V.H.)

Diagnosis
Only species of the genus found in waters of the USSR.
Description

Dorsal fin with a concave posterior margin and located above the anal
opening (Fig. 386). The flippers are larger in males than in females. The
bulge of the crescent-shaped blowhole faces the tail. Two (sometimes

655

656

886

<.

Fig. 386. Northern bottlenose whale, Hyperoodon ampullatus (figure by М.М. Kondakov).

four) longitudinal grooves extend along the throat. Four hairs are seen
on each side of the snout in the embryo.

The body is blackish-gray dorsally and light gray ventrally. The fins
are grayish-black. With advancing age, the color turns light, almost yel-
low. The skin is covered with numerous white spots, apparently caused
by a fungal disease.

Two pairs of teeth are seen only in some adult animals. In females
and many males the posterior pair of teeth does not emerge or, if it does,
these teeth are notably smaller than the anterior pair, which may reach
a height of 6.5 cm. The anterior teeth are set at the tip of the lower jaw
and project Outside when the mouth is closed. The second pair of teeth
(when present) is separated from the first pair by a considerable gap.

Males are larger than females. The largest of the measured males
was 9.4 m in length, of females 8.7 m.

Measurements (as percent of animal length) of a young male with a
body length of 518 cm and an adult female 716 cm long (Tomilin, 1962)
are respectively: from tip of snout to commencement of dorsal fin 52.9
and 57.4; height of dorsal fin 5 and 5.3, length of dorsal fin 7.8 and 8.5;
spread of caudal flukes between tips 27.1 and 29.1; length of flippers 10.7
and 8.5, and width of flippers 5.3 (female).

The skull dimensions are (Tomilin, 1957) (in cm): condylobasal

length of skull 145 (100%), zygomatic width 68 (46.9%), length of

655

887

Fig. 387. Skull of the northern bottlenose whale, Hyperoodon ampullatus (figure
by N.N. Kondakov).

rostrum 89 (61.4%), width of rostrum at base 35 (21.4%), length of lower
jaw 135 (93.1%), and length of mandibular symphysis 25 (17.2%). (V.S.)

Geographic Distribution

North Atlantic Ocean, usually at depths exceeding 1,000 meters (Jons-
gard, 1952).

Geographic Range in the USSR

Rare in our waters but sometimes encountered in the central part of the
Barents Sea, along the Murman coast, and less often along the western
coasts of Novaya Zemlya. Transgresses occasionally into the White Sea
where instances of some animals being caught are known. Transgres-
sions have also been observed in the southern part of the Baltic Sea (in
winter).

Geographic Range outside the USSR (Fig. 388)

In the eastern part of the North Atlantic, these animals inhabit
waters from the Cape Verde Islands in the south (15 to 18°N lat.),

656

657

888

NaS,
“PAR

30

Fig. 388. Species range of the northern bottlenose whale, Hyperoodon ampullatus
(V.A. Arsen’ev).

along the entire European coast, and up to the Arctic. They cover
the Mediterranean Sea, are encountered along the coasts of Holland,
Great Britain, Shetland and Faroe Islands, and coasts of Norway up to
Varangerfjord. They inhabit the waters of Iceland and Jan Mayen Island,
are common in the Norwegian and North seas, and penetrate the waters
of Spitsbergen in the zone of influence of the Gulf Stream (75-78° N
lat.) toward the coast of Greenland. They transgress into the southern
part of the Baltic Sea (Kiel and Lubeck bays, Rugen Island) but are
rare on the Baltic coasts of Sweden and Norway. Known in the west
from Rhode Island in the south up along the coast of North America
(New York and Massachusetts bays) to Newfoundland, Hudson, Davis,
etc. (Tomilin, 1957, 196210; Chapskii, 1963). (V.A.)

Geographic Variation

Not established.

10 References to the existence of this bottlenose whale in the Pacific Ocean, extensively
cited in the literature, including the latest reviews (see, in particular, Hershkovitz, 1966:
“Bering Sea and Japan’’), are erroneous and pertain to Вана’; beaked whale (Berardius
bairdi, Tomilin, 1952, 1957; see latter). (У. H.)

889
Biology

Population. The population of the northern bottlenose whale, compared
with that of large whales, is very small. Hunting has never exceeded 500
bottlenose whales per year.

Food. Cephalopods (Gonatus fabricu, Onychoteuthis sp., Sepia sp.,
and Loligo sp.) constitute the main food. Most of the stomachs inves-
tigated revealed only the beaks of cephalopods, which numbered up to
10,000 in one stomach. Herring, cod, sea-cucumbers, and starfish were
found as random food objects. The characteristic structure of the teeth
and the ability of the northern bottlenose whale to dive to great depths
confirm the view that these whales specialize in feeding on cephalopods
(Tomilin, 1957).

Daily activity and behavior. This whale is usually encountered in small
groups, sometimes up to 20 animals. Single animals are rare. Groups
most often comprise animals of different sexes and ages, including calves.
Such groups resemble the harems of sperm whales since usually a single
large male is seen in them. However, groups consisting exclusively of
adult males have also been observed. At places of food concentration,
groups of bottlenose whales sometimes gather into large herds of up to
a hundred or more animals.

Like the sperm whale, the northern bottlenose whale possesses the
ability to descend to great depths and remain submerged for a long while.
It has been suggested that the duration of its underwater residence may
be as much as one hour. Herein lies some similarity of behavior during
diving with the sperm whale. On surfacing after prolonged submergence,
the northern bottlenose whale inhales/exhales quite a number of times
and remains on the surface for a long time. It then undertakes another
long dive. While resting, up to thirty brief intermediate submergences,
one every 30 - 40 sec, producing small (50 -60 cm high), barely perceptible
blows have been recorded for this whale. A display of the caudal flukes
during diving has never been seen. The auditory faculty is thought to be
very well developed in the northern bottlenose whale and it is sensitive
to extraneous noises. The herd instinct is strong in these animals and
they do not usually abandon an injured mate even when confronted with
imminent danger themselves.

Seasonal migrations and transgressions. The northern bottlenose
whale performs regular seasonal migrations but the courses and periods
of its spring and autumn migrations have hardly been studied. It is only
known. that these whales inhabit the northern part of their range in the
summer months and enter the southern part in winter. Spring migrations
commence early since they are noticed off the coasts of Norway and

658

890

even Jan Mayen in March and April. The line of sharp increase in depth
represents the boundary of distribution of these whales and they usually
transgress no farther. Thus they never enter the fjords. Only one case
is known; a small herd of bottlenose whales appeared in Oslo Fjord in
September, 1939 and one whale beached (Jonsgard, 1952*). In June these
whales are sighted along ice edges up to 77° N lat. and have even been
encountered (sometimes in large numbers) among drifting ice. Off the
coasts of Norway and in other regions of their hunting, the maximum
number of bottlenose whales is caught in May and a slightly smaller
number in June @stby, 1956, 1957, 1958, 1959). It has been assumed
that this species penetrates farthest north in spring and later descends
slightly southward to inhabit the waters of the Falkland and Shetland
islands and east of Iceland. Most of the northern bottlenose whales are
confined to the region where the Gulf Stream waters mix with Arctic
currents, 1.е., the most productive regions. They transgress only into
those cold waters which fall under the influence of the Gulf Stream.
Autumn migrations have not been traced; the whales depart from the
northern regions commencing probably September and extending up to
late autumn. Wintering sites have not been established; it would appear
that they overwinter in the southern part of their range.

Many instances of beached bottlenose whales are known on the
coasts of France, the FRG, the GDR, Holland, Denmark, Sweden, Nor-
way, England, Ireland, Scotland, Faroe and Lofoten islands, and Ice-
land. Beaching has also been recorded on the American coast in Rhode
Island and Massachusetts states and in New York bay and the Gulf of
St. Lawrence. Beached bottlenose whales have even been found on the
coasts of Greenland and Spitsbergen. |

Finds of bottlenose whales in the White and Baltic seas can be
regarded as transgressions (Tomilin, 1957, 1962).

Reproduction. The period of mating has not been established; gesta-
tion extends for 12 and probably even for 15 months. Births occur in the
spring, evidently in March-April or in April-May. The sites of mating
and parturition have not been identified.

Growth and development. Data on these aspects are extremely frag-
mentary and few. The largest of the embryos studied was 305 and even
350 cm long. Apparently the newborn calf ranges from 200 to 250 cm in
length, although the smallest of those measured was only 183 cm long.
During lactation the calves double in length (duration of lactation is pre-
sumably five to seven months). The length of calves fed on milk varied
from 244 to 488 cm. By the third year these whales aitain a length of
6 т (Tomilin, 1957). Females become sexually mature at a body length

659

891

of 7 m. The smallest (of measured) pregnant females was 6.4 т long
(Jonsgard, 1952*).

Enemies, diseases, parasites, mortality, and competitors. The killer
whale is regarded as an enemy of bottlenose whales but there are no
direct observations of killer whale attacks on them. Diseases have not
been studied; some cases of bone tumors have been recorded in museum
skeletons.

The whale louse Platicyamus thompsoni Gosse, mainly infesting the
head, “beak,” and mouth corners, is a known ectoparasite of the northern
bottlenose whale. Among the barnacles, Conchoderma cuviert is more
commonly encountered and C. auritum rarely, in the teeth and light-
colored sections of the skin. Penella crassicornis penetrates the skin.

Seven species of helminths have been registered. The cestode Sirobil-
icephalus triangularis Diesing, parasitizing the intestines, has been found
in the northern bottlenose whale and three species of baleen whales
from the waters of Portugal and Brazil; Diphyllobothrium delphini Bosk
(larvae) has been detected in the adipose depots of these same whale
species. The nematode Anisakis simplex Rudolphi, localizing in the gul-
let, stomach, and intestines of the northern bottlenose whale, is very
widely distributed among marine mammals. It has been found in many
species of baleen and toothed whales and, among pinnipeds, in Steller’s
sea lion caught in the North Sea, off the coasts of Kamchatka, and in
the waters of Japan and New Zealand. Another nematode, Crassicauda
benneti Spaul, has been cited for “bottlenose whale” without mention
of the species. Of the two species of acanthocephalans, Bolbosoma tur-
binella Diesing parasitizes in addition to the northern bottlenose whale,
the intestine of six species of baleen whales from the waters of Iceland
and Australia. Finally, Bolbosoma turbinella [sic.| Diesing, found in the
intestine of the northern bottlenose whale, has been detected in five
species of baleen whales.

The mortality of bottlenose whales has not been established (Dela-
mure, 1955; Tomilin, 1957).

Field characteristics. Long, beak-shaped snout, steeply dropping high
forehead, and the white color of old males are prominent features. One
pair of teeth on the anterior tip of the lower jaw projects outward when
the mouth is closed (sometimes a second pair projects slightly among
very old males). The upper side of the body of young animals is black
or grayish-black and of old animals light brown. When diving deep, this
species flexes the body strongly but never displays the caudal flukes.
Between two prolonged submergences, it remains for a long time on
the water surface, producing 20-30 blows during this period. The blow

892

is low, bushy, 50-60 cm high, and bearly perceptible. The respiratory
sound is short and interrupted with a metallic timbre. (V.A.)

Economic Importance

Commencing from the 1880s, regular hunting of the northern bottlenose
whale was practiced in Norway but the output was low. Hunting inten-
sified due to a drop in catch of right whales and several hundreds of
bottlenose whales were caught in some years. However, by the begin-
ning of the twentieth century, as a result of the rapid development of
the hunting of large fin whales and reduction in the population of bot-
tlenose whales, the catch of the latter decreased sharply. Hunting was
initially carried out predominantly along the coasts of Greenland but
later most of the whales were caught in Norwegian waters (Table 67).

Commencing in 1929, modern specialized hunting of small whales
(including the bottlenose whale) employed small motor boats equipped
with small-caliber (50-60 mm) harpoon guns. Among the small whales
caught, the Minke whale occupied first place followed by the bottlenose
whale. In the 1930s, this hunting, especially of the Minke whale, was of
great economic importance to the Norwegian fishermen.

The maximum number of bottlenose whales was caught in the water
body between Jan Mayen, Iceland, and Faroe Islands, and later west of
Medvezhii Island and Spitsbergen and in the deepwater regions off the
Norwegian coast.

Table 67. Catch of northern bottlenose whales in Norway (international whaling

statistics)
Year Catch of whales Year Catch of whales
1938 70 1954 70
1939 45 1955 124
1940 8 1956 267
1941 21 1957 163
1942 9 1958 145
1943 34 1959 94 \
1944 40 1960 193
1945 22 1961 87
1946 22 1962 321
1947 108 1963 267
1948 59 1964 307
1949 220 1965 692
1950 48 1966 340
1951 7 1967 264
1952 17 1968 384

1953 49

660

893

A single northern bottlenose whale yields about a ton of fat, on
average (large animals up to two tons), in addition to 200 kg of sper-
maceti. The fat is used only for commercial purposes. The spermaceti of
the bottlenose whale is similar to that of the sperm whale in chemical
composition. The hollow of the lower jaw of the bottlenose whale con-
tains a small amount of oil with a typical chemical composition; it does
not congeal in the cold and can be used for lubricating fine mechanisms.
The raw meat causes disorders of the digestive tract but this adverse
phenomenon disappears with cooking and the meat is suitable for feed-
ing dogs and fur-bearing animals. It is not used for human consumption.
This whale is not hunted in the Soviet Union. (V.A.)

W
re

A)
Cato

661 - 684

LITERATURE CITED’

The references listed below include only works cited in the text. Works
cited under the synonymy of various species and groups have not been
included.

Russian Language Sources

Akimushkin, 1.1. 1954а. Osnovnoi ob”ekt риашуа butylkonosa
(Hyperoodon rostratus Muller) [Main food objects of the bottlenose
whale (Hyperoodon rostratus Muller)]. Dokl. AN SSSR, vol. 96, no. 2.

Akimushkin, ILI. 19546. Golovonogie mollyuski у pitanii kashalota
(Cephalopods in the food of the sperm whale). Dokl. AN SSSR,
vol. 96, no. 3.

Akimushkin, 1.1. 1955. О kharaktere pitaniya kashalota (Feeding
behavior of the sperm whale). Dokl. AN SSSR, vol. 101, no. 6.
Akinfov, I. 1848a. Otpiska yakutskogo voevody Ivana Akinfova o
tunguzakh, zhivushchikh bliz Okhoty reki, io morzhovom promysle
po ostrovam i zalivam Okhotskogo morya (Yakut Commander
Ivan Akinfov’s note on Tungus inhabiting near the Okhota River
and walrus hunting off the islands and in the bays of the Sea of
Okhotsk). Dopolneniya К aktam istoricheskim, sobrannye 1 izdannye

Arkheograficheskoyu Komissieyu, vol. 3.

Akinfov, I. 1848b. Rospis’ puti ot r. Okhoty do reki Motykhleya (Descrip-
tion of the course from Okhota River to Motykleya River). Dopol-
петуа К aktam istoricheskim, sobrannye 1 izdannye Arkheografich-
eskoyu Komissieyu, vol. 3.

Al’banov, У. 1917. Na учр, К Zemle Frantsa-losifa (South of Franz Josef
Land). Annexure in Zapisok po gidrografii, vol. 41, no. 2.

Aleev, У. 1913. Otchet о poezdke na morskoi zverinyi promysel у gorlo
Belogo morya v 1911 g. (An account of the 1911 expedition for
hunting marine animals in the neck of the White Sea). Mater. k.
poznan. russk. ryb-va, vol. 2, no. 2.

* Some references are incomplete in the Russian original.

896

Alekseev, Е.У. 1966. Vozrastnye izmeneniya nekotorykh morfologich-
eskikh pokazatelei belomorskogo lysuna (Pagophoca groenlandica
Erxleben) [Age-related changes of some morphological indices
of harp seal (Pagophoca groenlandica Erxleben)]. In: Ekologo-
morfologicheskie issledovaniya nekotorykh zhivotnykh. Kiev.

Ambroz, А.1. 1931. Nekotorye nablyudeniya nad morskimi mlekopi-
tayushchimi Vostochnogo Sakhalina (Some observations on marine
mammals of eastern Sakhalin). Rybn. Khoz. DV, nos. 3 -4.

Anbinder, E.M. 1969. Kariotipy baikal’skogo i kaspiiskogo tyulenei v
svyazi $ problemoi ikh proiskhozhdeniya (Karyotypes of Baikal and
Caspian seals in the context of the problem of their origin). IV Vses.
Soveshch. po Izuch. Morsk. Mlekop. (Tez. Dokl.). Moscow.

Anbinder, E.M. 1971. Khromosomnye nabory grenlandskogo, kaspi-
iskogo i baikal’skogo tyulenei 1 nekotorye voprosy evolyutsii nastoy-
ashchikh tyulenei (Chromosome numbers of Greenland, Caspian,
and Baikal seals and some aspects of the evolution of true seals).
Tsitologtya, vol. 3, no. 3.

Andriyashev, A.P. 1954. Ryby severnykh morei SSSR (Fishes of the
northern seas of the USSR). Opredelitel’ po Faune SSSR, 53, Zool.
In-t AN SSSR. Moscow-Leningrad.

Antipin, V.M. 1938. Morskoi zver’ na Novoi Zemle (Marine life on
Novaya Zemlya). Sov. Arktika, no. 7.

Antipin, V.M. 1938. Fauna pozvonochnykh severo-vostoka Novoi Zemle
(Vertebrate fauna of northeastern Novaya Zemlya). Probl. Arktiki,
no. 2.

Anufriev, [.Р. 1918. О tyulen’em promysle (Seal hunting). Tr. Sev. Коти.
dlya Pomoshchi Pomoram Rus. Sev.

Arsen’ev, V.A. 1935a. Morskoi promysel mlekopitayushchikh v Chukot-
skom i Vostochno-Sibirskom moryakh (Hunting for mammals in the
Chukchi and East Siberian seas). Sov. Sever., nos. 3-4.

Arsen’ev, У.А. 1935b. Nekotorye dannye о belukhe (Delphinapterus leu-
cas Pall.) i ee promysle na Sakhaline [Some data on beluga (Delphi-
napterus leucas Pall.) and its hunting in Sakhalin]. Rybn. Khoz. DV,
no. 13.

Arsen’ev, У.А. 1936a. Morfologicheskaya kharakteristika dal’nevosto-
chnoi belukhi (Morphological characteristics of Far East beluga).
Vestn. DV Filiala AN SSSR, no. 19.

Arsen’ev, V.A. 1936b. Promysel belukhi v Udskoi gube (Beluga hunting
in Uda Gulf). Sov. Kraevedenie, no. 7.

Arsen’ev, V.A. 1937a. O migratsii belukhi v Sakhalinskom raione (Beluga
migrations in Sakhalin region). Izv. TINRO, vol. 10.

897

Агзеп’еу, У.А. 19375. Nekotorye dannye о pitanii belukhi (Some data
on feeding of beluga). Izv. TINRO, vol. 10.

Arsen’ev, V.A. 1939. Raspredelenie 1 migratsii belukhi na Dal’nem Vos-
toke (Distribution and migration of beluga in the Far East). Izv.
TINRO, vol. 15.

Arsen’ev, V.A. 1940. Tekhnika promysla dal’nevostochnoi belukhi (Tech-
nique of hunting Far East beluga). /zv. TINRO, vol. 20.

Arsen’ev, V.A. 1941. Pitanie polosatogo tyulenya (Histriophoca fasciata)
[Feeding of ribbon seal (Histriophoca fasciata)|. Izv. TINRO, vol. 20.

Arsen’ev, V.A. 1958. Nablyudeniya nad morskimi mlekopitayushchimi i
ptitsami (Observations on marine mammals and birds). 7r. Kompl.
Antark. Eksp. AN SSSR. Opisanie Eksp. na D/E “Ob’” 1955 - 1956 gg.
Izd-vo AN SSSR, Moscow.

Arsen’ev, V.A. 1961a. Rasprostranenie kitov v Beringovom more i voz-
mozhnosti razvitiya kitoboinogo promysla [Distribution of whales
in the Bering Sea and the possibilities of developing the whaling
industry]. AN SSSR, Ikhtiol. Komis. “Tr. Soveshch.”’, по. 12.

Arsen’ev, V.A. 1961b. O malykh polosatikakh (Balaenoptera acutoros-
trata) [Minke whale (Balaenoptera acutorostrata)]|. AN SSSR, Ikhtiol.
Komis. “Tr. Soveshch.”’, по. 12.

Arsen’ev, V.A. 1964. O smeshivanii populyatsii morskikh kotikov (Inter-
mixing of fur seal populations). Izv. TINRO, vol. 54; Tr. VNIRO,
vol. 51.

Arsen’ev, V.A. 1968. Programma i metodika issledovanii po morskim
kotikam (Program and methodology of research on fur seals). Ту.
VNIRO, vol. 68; Izv. TINRO, vol. 62.

Arsen’ev, V.A. 1968b. Metodika opredeleniya promyslovogo ispol’zo-
vaniya pokolenii morskikh kotikov (Method for determining the
commercial exploitation of different generations of fur seals). Tr.
VNIRO, vol. 68; Izv. TINRO, vol. 62.

Arsen’ev, V.A. 1971. O lokal’nom raspredelenii morskikh kotikov v
Yaponskom more (Local distribution of fur seals in the Sea of
Japan). Tr. Atl. NIRO, no. 39.

Arsen’ev, У.А. 1972. Vozrastno-polovoi sostav morskikh kotikov,
zimuyushchikh v zapadnoi chasti Tikhogo okeana (Age and sex
composition of fur seals wintering in the western Pacific Ocean).
Tr. VNIRO, vol. 82; Izv. TINRO, vol. 80.

Arsen’ev, V.A., V.A. Zemskii, and I.S. Studenetskaya. 1973. Morskie
mlekopitayushchie (uch. pos.) [Marine Mammals (A Guide for Stu-
dents)]. Pishchevaya Promyshlennost’. Moscow.

Arsen’ev, V.A. and S.G. Fedorov (eds.). 1964. Dokl. Komissii po kotikam
sev. chasti Tikh. okeana ob issled. za 1958-1961 gg. (Report of

898

the Commission on Fur Seals Research in 1958-1961 in the North
Pacific Ocean). Tokyo.

Arsen’ev, У.К. 1927. Tikhookeanskii morzh (Pacific walruses). Knizhnoe
Delo. Khabarovsk- Vladivostok.

Ass, M.Ya. 1934. Ektoparazity morzha (Ectoparasites of walrus). Tr.
Arkt. In-ta, vol. 9.

Ass, M.Ya. 1935. Ektoparazity baikal’skogo tyulenya (Ectoparasites of
Baikal seal). Tr. Baik. Limnol. St., vol. 6.

Averin, Yu.V. 1948. Nazemnya pozvonochnya Vostochnoi Kamchatki
(Land vertebrates of eastern Kamchatka). Tr. Kronotsk. Gos.
Zapovedn., no. 1.

Averkov, LI. 1914. Tyulenii promysel у Kaspiisko-Volzhskom raione
(Seal hunting in the Caspian-Volga region). Vest. Rybopromyshl,
St. Petersburg, nos. 6-7.

Badamshin, В.Г. 1948. Kaspiiskii tyulen’ (Caspian seal). Rybn. Khoz.,
no. 3.

Badamshin, B.I. 1948. О pitanii kaspiiskogo tyulenya (Feeding of Caspian
seal). Tr. Volgo-Kasp. Nauch. Rybokhoz. St., vol. 10.

Badamshin, B.I. 1949. Nekotorye novye dannye о biologii kaspiiskogo
tyulenya у I’dakh (Some new data on the biology of Caspian seal on
ice). Rybn. Khoz., no. 3. iM,

Badamshin, B.I. 1950. Nekotorye dannye ob ostrovnykh zalezhkakh tyu-
lenya v Severnom Kaspii (Some data on island breeding grounds of
seals in the northern Caspian). 7’. Kasp. Bass. Fil. VNIRO, vol. 11.

Badamshin, B.I. 1950. Opyt lecheniya kaspiiskogo tyulenya (Experience
in treating a Caspian seal). Priroda, no. 7.

Badamshin, В.Г. 1959. Ору! soderzhaniya kaspiiskogo tyulenya у nevole
(Experience in raising a Caspian seal in captivity). Tr. Kasp. NII
Morsk. Rybn. Khoz. i Okeanogr., vol. 15.

Badamshin, B.I. 1959. Kaspiiskii tyulen’ pod Stalingradom (Caspian seal
near Stalingrad). Prikaspiiskaya Kommuna, по. 230 (7405).

Badamshin, B.I. 1960. Sostoyanie zapasov kaspiiskogo tyulenya (Status
of Caspian seal reserves). Zool. Zhurn., vol. 39, no. 6.

Badamshin, В.Г. 1961. Zapasy kaspiiskogo tyulenya 1 рий ikh
ratsional’nogo ispol’zovaniya (Caspian seal reserves and methods for
their rational exploitation). Tr. Soveshch. Ikhtiol. Komis. AN SSSR,
no. 12.

Badamshin, B.I. 1964. Lin’ka kaspiiskogo tyulenya (Molting of Caspian
seal). Sb. Morsk. Mlekopit. Moscow.

Badamshin, В.1. 1966. Biologiya i promysel kaspiiskogo tyulenya (Biology
and hunting of Caspian seal). Kazakh. NII Rybn. Khoz., no. 5.

899

Badamshin, B.I. 1966. Vozrastnoi sostav produtsiruyushchikh samok
kaspiiskogo tyulenya kak pokazatel’ sostoyaniya ego zapasov (Age
composition of productive female Caspian seals as an index of the
state of reserves). Tr. Kasp. МП Rybn. Khoz., vol. 22.

Badamshin, B.I. 1966. Kaspiiskii tyulen’ 1 ego promyslovoe ispol’zovanie
(Caspian seal and its commercial exploitation). Sb. Biol. Osnovy
Rybn. Khoz. у Vodoemakh Sr. Аги 1 Kazakh. Alma Ata.

Badamshin, В.1. 1969. Godovoi tsikl zhizni kaspiiskogo tyulenya (Annual
life cycle of Caspian seal). IV Vses. Soveshch. po Izuch. Morsk.
Mlekop. (Tez. Dokl.). Moscow.

Badamshin, B.I. 1969. Chislennost’ i promyslovye zapasy kaspiiskogo
tyulenya (Population and exploitable reserves of Caspian seal). Sb.
Morsk. Mlekopit. Moscow.

Barabash-Nikiforov, [.1. 1936. Lastonogie Komandorskikh ostrovov (Pin-
nipeds of Commander Islands). Tr. ИМКО, vol. 3.

Barabash-Nikiforov, LI. 1940. Fauna kitoobraznykh Chernogo morya,
ее sostav i proiskhozhdenie (Cetacean Fauna of the Black Sea, Its
Composition and Origin). Voronezh.

Barabash-Nikiforov, I.I. 1960. Razmery i okraska del’finov-afalin kak kri-
теги у voprose ikh podvidovoi differentsiatsii (Dimensions and color
of bottlenose dolphins as criteria of their subspecific differentiation).
Nauch. Dokl. Vyssh. Shk., Biol. Nauki, no. 1.

Belkin, A.N. 1964. Novyi vid tyulenya s Kuril’skikh ostrovov (A new
species of seal from the Kuril Islands). Dokl. AN SSSR, vol. 158, no. 5.

Belkin, А.М. 1963. Ob utrobnoi lin’ke morskogo kotika (Callorhinus ursi-
nus L.) [Uterine molting of northern fur seal (Callorhinus ursinus
L.)]. Sb. D/V Filiala Sib. Otd. AN SSSR, no. 17.

Belkin, A.N. 1965. Sovremennoe sostoyanie lezhbishch i chislennost’
morskikh kotikov i sivuchei na ostrovakh Lovushki i Sredneva
(Present status of rookeries and population of fur seals and Steller’s
sea lions on Lovushka and Srednev Islands). Sb. Morsk. Mlekop.
Moscow.

Belkin, A.N. 1966a. Letnee raspredelenie, zapasy, perspektivy promysla
i nekotorye cherty biologii sivucha, obitayushchego na Kuril’skikh
ostrovakh (Summer distribution, reserves, hunting prospects, and
some features of the biology of Steller’s sea lion inhabiting Kuril
Islands). Izv. TINRO, vol. 58.

Belkin, A.N. 1966b. O vzaimnykh otnosheniyakh sivucha i kotika na
sovmeéstnykh lezhbishchakh razmnozheniya (Interrelations between
Steller’s sea lion and fur seal in common breeding colonies). Izv.
TINRO, vol. 58.

900

Belkin, A.N., С.М. Kosygin, and K.I. Panin. 1969. Novye materialy po
kharakteristike ostrovnogo tyulenya (New data on the characteristics
of island seal). Sb. Morsk. Mlekop. Moscow.

Bel’kovich, V.M. 1959. O mekhanizme smeny okraski belukhi (Color
variation mechanism in belugas). DAN SSSR, vol. 127, no. 4.

Bel’kovich, V.M. 1960. Nekotorye biologicheskie nablyudeniya za
belukhoi s samoleta (Some biological observations of beluga from
an airplane). Zool. Zhurn., vol. 39, no. 9.

Belkovich, V.M. 1964. Stroenia kozhnogo pokrova nekotorykh
lastonogikh (Structure of the integument of some pinnipeds).
Morfologicheskie Osobennosti Vodnykh Mlekop. Moscow.

Bel’kovich, V.M. and R.Sh. Khuzin. 1960. Atlanticheskii morzh (Atlantic
walrus). Priroda, no. 2.

Bel’kovich, V.M. and R.Sh. Khuzin. 1960. Opyt ispol’zovaniya samoleta
dlya razvedki belukhi (Experience in using an airplane for scouting
belugas). Tr. PINRO, по. 12.

Bel’kovich, V.M. and A.V. Yablokov. 1960. Osobennosti stroeniya zub-
noi sistemy morzha (Structural features of dentition in walrus). ГУ
Molodezhnaya Nauch. Konf. In-ta Morfol. Zhiv. AN SSSR. Moscow.

Bel’kovich, V.M. and A.V. Yablokov. 1961. Sredi morzhei (Among wal-
ruses). Priroda, no. 3.

Bel’kovich, V.M. and V.S. Gurevich. 1971. Voprosy otlova 1 dlitel’nogo
soderzhaniya del’finov v nevole (Aspects of trapping and prolonged
rearing of dolphins in captivity). Tr. Atlant NIRO, no. 39.

Beloborodov, A.G. 1969. Ob osenne-zimnikh migratsiyakh grenland-
skogo tyulenya k Belomy moryu (Autumn-winter migrations of harp
seal to the White Sea). In: 8-ya Sessiya Uch. Sov. po Probl. “Biol.
Res. Bel. Morya i Vnutr. Vod Катеш”’. Petrozavodsk.

Beloborodov, А.С. and V.A. Potelov. 1965. Rezul’taty izucheniya tyu-
lenei Grenlandskogo morya v 1965 g. (Results of studies on seals
of the Greenland Sea in 1965). “Mater. Rybokhoz. Issl. Sev. Bass.”
PINRO, no. 9.

Beloborodov, А.С. and V.A. Potelov. 1968. О detnom periode tyulenei
v Belom more (Juvenile period of seals in the White Sea). In: 7-
ya Sessiya Uch. Sov. po Probl. “Biol. Res. Bel. Morya i Vnutr. Vod
Karelii”’. Petrozavodsk.

Beloborodov, А.С. and V.A. Potelov. 1969. Nekotorye osobennosti
nachal’nogo perioda zhizni detenyshei khokhlacha i kharakteristika
ikh volosyanogo pokrova (Some features of the juvenile period of
hooded seal calves and the characteristics of their hair coat). Sb.
Morsk. Mlekop. Moscow.

901

Belopol’skaya, М. М. 1960. Gel’mintofauna largi (Phoca vitulina largha
Pall.) [Helminth fauna of larga seal (Phoca vitulina largha Pall.)].
Vest. Leningr. Un-ta, no. 3.

Belopol’skii, L.O. 1939. О migratsiyakh i ekologii razmnozheniya
tikhookeanskogo morzha (Odobaenus rosmarus divergens Г. Ilig.)
[Migrations and ecology of reproduction of the Pacific walrus
(Odobaenus rosmarus divergens |. Шв.)|. Zool. Zhurn., vol. 18, no. 5.

Belopol’skii, L.O. 1941. Shchennye zalezhki tevyaka Halichoerus grypus
Fabr. у zapovednike “Sem’ ostrovov” (Nurseries of Atlantic seal,
Halichoerus grypus Fabr., in “Зет” Ostrovov” sanctuary). Ти. Gos.
Zap. “Sem’ Ostrovov’’, no. 1.

Belopol’skii, L.O. 1951. Zapovednik “Зет?” ostrovov” (“Sem’ Ostrovov”
sanctuary). Sb. Zapovedniki SSSR, vol. 1. Moscow.

Berzin, A.A. 1959. O pitanii kashalota v Beringovom more (Feeding of
sperm whale in the Bering Sea). Izv. TINRO, vol. 47.

Berzin, A.A. 1961. Materialy po razvitiyu zubov i opredeleniyu vozrasta
kashalotov (Data on dental growth and age determination of sperm
whales). Tr. Soveshch. Ikhtiol. Komis. AN SSSR, no. 12.

Berzin, А.А. 1964a. Rost Kashalotov severnoi chasti Tikhogo okeana
(Growth of sperm whales in the North Pacific Ocean). Tr. VNIRO,
vol. 53; Izv. TINRO, vol. 52.

Berzin, A.A. 1964b. Opredelenie vozrastnogo sostava stada kashalotov
Beringova morya i prilezhashchikh chastei Tikhogo okeana (Deter-
mining the age composition of sperm whale herds in the Bering Sea
and adjoining parts of the Pacific Ocean). Tr. VNIRO, vol. 53; Izv.
TINRO, vol. 52.

Berzin, A.A. 1965. Opredelenie vozrasta nastupleniya polovoi zrelosti
samtsov kashalota severnoi chasti Tikhogo okeana (Determining the
age of onset of sexual maturity among sperm whales of the North
Pacific Ocean). Sb. Morsk. Mlekop. Moscow.

Berzin, A.A. 1971. Kashalot (Sperm Whale). Pishchevaya Promyshlen-
nost’. Moscow.

Betesheva, E.I. 1954. Dannye о pitanii usatykh kitov у raione Kuril’skoi
gryady (Data on the food of baleen whales in the region of the Kuril
range). Tr. In-ta Okean. AN SSSR, vol. П.

Betesheva, Е.1. 1955. Pitanie usatykh kitov у raione vod Kuril’skoi gryady
(Feeding of baleen whales in the region of the Kuril range). Tr. In-ta
Okean. AN SSSR, vol. 18.

Betesheva, E.I. 1960. Pitanie kashalota (Physeter catodon L.) 1 berardiusa
(Berardius Батай Stejneger) у raione Kuril’skoi gryady [Feeding
of sperm whale (Physeter catodon Г.) and Baird’s beaked whale

902

(Berardius Батаи Stejneger) in’ the region of the Kuril range]. 7r.
Vses. Gidrobiol. Ob-va, vol. 10.

Betesheva, E.I. 1961. Pitanie promyslovykh kitov prikuril’skogo raiona
(Feeding of game whales in the Kuril region). Tr. In-ta Morf. Zhiv.
AN SSSR, no. 34.

Betesheva, E.I. and 1.1. Akimushkin. 1955. Pitanie kashalota у raione vod
Kuril’skoi gryady (Feeding of sperm whale in the region of the Kuril
range). Tr. In-ta Okean. AN SSSR, vol. 18.

Bianki, V.V. 1965. O chislennosti morskikh mlekopitayushchikh v ver-
shine Kandalakshskogo zaliva (Population of marine mammals at
the head of Kandalakshsk Вау). “Tr. V Sess. Uch. 5оу.” GosKIOSKh,
Petrozavodsk.

Bianki, V.V. and V.N. Karpovich. 1968. O vliyanii anomal’noi ledovi-
tosti 1968 g. na ptits i mlekopitayushchikh Belogo morya i Murmana
(Effect of anomalous ice conditions in 1968 on birds and mammals
of the White and Murman seas). In 7-ya Sesstya Uch. Sov. po Probl.
“Biol. Res. Bel. Morya i Vnutr. Гоа Karelii (Tez. Dokl.)”. Petrozavodsk.

Birman, Т.В. 1966. Novye dannye о rasprostranenii 1 migratsiyakh
morskikh kotikov (New data on the distribution and migrations of
fur seals). Vopr. Geogr. Kamchatki, no. 4.

Bobrinskii, N.A., B.A. Kuznetsov, and A.P. Kuzyakin. 1965. Opredeli-
tel’ mlekopitayushchikh SSSR (Key to Mammals of the USSR).
Prosveshchenie, Moscow.

Bodrov, V.A., S.N. Grigor’ev, and У.А. Tver’yanovich. 1958. Tekhnikai .
tekhnologiya obrabotki morskikh mledopitayushchikh (kity, del’finy,
lastonogie) [Technique and Technology of Processing Marine
Mammals (Whales, Dolphins, and Pinnipeds)]. Pishchepromizdat,
Moscow.

Boguslav, I. 1847. Vzglyad na Belomorskie zverinye promysly (White Sea
animal hunting). Tr. Imp. Vol’no-ekonomich, Ob-va, vol. 4, no. 1.
Boitsov, L.V. 1934. Kotikovoe khozyaistvo (Seal Fisheries). Wneshnet-

orgizdat.

Borisov, V.I. 1966. Nekotorye dannye serologicheskogo analiza stad gren-
landskogo tyulenya (Pagophilus groenlandicus Erxb.) [Some data on
serological analysis of Greenland harp seal (Pagophilus groenlandicus
Erxb.) herds]. Zool. Zhurn., vol. 45, no. 12.

Brandt, I.F. 1856. Pozvonochnye zhivotnye Severoevropeiskoi Rossii i
osobenno Severnogo Urala (Vertebrates of North European Russia,
Especially of the Northern Urals). St. Petersburg.

Breitfus, L.L. 1903. Ekspeditsiya dlya nauchno-promyslovykh issledovanii
и beregov Murmana. Otchet о ee deyatel’nosti $ 1 yanvarya po 1
iyunya 1902 g. (Scientific-hunting Research Expedition on Murman

903

Coasts. An Account of Its Activity from January 1 through June 1,
1902). St. Petersburg.

Braitfus, L.L. 1906. Ekspeditsiya dlya nauchno-promyslovykh issledovanii
и beregov Murmana. Otchet о ee rabotakh у 1903 g. (Scientific-
hunting Research Expedition on Murman Coasts. An Account of Its
Activity in 1903). St. Petersburg.

Breitfus, L.L. 1908. Ekspeditsiya Фуа nauchno-promyslobykh issle-
dovanii и beregov Murmana. Otchet о rabotakh у 1904 р. (Scientific-
hunting Research Expedition on Murman Coasts. An Account of
Activity in 1904). St. Petersburg.

Breitfus, L.L. 1912. Otchet nachal’nika ekspeditsii (Report by the expe-
dition leader). Tr. Murmanskoi Nauch.-Promysl. Ekspeditsii 1905 р.
St. Petersburg.

Breitfus, L.L. 1915. Trudy Murmanskoi nauchno-promyslovoi ekspeditsii
1906 g. (Proceedings of the Murman Scientific-hunting Expedition
of 1906). St. Petersburg.

Budylenko, G.A. and Yu.A. Mikhalev. 1970. K biologii razmnozheniya
seivala Antarktiki (Reproduction biology of Antarctic sei whale). 7r.
AtlantNIRO, no. 25.

Bukin, V.N. and E.P. Skorobogatova. 1951. Pechen’ kitov kak syr’e dlya
polucheniya vitamina A (Whale liver as raw material for producing
vitamin А). Sb. Vitaminnye Resursy i Ikh Ispol’zovanie, no. 1. Izd-vo
AN SSSR, Moscow.

Bulgakov, М.У. et al. 1954. Киоуое kozhevennoe syr’e i ego ispol’zovanie
у kozhevenno-obuvnoi promyshlennosti (Whale skin and its use in
the leather-shoe industry). Tr. Tsentr. NII Kozhev.-obuvnoi Prom-ti,
Nauchno-Issl. Trudy, по. 22.

Butorin, О.А. 1957. Razvivat’ promysel belukhi (Developing beluga
hunting). Rybn. Khoz., no. 2.

Buturlin, S.A. 1913. Nablyudeniya nad mlekopitayushchimi, sdelannye vo
vremya Kolymskoi ekspeditsii 1905 g. (Observations on mammals
during the 1905 Kolyma expedition). Dnevnik Zool. Otdel. Ob-va
Lyubit. Est., Antrop. i Etnogr., Nov. Ser., vol. 1, no. 5.

Bychkov, V.A. 1960. Novye dannye о lin’ke lakhtaka (New data on molt-
ing of bearded seals). Inf. Sb. VNIRO, no. 8.

Bychkov, V.A. 1964a. Materialy po lin’ke morskikh kotikov na ostrove
Tyulen’em (Data on the molting of fur seals on Tyulen’ Island). Izv.
TINRO, vol. 54; Tr. VNIRO, vol. 51.

Bychkov, V.A. 1964b. Nablyudeniya za samkami morskogo kotika na
ostrove Tyulen’em (Observations on female fur seals on Tyulen’
Island). Izv. TINRO, vol. 54; Tr. VNIRO, vol. 51.

904

Bychkoy, V.A. 1964c. Promysel morskikh kotikov па ostrove Tyulen’em
(Hunting of fur seals on Tyulen’ Island). Izv. TINRO, vol. 54; Tr.
VNIRO, vol. 51. ;

Bychkov, V.A. 1968. Kharakteristika razmerov i okraski volosyanogo
pokrova morskikh kotikov (Size characteristics and color of hair
coat of fur seals). Tr. VNIRO, vol. 68; Izv. TINRO, vol. 62.

Bychkov, V.A. 1969. Sostav i razmeshchenie samok morskikh kotikov na
garemnom lezhbishche ostrova Tyulen’ego (Composition and dispo-
sition of female fur seals in the harem rookeries on Tyulen’ Island).
Sb. Morsk. Mlekop. Nauka, Moscow.

Bychkov, V.A. and S.V. Dorofeev. 1962. O povedenii sekachei morskikh
kotikov v garemnyi period ikh zhizni (Behavior of bull fur seals in
their harem period). Zool. Zhurn., vol. 41, no. 9.

Chapskii, К.К. 1930. О sbore materiala po tyulenyu na dagestanskom
poberezh’e Kaspiya (Collection of seal material on the Dagestan coast
of the Caspian). Soveshch. ро 1551. Kasp. Туш. 1 Ego Prom. Astrakhan.

Chapskii, К.К. 1931. Ladozhskii tyulen’ 1 vozmozhnost’ ego promysla
(Ladoga seal and its possible hunting). Izv. Leningr. N.-I. Ikhtiol.
In-ta, vol. 13, no. 2.

Chapskii, K.K. 1931. O kaspiiskom tyulene “plyvune” (Caspian seal
“plyvun”’). Izv. Len. [khtiol. In-ta, vol. 13, no. 2.

Chapskii, К.К. 1936. Morzh Karskogo morya (Kara Sea walrus). Tr. Arkt.
In-ta, vol. 67.

Chapskii, К.К. 1937. “Belukha” ili “beluga” (“Belukha” or “beluga’’).
Probl. Arktiki, no. 4.

Chapskii, К.К. 1937. Issledovaniya belomorskogo stada grenlandskogo
tyulenya 1937 g. (Study of White Sea harp seal herds in 1937). Probl.
Arktiki, no. 2.

Chapskii, К.К. 1937. Migratsii i promysel belugi (De/phinapterus leucas
Pall.) v severnoi chasti Obskoi guby [Migrations and hunting of bel-
uga (Delphinapterus leucas Pall.) in the northern part of Obsk Gulf].
Tr. Arkt. In-ta, vol. 71.

Chapskii, К.К. 1938. Morskoi zayats (Erignathus barbatus Fabr.)
Karskogo i Barentseva morei [Bearded seal (Erignathus barbatus
Fabr.) of the Kara and Barents seas]. Tr. Arkt. In-ta, vol. 123.

Chapskii, К.К. 1938. Noveishie dannye о respredelenii belomorskoi газу
grenlandskogo tyulenya vne belomorskogo basseina (Latest data on
distribution of local White Sea harp seal outside the White Sea
basin). Probl. Arktiki, no. 4.

Chapskii, К.К. 1939. Kratkii istoricheskii analiz sovremennogo sostoy-
aniya zapasov morzha v Barentsevom i Karskom moryakh (Brief

\

905

Historical Analysis of the Status of Contemporary Walrus Reserves
in the Barents and Kara Seas). Izd-vo Glavsevmorputi, Leningrad.

Chapskii, К.К. 1940. Nerpa zapadnykh morei Sovetskoi Arktiki (Ringed
seals of western Soviet Arctic seas). Tr. Arkt. In-ta, vol. 145.

Chapskil, К.К. 1940. Rasprostranenie morzha у moryakh Laptevykh 1
vostochno-Sibirskom (Walrus distribution in the Lepteva and East
Siberian seas). Probl. Arktiki, no. 3.

Chapskii, K.K. 1941. Morskie zveri Sovetskoi Arktiki (Soviet Arctic
Marine Animals). [24-уо Glavsevmorputi, Moscow-Leningrad.

Chapskii, К.К. 1948. Tyuleni podsemeistva Phocinae 1 ikh khozyaistven-
noe znachenie (Seals of the subfamily Phocinae and their economic
importance). Tez. diss., Zool. In-ta AN SSSR. Leningrad.

Chapskii, K.K. 1949. Mlekopitayushchie vysokikh shirot Severnogo
Ledovitogo okeana (High altitude mammals of the northern Arctic
Ocean). Tr. 3 Ekspeditsii Glavsevmorputi na Ledokole “Sedov’’, pt. 3.

Chapskii, K.K. 1952. Vozrastno-polovaya izmenchivost’ kraniologich-
eskikh priznakov i ee vliyanie na diagnostiku nekotorykh lastonogikh
(Age- and sex-related variation of craniological characteristics and
its influence on recognizing some pinnipeds). Izv. Est.-nauch. In-ta
im. Lesgafta, vol. 25.

Chapskii, К.К. 1955a. Opyt peresmotra sistemy 1 diagnostiki tyulenei
podsemeistva Phocina (Review of the system and diagnosis of seals
of the subfamily Phocinae). Tr. Zool. In-ta AN SSSR, vol. 17.

СрарзКи, К.К. 1955b. К voprosu ob istorii formirovaniya kaspiiskogo
i baikal’skogo tyulenei (Historical evolution of Caspian and Baikal
seals). Tr. Zool. In-ta SSSR, vol. 17.

Chapskii, K.K. 1961. Nekotorye aspekty morfogeneza lastonogikh v
svyazi $ formirovaniem arealov (Some aspects of morphogenesis of
pinnipeds in relation to the formation of their ranges). Tez. Dokl. I
Vses. Soveshch. po Morsk. Mlekop. Moscow.

Chapskii, K.K. 1961. Nekotorye ekologicheskie obosnovaniya sezonnoi
dinamichnosti areala belomorskoi populyatsii grenlandskogo tyu-
lenya (Some ecological features of seasonal dynamics of the range of
the White Sea harp seal population). Tez. I Vses. Soveshch. po Izuch.
Morsk. Mlekop. Moscow.

Chapskii, К.К. 1961. Sovremennoe sostoyanie i problemy sistematiki
lastonogikh (Present status and taxonomic problems of pinnipeds).
Tr. Soveshch. Ikht. Komis. AN SSSR, no. 12.

Chapskii, К.К. 1963. Otryad lastonogikh (Order of pinnipeds). In:
Mlekop. Fauny SSSR, pt. 2. Izd-vo AN SSSR, Moscow-Leningrad.
Chapskii, K.K. 1963. Resursy zveroboinogo promysla, ikh sostoyanie,

ispol’zovanie 1 izuchenie (Animal resources for hunting, their status,

906

exploitation, and study). Tez. Dokl. II Vses. Soveshch. po Izuch. Morsk.
Mlekop. Moscow.

Chapskii, К.К. 1963. K izucheniyu vosproizvoditel’noi sistemy grenland-
skogo tyulenya i intensivnosti razmnozheniya ego belomorskoi pop-
ulyatsii (A study of the reproductive system of harp seal and the repro-
duction intensity of its White Sea population). Tr. PINRO, no. 15.

Chapskii, К.К. 1964. Nachal’nyi period postnatal’nogo rosta_ belo-
morskogo lysuna (Pagophoca groenlandica) [Early postnatal develop-
ment of White Sea harp seal (Pagophoca groenlandica)|. Sb. Morsk.
Mlekop. Nauka, Moscow.

Chapskii, K.K. 1965. Rost 1 nastuplenie polovoi zrelosti и samok kaspi-
iskogo tyulenya (Growth and onset of sexual maturity in female
Caspian seals). Sb. Morsk. Mlekop. Moscow.

Chapskii, K.K. 1966. Sovremennoe sostoyanie 1 zadachi vosstanovleniya
resursov morskogo zveroboinogo promysla (Present status and
problems of restoring the reserves of marine animals for hunting).
Tez. Dokl. III Vses. Soveshch. po Izuch. Morsk. Mlekop. Moscow-
Leningrad.

Chapskii, K.K. 1966. Sovremennye dannye o taksoncmicheskom sostave
tyulenei roda Phoca (Present data on taxonomic composition of seals
of the genus Phoca). Tez. Dokl. Ш Vses. Soveshch. po Izuch. Morsk.
Mlekop. Moscow-Leningrad.

Chapskii, K.K. 1966. Ekologicheskaya raznorodnost’ i taksonomich-
eskaya struktura tyulenei roda Phoca (Ecological heterogeneity and
taxonomic structure of seals of the genus Phoca). Tez. Dokl. Otch.
Nauch. Sessii po Поват Rabot 1965 в. Leningrad.

Chapskii, K.K. 1967. Morfotaksonomicheskaya kharakteristika pagetod-
noi formy largi Beringova morya (Morphotaxonomic characteristics
of the pagetode form of Bering Sea larga seal). Tr. PINRO, no. 21.

Chapskii, К.К. 1967а. О perekhodnykh tipakh okraski volosyanogo
pokrova samtsov grenlandskogo tyulenya (Transitional hair coat
color types of male harp seals). Tr. PINRO, no. 21.

Chapskii, K.K. 1968. Rol’ Polyarnogo basseina v istorii lastonogikh.
Kainozoiskaya istoriya Polyarnogo basseina i ee vliyanie na razvitie
landshaftov severnykh territorii (Role of the Polar Basin in the
history of pinnipeds. Cenozoic history of the Polar Basin and its
influence on the development of northern territories). Mater. k
Simpoz. Leningrad.

Chapskii, K.K. 1969. Voprosy taksonomicheskoi struktury semeistva
Phocidae v svete kraniologicheskikh dannykh (Taxonomic structure
of the family Phocidae in the light of craniological data). Tez. Dokl.
IV Vses. Soveshch. po Izuch. Morsk. Mlekop. Moscow.

907

Chapskii, К.К. 1969. Taksonomiya tyulenei года Phoca sensu stricto у
svete sovremennykh kraniologicheskikh dannykh (Taxonomy of seals
of the genus Phoca sensu stricto in the light of recent craniological
data). Sb. Morsk. Mlekop. Moscow.

Chapskii, К.К. 1970. Kontseptsiya arkticheskogo proiskhozhdeniya
lastonogikh i drugie puti resheniya etoi problemy (Concepts оп
the arctic genesis of pinnipeds and other methods of resolving
this problem). Sb. Sev. Led. Okean i Ego Pober. у Kainozoe.
Gidrometeoizdat, Leningrad.

Chelnokov, Е.С. 1971. Peremeshcheniya morskikh kotikov na
lezhbishchakh ostrova Mednogo (Fur seal movements in rookeries
on Mednyi Island). Tr. AtlantNIRO, no. 39.

Chentsov, P. 1959. Nerpa v tundre (Ringed seal in the tundra). Okhota
т Okhot. Khoz., по. 2.

Chernigovskii, N.T. 1935. Pervaya zimovka na o. Uedineniya (First win-
tering on Uedineniya Island). Byull. Аг. In-ta, по. 10.

Chugunkov, D.I. 1966. O lokal’nosti stad morskikh kotikov, obitayush-
chikh na ostrovakh Beringa i Mednyi (Localization of fur seal herds
on Bering and Mednyi islands). Izv. TINRO, vol. 58.

Chugunkov, D.I. 1967. Opyt soderzhani shchenka largi у nevole (Raising
larga seal calves in captivity). Vopr. Geogr. Kamchatki, no. 5.

Chugunkov, Р.Г. 1968. Ob opredelenii vozrastnogo sostava dobychi
promyslovykh kotikov po vozvratu metok (Determining the age
composition of fur seals caught, based on the tags returned). Ти.
VNIRO, vol. 68; Izv. TINRO, vol. 62.

Chugunkov, D.I. 1968. Sivuchi i kotikovoe khozyaistvo Komandorskikh
ostrovov (Steller’s sea lion and fur seal trade on Commander
Islands). Tr. VNIRO, vol. 68; Izv. TINRO, vol. 62.

Chugunkovy, D.I. 1969. Dal’nevostochnaya larga (Far East larga seal).
Okhota i Okhotn. Khoz., no. 10.

Chugunkov, D.I. 1970. Morzhi na Kamchatka (Walrus on Kamchatka).
Vopr. Geogr. Kamchatki, no. 6.

Chugunkov, О.Г. 1970. Nablyudeniya za pyatnistym tyulenem utkinskogo
lezhbishcha (Observations on common seal of Utkinsk rookery). Jzv.
TINRO, vol. 70.

Chugunkoy, D.I. and V.G. Prokhorov. 1966. Novye svedeniya о zimovke
kotikov v Beringovom more (Recent information on the wintering
of fur seals in the Bering Sea). Izv. TINRO, vol. 58.

Chuzhakina, E.S. 1955. Akiba, kol’chataya nerpa (Akiba—ringed seal).
Tr. In-ta Okeanol. AN SSSR, vol. 14.

Chuzhakina, E.S. 1955. К voprosu о ‘tsikle razmnozheniya kashalota
(Sperm whale reproduction cycle). Tr. In-ta Okeanol. AN SSSR, vol. 18.

908

Danilevskii, М.Уа. 1862. Rybnye i zverinye promysly у Belom i Ledovitom
moryakh (Fish and animal hunting in the White and Arctic seas).
Issl. о Sost. Rybolovstva у Rossii, vol. 4. St. Petersburg.

Delamure, S.L. 1955. Gel’mintofauna morskikh mlekopitayushchikh v
sveta ikh ekology 1 filogenii (Helminth Fauna of Marine Mammals
in Terms of Their Ecology and Phylogenesis). Moscow.

Delamure, S.L. and E.V. Alekseev. 1965. Obzor gel’mintofauny
kol’chatoi nerpy (Phoca hispida Schreber) [Review of helminth fauna
of ringed seal (Phoca hispida Schreber)]. Sb. Рагагиу i Parazitozy
Cheloveka 1 Zhivotnykh. Kiev.

Delamure, S.L., D.D. Zavaleeva, and G.A. Fedoseev. 1965. Materialy
k faune paraziticheskikh chervei kol’chatoi nerpy (Pusa hispida
Krascheninnikovi) [Data on the fauna of parasitic worms of ringed
seal (Pusa hispida Krascheninnikovi)]. Пор. Nauch. Konf. Prof.-prep.
Sostava Krym. Ped. In-ta (Tez. Dokl.). Simferopol’.

Delamure, S.L., $.Z. Dubovyi, and A.S. Skrjabin. 1961. К izucheniyu
filyarii-parazitov morskikh kotikov (A study of the filarial parasites
of fur seals). Izv. Krym. Otd. Geogr. Ob-va SSSR, no. 6.

Delamure, S.L., Yu.V. Kurochkin, and A.S. Skrjabin. 1964. O gel’minto-
fauna kaspiiskogo tyulenya (Helminth fauna of Caspian seal). ТУ.
Astrakh. Zap-ka, no. 9.

Delamure, S.L. and A.S. Skrjabin. 1960. Gel’mintofauna komandorskikh
kotikov (Helminth fauna of Commander Island fur seals). Nauch.
Dokl. Vyssh. Shk., Biol. Nauki, no. 2.

Delamure, S.L. and V.V. Treshchev. 1966. O gel’mintofaune khokhlacha
(Cystophora cristata Erxleben), v Grenlandskom more [Helminth
fauna of hooded seal (Cystophora cristata Erxleben) inhabiting the
Greenland Sea]. Zool Zhurn., vol. 14, no. 12.

Dmokhovskii, A.V. 1933. Materialy k poznaniyu geograficheskogo
rasprostraneniya 1 biologii belukhi (Delphinapterus leucas Pall.) у
Belom more [Data on the geographic distribution and biology of
beluga (Delphinapterus leucas Pall.) in the White Sea]. Byull. MOIP,
vol. 42, no. 1.

Dogel’, V.A. 1958. Parazitofauna i okruzhayushchaya sreda. Nekotorye
voprosy ekologii parazitov presnovodnykh ryb (Parasitic fauna and
environment. Some aspects of the ecology of the parasites of fresh-
water fish). Sb. Osnovnye Problemy Parazitologii Ryb. Leningrad.

Dolzhenkov, V.N. 1969. Zooplankton. kak indikator skoplenii usatykh
kitov (Zooplankton as an indicator of congregations of baleen
whales). Sb. Morsk. Mlekop. Nauka, Moscow.

Dormenko, V.V. 1952. Opyt pressovaniya sala kashalota Фуа
polyucheniya zhira i kozhevennogo syr’ya (Pressing of sperm

909

whale blubber for recovery of oil and raw skin). Ти’ VNIRO,
vol. 22.

Dorofeev, S.V. 1936a. Nablyudeniya nad periodom razmnozheniya gren-
landskogo tyulenya (Histriophoca groenlandica oceanica Lepechin)
[Observations on the reproduction period of harp seal (Histriophoca
groenlandica oceanica Lepechin)]. DAN SSSR, vol. 11, no. 1.

Dorofeev, S.V. 1936b. Materialy po detnomu periodu zhizni grenland-
skogo tyulenya (Data on the juvenile period of harp seal). Tr. Polyar.
Komis., no. 31.

Dorofeev, S.V. 1936. Materialy k promyslovoi biologii lastonogikh v
vesennii ledovyi period v Tatarskom prolive (Data on biology of pin-
nipeds relative to hunting in the autumn icy period in Tatar Strait).
Tr. VNIRO, vol. 3.

Dorofeev, 5.У. 1939a. Sootnoshenie vozrastnykh grupp u tyulenei kak
pokazatel’sostoyaniya zapasov (Ratio of age groups of seals as an
index of state of reserves). Sb. Posvyashch. Nauch. Deyat. Akad.
N.M. Knipovicha. Moscow.

Dorofeev, S.V. 1939b. Vliyanie ledovykh uslovii na povedenie grenland-
skogo tyulenya (Effect of ice conditions on the behavior of harp
seal). Zool. Zhurn., vol. 18, no. 5.

Dorofeev, S.V. 1947. Vliyanie ledovykh uslovii na dislokatsiyu zalezhek
kaspiiskogo tyulenya (Effect of ice conditions on the dislocation of
Caspian seal rookeries). Dokl. VNIRO, no. 9.

Dorofeev, 5.У. 1952. Belomorskii tyulen’-lysun (promyslovaya biol.)
[White Sea Harp Seal (Biological Aspects Related to Hunting)].
Moscow.

Dorofeev, S.V. 1956. Zapasy grenlandskogo tyulenya i ikh ispol’zovanie
(Reserves of harp seal and their exploitation). Rybn. Khoz. Severa,
no, 12.

Dorofeev, S.V. 1958. Biologicheskoe obosnovanie srokov zimnego
promysla kasp. tyulenya (Biological rationale of winter hunting of
Caspian seal). Inf. Sb. VNIRO.

Dorofeev, S.V. 1958. Metodika ucheta chernykh kotikov па ostrove
Tyulen’em (Method of counting the Black Sea fur seals on Tyulen’
Island). Inf. Sb. VNIRO, no. 3.

Dorofeev, S.V. 1960. Materialy k dinamike chislennosti morskikh kotikov
na ostrove Tyulen’em (Data on the population dynamics of fur seals
on Tyulen’ Island). Byull. MOIP, Otd. Biol., vol. 65 (1).

Dorofeev, S.V. 1960. Osnovnye cherty godovogo 15а zhizni
belomorskogo lysuna (Main features of the life cycle of White Sea
harp seal). Sb. Sov. Rybokhoz. Issl. v Moryakh Evr. Severa. Moscow.

910

Dorofeev, S.V. 1964. Severnye morskie kotiki (Callorhinus ursinus Г.)
[Northern fur seals (Callorhinus ursinus L.)]. Izv. TINRO, vol. 54; Tr.
VNIRO, vol. 51.

Dorofeev, S.V. and V.A. Bychkov. 1964. Biologicheskie predposylki dlya
regulirovaniya chislennosti sekachei na ostrove Tyulen’em (Biologi-
cal premises for controlling the bull populations on Tyulen’ Island).
Izv. TINRO, vol. 54; Tr. VNIRO, vol. 51.

Dorofeev, S.V. and S.K. Klumov. 1936a. Promyslovaya kharakteristika
migratsii belukhi v raione Sakhalina (Migration characteristics of
belugas in Sakhalin region). Tr. VNIRO, vol. 3.

Dorofeev, S.V. and S.K. Klumov. 1936b. К voprosu ob opredelenii
vozrasta belukhi i sostava kosyakov (Determining the age and herd
composition of belugas). Tr. VNIRO, vol. 3.

Dorofeev, S.V. and S.Yu. Freiman. 1928. Opyt kolichestvennogo ucheta
zapasov belomorskogo stada grenlandskogo tyulenya metodom
aerofotografirovaniya (Quantitative determination of the reserves of
White Sea harp seal herds by aerial photography). Tr. Nauch. In-ta
Rybn. Khoz., pt. 2, no. 4.

Dorofeev, S.V. and S. Yu. Freiman. 1928. Kaspiiskii tyulen’ 1 ego promy-
sel vo l’dakh (Caspian seal and its hunting on ice). Tr. Nauch. In-ta
Rybn. Khoz-va, vol. 3, no. 3.

Doroshenko, N.V. 1969. O raspredelenii i migratsiyakh gorbatykh kitov
у severo-vostochnoi chasti Tikhogo okeana, Beringovom i Chukot-
skom moryakh (Distribution and migrations of humpback whales
in the northeastern Pacific Ocean, Bering and Chukchi seas). Sb.
Morsk. Mlekop. Moscow.

Dragunov, A.M. and N.E. Kasinova. 1951. Vesovoi i khimicheskii sostav
del’finov Chernogo morya (Quantitative and chemical een set
of Black Sea dolphins). Rybn. Khoz., no. 11.

Dragunov, A.M. and N.E. Kasinova. 1954. Tekhnologicheskaya kharak-
teristika del’fina i skhema ratsional’noi ego obrabotki (Technological
characteristics of dolphin and a method for its rational exploitation).
Tr. VNIRO, vol. 29.

Druzhinin, А.М. 1924. К voprosu о stroenii funktsii 1 genezise perednego
poyasa konechnostei u del’fina (Structure, function, and genesis of
the fore girdle of dolphin limbs). Rus. Zool. Zhurn., vol. 4, nos. 3
and 4.

Druker, С.Е. and S.I. Gakichko. 1936. Tikhookeanskaya belukha
kak promyshlennoe syr’e (Pacific Ocean beluga as an exploitable
resource). Tr. VNIRO, vol. 3.

Dubrovskii, A.N. 1937. O brachnom krike morskogo zaitsa (Mating call
of bearded seal). Priroda, no. 4.

911

Dukhovnyi, М.М. 1933. Belukha у Obskoi gube (Delphinapterus leucas
Pall.) [Beluga (Delphinapterus leucas Pall.) in Obsk Gulf]. Вуий. МП
Zool. MGU.

Dukhovnyi, М.М. 1934. Promysel belukhi Delphinapterus leucas у Obskoi
gube letom 1931 g. (Hunting of beluga Delphinapterus leucas in Obsk
Gulf in summer of 1931). Rab. Obsko-Tazovskoi Nauch. о
St. VNIRO, vol. 2, по. 1.

Egorova, L.N. 1953. Podzheludochnaya zheleza kitov kak endokrinnoe
i fermentnoe syr’e (soobshch. 1) [Pancrease of whales as endocrine
and enzyme raw material (report 1)]. Tr. VNIRO, vol. 25.

Egorova, L.N. and T.M. Lebedeva. 1953. Mozg kitov kak istochnik
polucheniya kholesterina (soobshch. 1) [Whale brain as a source
of producing cholesterol (report 1)]. Tr. VNIRO, vol. 25.

Esipov, V.K. 1935. Zemlya Frantsa-losifa (Franz Josef Land).
Arkhangel’sk.

Esipov, V.K. 1937. Zveri, ptitsy i ryby Arktiki (Animals, Birds, and Fish
of the Arctic). Sev. Obl. Izd-vo, Arkhangel’sk.

Esipov. V.K. 1937. Poteplenie Arktiki i Subarktiki (Warming of the Arc-
tic and sub-Arctic). Nauka 1 Tekhnika, no. 1.

Faingersh, R. Ya. et al. 1953. Poluchenie belkovogo preparata iz myasa
kitoobraznykh (Protein preparation from flesh of cetaceans). Tr.
VNIRO, vol. 25.

Fedoseev, С.А. 1962. О sostoyanii zapasov 1 raspredelenii tikhookaan-
skogo morzha (Reserves and range of Pacific walrus). Zool. Zhurn.,
vol. 41, no. 7.

Fedoseev, С.А. 1963. Polovoe sozrevanie 1 tempy razmnozheniya okhot-
skoi kol’chatoi nerpy (Sexual maturity and reproduction tempo of
Okhotsk ringed seal). Sb. Morsk. Mlekop. Moscow.

Fedoseev, С.А. 1964. Ob embrional’nom, postembrional’nom roste 1
polovom sozrevanii okhotskoi kol’chatoi nerpy (Embryonal and
postembryonal growth and sexual maturity of Okhotsk ringed seal).
Zool. Zhurn., vol. 43, no. 8.

Fedoseev, G.A. 1964a. Ob osnovnykh pokazatelyakh dinamiki chislen-
nosti kol’chatoi nerpy Okhotskogo morya (Main indices of the pop-
ulation dynamics of ringed seal in the Sea of Okhotsk). Sb. Sovr.
Probl. Izuch. Dinamiki Chisl. Populyatsii Zhivotnykh. Moscow.

Fedoseev, С.А. 1965a. Vozrastno-polovoi sostav poboek okhotskoi
kol’chatoi nerpy Phoca hispida ochotensis Pall. kak pokazatel’
vozrastnoi struktury populyatsii (Age and sex composition of herds
of Okhotsk ringed seal Phoca hispida ochotensis Pall. as an index to
the age structure of the population). Sb. Morsk. Mlekop. Moscow.

912

Fedoseev, С.А. 19655. Sravnitel’naya kharakteristika populyatsii
kol’chatoi nerpy pribrezhnykh vod Chukotskogo poluostrova
(Comparative characteristics of ringed seal populations in the coastal
waters of Chukchi Peninsula). Izv. TINRO, vol. 59.

Fedoseev, G.A. 1965c. K ekologii razmnozheniya tyulenei severnoi chasti
Okhotskogo morya (Reproduction ecology of seals in the northern
part of the Sea of Okhotsk). Izv. TINRO, vol. 59.

Fedoseev, С.А. 19654. Pitanie kol’chatoi nerpy (Phoca hispida ochotensis
Pall.) [Feeding of ringed seal (Phoca hispida ochotensis Pall.)]. Izv.
TINRO, vol. 59.

Fedoseev, G.A. 1965e. Opredelenie vozrastno-polovoi struktury populy-
atSil i sostoyanie zapasov okhotskikh tyulenei (Determining the age
and sex structure of populations and status of reserves of Okhotsk
seals). Zool. Zhurn., vol. 44, no. 6.

Fedoseev, G.A. 1965f. Polovoe sozrevanie i tempy razmnozheniya okhot-
skoi kol’chatoi nerpy (Sexual maturity and reproduction tempo of
Okhotsk ringed seal). Sb. Morsk. Mlekop. Nauka, Moscow.

Fedoseev, G.A. 1966a. Aerovizual’nye nablyudeniya za morskimi mleko-
pitayushchimi v Beringovom i Chukotskom moryakh (Aerial obser-
vations of marine mammals in the Bering and Chukchi seas). Izv.
TINRO, vol. 58.

Fedoseev, С.А. 1966b. О sostoyanii zapasov 1 ratsional’nom ispol’zovanii
tyulenei Okhotskogo morya (Status of reserves and rational exploita-
tion of seals in the Sea of Okhotsk). Izv. TINRO, vol. 58.

Fedoseev, С.А. 1966c. Opyt opredeleniya chislennosti 1 obosnovanie
normy vyboya kol’chatoi nerpy (akiby) v Okhotskom more (Deter-
mining the population and rationalizing the killing norms of ringed
seal in the Sea of Okhotsk). Tez. Dokl. III Vses. Soveshch. po Izuch.
Morsk. Mlekop. Moscow.

Fedoseev, G.A. 1968. Opredelenie chislennosti i obosnovanie normy
vyboya kol’chatoi nerpy (akiby) v Okhotskom more (Determining
the population and rationalizing the killing norms of ringed seal in
the Sea of Okhotsk). Tr. VNIRO, vol. 68; Izv. TINRO, vol. 62.

Fedoseev, С.А. 1969. Promyslovoe iz’yatie 1 chislennost’ otdel’nykh
pokolenii akiby (Hunting rate and population of different
generations of ringed seal). Tez. Dokl. IV Vses. Soveshch. po Izuch.
Morsk. Mlekop. Moscow.

Fedoseev, G.A. 1970. Morfekologicheskie tipy populyatsii kol’chatoi
пегру Severnoi Patsifiki. Populyats. struktura vida и mlekopit.
(Morpho-ecological Types of Ringed Seal Populations in the North
Pacific. Types of Population Structure of Mammals). Moscow.

IS)

Fedoseev, С.А. 1970. Raspredelenie 1 chislennost, tyulenei и о. Sakhalina
(Distribution and population of seals on Sakhalin Island). Izv.
TINRO, vol. 71.

Fedoseev, С.А. 1971. Raspredelenie 1 chislennost’ tyulenei na detnykh i
linnikh zalezhkakh v Okhotskom more (Distribution and population
of seals in nurseries and molting rookeries in the Sea of Okhotsk).
Tr. AtlantNIRO, no. 39.

Fedoseev, С.А. 1972. Nekotorye morfofiziologicheskie osobennosti
tyulenei Okhotskogo morya 1 ikh ekologicheskaya obuslovlennost’
(Some morpho-physiological features of seals of the Sea of Okhotsk
and their ecological background). 7r. VNIRO, vol. 82; Izv. TINRO,
vol. 80.

Fedoseev, G.A., V.N. Gol’tsev, and G.M. Kosygin. 1970. Aerovizual’nyi
uchet na shchennykh zalezhkakh v Okhotskom more (Aerial census
in nurseries of the Sea of Okhotsk). /zv. TINRO, vol. 70.

Fedoseev, С.А. and Yu.I. Nazarenko. 1970. К voprosu о vnutrividovoi
strukture kol’chatoi nerpy Arktiki (Intraspecific structure of ringed
seal of the Arctic). Izv. TINRO, vol. 70.

Fedoseev, С.А. and A.V. Yablokov. 1965. Morfologicheskaya kharak-
teristika nerpy Okhotskogo morya (Morphological characteristics of
ringed seal of the Sea of Okhotsk). Zool. Zhurn., no. 5.

Filimonovy, N.I. 1949. Sravnitel’naya anatomiya kory bol’shogo mozga
mlekopitayushchikh (Comparative Anatomy of Cerebral Cortex of
Mammals). Izd-vo Ak. Med. Nauk SSSR.

Formozov, A.N. 1929. Ostrov. Kil’din i ego fauna (Kil’din Island and its
fauna). Tr. po Lesn. Opytn. Delu TsLOS, Otd. Biol. i Prom. Okhoty,
vol. 6.

Freiman, S.Yu. 1936a. Respredelenie lastonogikh v moryakh Dal’nego
Vostoka (Distribution of pinnipeds in the Far East seas). Tr. ИМКО,
vol. 3.

Freiman, S.Yu. 1936b. Materialy К promyslovoi biologii tyulenei
Dal’nego Vostoka (Data on hunting biology of seals in the Far East).
Tr. VNIRO, vol. 3.

Freiman, S.Yu. 1936c. Promyslovaya kharakteristika severnoi chasti
Okhotskogo morya (Hunting characteristics in the northern part of
the Sea of Okhotsk). Tr. VNIRO, vol. 3.

Freiman, S.Yu. 1939. О migratsii grenlandskogo tyulenya (Harp
seal migration). Sb. Posvyashch. Nauch. Deyat. N.M. Knipovicha.
Moscow.

Freiman, S.Yu. 1941. Materialy po biologii Chukotskogo morzha (Data
on biology of Chukchi walrus). Izv. TINRO, vol. 20.

914

Freiman, S.Yu. 1950. О raspredelenii del’fina-belobochki у letnie mesy-
atsy u beregov Kryma i Severnogo Kavkaza (Distribution of com-
mon dolphin in summer along the coasts of Crimea and northern
Caucasus). Tr. Az-Chern. NIRO, vol. 14.

Gadzhiev, D.V. 1957. Morfologicheskie cherty stroeniya nosovykh kostei
kaspiiskogo tyulenya i ikh taksonomicheskoe znachenie (Morpho-
logical features of the structure of the nasal bones of Caspian seal
and their taxonomic significance). Dokl. AN Az. SSR, vol. 13, no. 4.

Gadzhiev, D.V. 1959. Novye materialy po el’darskoi faune (New data on
Eldar fauna). Izv. AN Az. SSR, Ser. Geol.-Geogr. Nauk, no. 4.

Gakichko, S. 1931. Osennii promysel lastonogikh v zalive Tyk (Autumn
hunting of pinnipeds in Tuk Вау). Rybn. Khoz. DV, nos. 1-2.

Gakichko, S.I. and S.N. Surzhin. 1936. Lastonogie Tatarskogo proliva
i Amurskogo limana kak promyshlennoe syr’e (Pinnipeds of Tatar
Strait and Amur liman as exploitable raw material). Tr. VNIRO,
vol. 3.

Gavrilenko, N.M. 1951. Soderzhanie vitamina A v solenoi pecheni kitov
(Vitamin A content in the saline liver of whales). Izv. TINRO, vol. 34.

Gebel’, Е. 1888. О kitakh murmanskogo berega $ opisaniem dvukh
novykh vidov (Whales on Murman coast with a description of two
new species). Vestn. Ryboprom., no. 3.

Geptner (Heptner), V.G. 1930. Materialy k poznaniyu raspredeleniya i
biologii belukhi (Delphinapterus leucas Pall.) [Data on distribution
and biology of beluga (Delphinapterus leucas Pall.)]. Tr. МП Zoologi
pri MGU, vol. 4, no. 2.

Geptner (Heptner), V.G. 1936. Material, po mlekopitayushchim
ostrova Dikxona, prilezhashchei chasti severo-zapadnogo Taimyra 1
Karskogo morya (Data on mammals of Dikson Island, adjacent part
of northwestern Taimyr, and the Kara Sea). Tr. Gos. Zool. Muzeya
pri MGU, vol. 3.

Golenchenko, A.P. 1935. Promysel nerpy v Ladozhskom ozere (Hunting
of ringed seal in Lake Ladoga). Za Rybnuyu Industriyu Severa, no. 2.

Golenchenko, A.P. 1949a. Raspredelenie ryby i morzverya v Tatarskom
prolive i Okhotskom more (Distribution of fish and marine animals
in Tatar Strait and Sea of Okhotsk). Rybn. Khoz., no. 2.

Golenchenko, A.P. 1949b. Raspredelenie i migratsii chernomorskogo
del’fina-belobochki v vostochnoi chasti Chernogo morya (Distribu-
tion and migration of Black Sea common dolphin in the eastern part
of the Black Sea). Rybn. Khoz., no. 3.

Golenchenko, A.P. 1961. Nerpa Belogo, Barentseva i Karskogo morei
(Seals of the White, Barents, and Kara seas). Ses. Uch. Sov. po Probl.

ЭЛЬ

Teoret. Osnovy Rats. Ispol’zov. Vosproizv. 1 Povysh. Rybn. 1 Nerybn.
Res. (Tez. Dokl.). Petrozavodsk.

Golovlev, I.F. 1960. Tekhnika kitobioinogo promysla (Whale Hunting
Technique). Kaliningradskos Obl. Izd-vo.

Gol'tsev, У.М. 1968. Dinamika beregovykh lezhbishch morzha У svyazi
a ego raspredeleniem i chislennost’yu (Dynamics of walrus coastal
rookeries in relation to animal distribution and population). Ту.
VNIRO, vol. 68; Izv. TINRO, vol. 62.

Gol’tsev, V.N. 1969. Pitanie beringovomorskoi largi у vesenne-letnii
period (Feeding of Bering Sea larga seal in the autumn-summer
period). IV Vses. Soveshch. po Izuch. Morsk. Mlekop. (Tez. Dokl.).
Moscow.

Gol’tsev, V.N. 1971. Pitanie largi (Feeding of larga seal). Ekologiya, no. 2.

Gol’'tsev, V.N. and С.А. Fedoseev. 1970. Dinamika vozrastnogo sostava
zalezhek 1 vosproizvoditel’naya sposobnost’ populyatsii largi (Age
composition dynamics in rookeries and the reproductive faculty of
larga seal populations). Izv. TINRO, vol. 71.

Gondatti, N.L. 1897. Osedloe naselenie r. Anadyra (Settled population
of Anadyr River). Zap. Priamursk. Ош. Imp. Russk. Сеовта]. Ob-va,
vol. 3, no. 1.

Gorbunov, G.P. 1929. Material po faune mlekopitayushchikh i ptits
Novoi Zemli (Data on the fauna of mammals and birds of Novaya
Zemlya). Tr. In-ta po Izuch. Severa, no. 40.

Gorbunov, G.P. 1940. Nablyudeniya nad narvalami (Observations on nar-
whals). Priroda, no. 6.

Govorkov, I.V. 1934. Belukha. Lov i obrabotka (Beluga. Trapping and
Processing). Snabtekhizdat, Moscow-Leningrad.

Grebnitskii, N.A. 1902. Komandorskie ostrova (Commander Islands).
St. Petersburg.

Grimm, О.А. 1907. Vopros о kaspiiskom tyulene (About Caspian seal).
Vest. Ryboprom., nos. 4 and 5.

Gudkoyv, У.М. 1963. Ob osobennostyakh okraski kashalotov vod
Dal’nego Vostoka (Color characteristics of sperm whales in the
waters of the Far East). Tr. In-ta Okeanol. AN SSSR, Vol. 71

IV’ina, Е.О. 1950. Ostrovno zverovodstvo (Island Fur Farming). Mezh-
dunarodnaya Kniga.

lokhel’son, V.I. 1898. Ocherk zveropromyshlennosti i torgovli mekhami у
Kolymskom okruge (Résumé of fur farming and fur trade in Kolyma
division). Tr. Yakutskoi Ekspeditsii, sect. 3, vol. 10, pt. 3.

Ivanov, Т.М. 1938. Baikal’skaya nerps, ee biologiya i promysel (Baikal
seal, its biology and hunting). [2у.: Biol.-Geogr. In-ta pri Irkut. Un-te,
vol. 8, nos. 1 and 2.

916

Ivanova, Е.Г. 1955. Kharakteristika proportsii tela kashalotov (Body pro-
portions of sperm whales). Tr. In-ta Okeanologii AN SSSR, vol. 18.

Ivanova, E.I. 1961a. Proportsii tela i rost finvala (Balaenoptera physalus
L.), dobyvaemogo u Kuril’skikh ostrovov [Body proportions and
growth of fin whale (Balaenoptera physalus L.) caught off the Kuril
Islands]. Tr. In-ta Мог). Zhiv. AN SSSR, по. 34.

Ivanova, Е.1. 1961b. Proportsii tela seival (Balaenoptera borealis), doby-
vaemogo u Kuril’skikh ostrovov [Body proportions of sei whale
(Balaenoptera borealis) caught off the Kuril Islands]. Tr. In-ta Morf.
Zhiv. AN SSSR, no. 34.

Ivanova, Е.1. 1961с. О tikhookeanskoi kosatka (Orcinus orca Г.) [ Paci-
fic killer whale (Orcinus orca L.)]. Tr. In-ta Morf. Zhiv. AN SSSR,
no. 34. |

Ivanova, Е.Г. 19614. Proportsii tela i kharakter rosta kitov Dal’nego
Vostoka (Body proportions and growth characteristics of Far East
whales). [khtiol. Komis. AN SSSR. Tr. Soveshch., no. 12.

Ivashin, M.V. 1958a. Obrestanie finvalov diatomovymi vodoroslyami
v Antarktike (Choking of fin whales with diatomaceous algae in
Antarctica). Tr. VNIRO, vol. 33.

Ivashin, M.V. 1958b. Metodika opredeleniya sledov zheltykh tel bere-
mennosti i oOvulyatsii и gorbatogo kita (Method for determining
traces of the corpus luteum of pregnancy and ovulation in humpback
whale). Tr. VNIRO, vol. 33.

Ivashin, M.V. 1958c. K biologii razmnozheniya gorbatogo kita v yuzhnoi
chasti Atlanticheskogo okeana (Reproduction biology in humpback
whale in the South Atlantic Ocean). Inf. Sb. VNIRO, по. 2.

Ivashin, M.V. 1958d. Okraska tela gorbatogo kita (Megaptera nodosa
Bonnaterre, 1789) yuzhnoi chasti Atlanticheskogo okeana [Body
color of humpback whale (Megaptera nodosa Bonnaterre, 1789) in
the South Atlantic Ocean]. Inf. Sb. VNIRO, no. 2.

Ivashin, M.V. 1958e. O sistematicheskom polozhenii gorbatykh kitov
(Megaptera nodosa lalandii) yuzhnogo polushariya [Taxonomic posi-
tion of humpback whale (Megaptera nodosa lalandii) in the Southern
Hemisphere]. Inf. Sb. VNIRO, no. 3.

Ivashin, M.V. 1959a. О rasprostranenii stad gorbatogo kita у yuzhnom
розваги (Range of humpback whale herds in the Southern Hemi-
sphere). Inf. Sb. VNIRO, no. 7.

Ivashin, M.V. 1959b. Orezmnozhenii gorbatogo kita (Megaptera nodosa
lalandii) у yuzhnoi chasti Atlanticheskogo okeana [Reproduction of
humpback whale (Megaptera nodosa lalandii) in the South Atlantic
Ocean). Inf. Sb. VNIRO, no. 7.

917

Ivashin, M.V. 1960. Znachenie razlichnykh kormovykh ob’’ektov у pitanii
gorbatogo kita (Importance of various food objects to humpback
whale). Inf. Sb. VNIRO, no. 8.

Ivashin, M.V. 1961. O periodichnosti pitaniya gorbatogo kita v yuzhnoi
chasti Atlanticheskogo okeana (Feeding cycle of humpback whale in
the South Atlantic Ocean). Byull. MOIP, Otd. Biol., vol. 66, no. 6.

Ivashin, M.V. 1965a. O prodolzhitel’nosti beremennosti i razmere
novorozhdennykh detenyshei gorbatogo kita у yuzhnoi chasti
Atlanticheskogo okeana (Extent of gestation and size of newborn
claves of humpback whales in the South Atlantic Ocean). Sb. Morak.
Mlekop. Moscow.

Ivashin, M.V. 1965b. Obrastaniya i ektoparazity u gorbatykh kitov
(Epigrowths and ectoparasites among humpback whales). Sb. Morsk.
Mlekop. Moscow.

Ivashin, M.V. 1966. Ambra i promysel kashalotov (Ambergris and sperm
whale hunting). Priroda, no. 7.

Ivashin, M.V., L.A. Popov, and AS. Tsapko. 1972. Morskie
mlekopitayushchie (Spravochnik) [Marine Mammals (Reference
Work)]. Moscow.

Karpovich, V.N. 1969. Ob uperyadochenii dobychi tyulenei v raienakh
ikh nizkoi chislennosti na Belem i Barentsevom moryakh (System-
atizing seal hunting in low population areas in the White and Barents
seas). Sb. Morsk. Mlekop. Moscow.

Karpovich, V.N., V.D. Kokhanov, and N.P. Tatarinkova. 1967. Seryi
tyulen’ na Murmane (Gray seal in Murman). 7r. PINRO, no. 21.
Kessler, K. 1861. Puteshestvie s zoologicheskoi tsel’yu k severnomu
beregu Chernogo morya i v Krym v 1858 g. (Zoological Expedition
along the North Coast of the Black Sea and in Crimea in 1858).

Kiev.

Kester, B.V. and V.N. Karpovich. 1969. Kandalakshskii zapovednik
(Kandalakshsk sanctuary). In: Zapovedniki Sovetskogo Soyuza. Kolos,
Moscow.

Khar’kov, I.I. 1940. Materialy po vesovomu i khimicheskomu sostavu
kitov (Data on weight and chemical composition of whales). Tr.
VNIRO, vol. 15.

Khar’kov, [.1. 1952. Ispol’zovanie podkozhnogo sala kashalota dlya
polucheniya zhira i kozhevennogo syr’ya (Sperm whale blubber for
producing oil and leather). Rybn. Khoz., no. 7.

Khlebtsov, M. 1902. Tyulen’ est belorybitsu (Seal is a white fish). Vest.
Ryboprom., no. 7.

Khmelin, A. 1950. Kotiki v Okhotskom more (Fur seals in the Sea of
Okhotsk). Priroda, no. 8.

918

Khuzin, R.Sh. 1960. Zimne-vesennee raspredelenie belukhi у Belom
more (Winter-spring distribution of beluga in the White Sea). Tr.
PINRO, no. 12.

Khuzin, R.Sh. 1963. K voprosu o samostoyatel’nosti trekh stad gren-
landskogo tyulenya (Independence of three harp seal herds). Sb.
Nauchn.-issl. Rabot 1962 р. (Grenl. Туш. 1 Khokhlach). Sev. PINRO,
Arkh. Ter.-proizv. Орг. Rybn. Promysh. Arkhangel’sk.

Khuzin, R.Sh. 1963a. Materialy po biologii i promyslu grenlandskogo tyu-
lenya (Pagophilus groenlandicus Erxl., 1777) u N’yufaundlenda [Data
on the biology and hunting of harp seal (Pagophilus groenlandicus
Erxl., 1777) in Newfoundland]. Tr. PINRO, no. 15.

Khuzin, R.Sh. 1963b. Materialy po morfologicheskoi kharakteristike
trekh stad grenlandskogo tyulenya Pagophilus groenlandicus Erxl.,
1777 (Data on morphological characteristics of three herds of harp
seal Pagophilus groenlandicus Erxl., 1777). Mat. Rybokhoz. Issl. Sev.
Bass., Murmansk, no. 1.

Khuzin, R.Sh. 1964. Morfoekologicheskaya kharakteristika greniand-
skogo tyulenya yanmaienskogo stada (Morphoecological character-
istics of harp seal herd of Jan Mayen). Mat. Sessii Uch. Sov. PINRO
po Rezul'’t. Issl. у 1962 - 1963 gg. Murman.

Khuzin, R.Sh. 1967. Vozrastnoi sostav khokhlacha v Grenlandskom more
(Age composition of hooded seal in the Greenland Sea). Tr. PINRO,
no. 21.

Khuzin, R.Sh. 1969a. Razmery i temp rosta grenlandskogo tyulenya belo-
morskoi, yanmaienskoi 1 n’yufaundlendskoi populyatsii (Dimensions
and growth tempo of harp seal populations of the White Sea, Jan
Mayen, and Newfoundland). Tez. IV Vses. Soveshch. po Izuch. Morsk.
Mlekop. Moscow.

Khuzin, R.Sh. 1969b. Vozrastnye izmeneiya okraski grenlandskogo
tyulenya belomorskoi, yanmaienskoi i n’yufaundlendskoi populyatsii
(Age-related color changes of harp seal populations of the White
Sea, Jan Mayen, and Newfoundland). Tez. IV Vses. Soveshch. po
Izuch. Morsk. Mlekop. Moscow.

Khuzin, R.Sh. 1969c. Polovoe sozrevanie i popolnenie stada vzroslykh
samok grenlandskogo tyulenya (Sexual maturity and herd recovery
of adult female harp seals). Tez. [V Vses. Soveschch. po Izuch. Morsk.
Mlekop. Moscow.

Khuzin, R.Sh. 1969. Promysel i sostoyanie zapasov grenlandskogo tyu-
lenya у raione о-уа Yan-Maien (Harp seal hunting and reserves in
Jan Mayen region). Sb. Morsk. Mlekop. Moscow.

Khuzin, R.Sh. 1970. Issledovaniya belomorskogo lysuna у zimne-vesennii
period 1968 g. na bortu zverob. shkhuny “Chistopol”’ (Study of

919

White Sea harp seal in winter-spring of 1968 on board animal hunt-
ing schooner “Chistopol”’). Sb. Mat. Rybokhoz. Issl. Sev. Bass., Mat.
Otch. Sessii Uch. Sov. PINRO, no. 16 (pt. 2). Murman.

Khuzin, R.Sh. and V.A. Potelov. 1963. Issledovanie tyulenei v raione
Yan-Maiena (Study of seals in Jan Mayen region). Mater. Rybokhoz.
Issl. Sev. Bass., по. 1. Murman.

Khuzin, R.Sh. and М.Уа. Yakovenko. 1963. О raspredelenii г biologii
khokhlacha (Cystophora cristata Erxl., 1777) Grenlandskogo morya
[Distribution and biology of hooded seal (Cystophora cristata Exrl.,
1777) in the Greenland Sea]. Tr. PINRO, no. 15.

Khuzin, R.Sh. and A.V. Yablokov. 1963. O nekotorykh chertakh
funktsionirovaniya pishchevaritel’nogo trakta khokhlacha v period
molochnogo pitaniya (Some functional features of the alimentary
canal of hooded seal during the suckling stage). Zool. Zhurn., vol. 42,
no. 8.

Kirpichnikov, A.A. 1932. Nekotorye materialy po zveroboinomu
promyslu v Belom more (Some data on animal hunting in the White
Sea). Izv. Len. N.-I. Ikhtioi. In-ta, vol. 13, no. 2.

Kirpichnikov, А.А. 1937. Nablyudeniya nad morskimi mlekopi-
tayushchimi v yugo-vostochnoi chasti Karskogo morya (Observations
on marine mammals in the southeastern Kara Sea). Byull. MOIP, Otd.
Biol., vol. 46, no. 4.

Kirpichnikov, A.A. 1949a. Krupnye kitoobranznye v Sredizenom more
(Large cetaceans in the Mediterranean Sea). Priroda, no. 8.

Kirpichnikov, A.A. 1949b. Kashalot v vodakh Antarktiki (Sperm whale
in Antarctic waters). Rybn. Khoz., no. 4.

Kirpichnikov, A.A. 1950a. Nablyudeniya nad raspredeleniem krupnykh
kitoobranznykh v Atlanticheskom okeane (Observation on the range
of large cetaceans in the Atlantic Ocean). Priroda, no. 10.

Kirpichnikov, A.A. 1950b. O sovremennom rasprostranenii kashalota v
Mirovom okeane po promyslovym dannym (Present range of sperm
whale in the World Ocean based on hunting data). Byull. MOIP, Nov.
Ser., Otd. Biol., vol. 55, no. 5.

Kirpichnikov, A.A. 1964. O proiskhozhdenii kaspiiskogo tyulenya (Ori-
gin of Caspian seal). Byull. MOIP, vol. 19 (5).

Kizevetter, I. 1966. Lastonogie kak promyshlennoe syr’e (Pinnipeds as
exploitable raw material). In: Dal’nevostochnye Lastonogie. V\adivos-
tok.

Kleinenberg, S.E. 1936. Materialy k izucheniyu pitaniya del’finov
Chernogo morya (Data on the feeding of Black Sea dolphins). Byull.
MOIP, Nov. Ser., Otd. Biol., vol. 45, no. 5.

920

Kleinenberg, S.E. 1938. Nekotorye dannye о рцапи Tursiops tursio Fabr.
v Chernom more (Some data on the food of Tursiops tursio Fabr.
in the Black Sea). Byull. MOIP, Nov. Ser., Otd. Biol., vol. 47, nos. 5
and 6.

Kleinenberg, S.E. 1939. Zalezhki 1 promysel kaspiiskogo tyulenya na
ostrovakh Apsheronskogo arkhipelaga (Caspian seal rookeries and
hunting in the Apsheron archipelago islands). Rybn. Khoz., no. 5.

Kleinenberg, S.E. 1940. Pitanie i dinamika upitannosti Delphinus delphis
ponticus Baranasch (Feeding and Its Dynamics in Delphinus delphis
ponticus Baranasch). Izd-vo MOIP, Moscow.

Kleinenberg, S.E. 1951. O faune mlekopitayushchikh Chernogo morya
(Black Sea mammalian fauna). ВуиЙ. MOIP, Ош. Biol., Vol. 56 (1).

Kleinenberg, S.E. 1956a. Mlekopitayushchie Chernogo i Azovskogo
morei (Mammals of the Black and Azov seas). Izd-vo AN SSSR,
Moscow.

Kleinenberg, S.E. 1956b. Osobennosti dykhaniya kitoobraznykh (Respi-
ratory characteristics of cetaceans). Usp. Sovr. Biol., vol. 41, no. 3.

Kleinenberg, S.E. 1957. Ispol’zovanie morfologicheskikh i fiziologich-
eskikh pokazatelei у analize ekologii nekotorykh vodnykh mlekopi-
tayushchikh (Utilization of morphological and physiological indices
in analyzing the ecology of some aquatic mammals). Vopr. Ekol.,
vol. 1.

Kleinenberg, S.E. 1957. Ob okhrane morzha (Walrus conservation).
Priroda, no. 7.

Kleinenberg, S.E. 1958. K voprosu o proiskhozhdenii kitoobraznykh
(Origin of cetaceans). DAN SSSR, vol. 122, no. 5.

Kleinenberg, S.E. V.M. Bel’kovich, and A.V. Yablokov. 1960. Materialy k
izucheniyu raspredeleniya belukhi v severnykh moryakh SSSR (Data
on range of beluga in the northern seas of the USSR). Tr. PINRO,
no. 12.

Kieinenberg, S.E., V.M. Bel’kovich, and A.V. Yablokov. 1964. Materialy
k izucheniyu rasprostraneniya i sostoyaniya populyatsii morzhei
Sovetskoi Arktiki (Data on range and population of Soviet Arctic
walruses). In: Opredelenie Vozrasta Promyslovykh Lastonogikh 1
Ratsional’noe Ispol’zovanie Morskikh Mlekopitayushchikh. Moscow.

Kleinenberg, S.E. and A.V. Yablokov. 1958. K morfologii verkhnikh
dykhatel’nykh putei kitoobranznykh (Morphology of upper respira-
tory tract among cetaceans). Zool. Zhurn., vol. 37, no. 7.

Kleinenberg, S.E. and A.V. Yablokov. 1960. Materialy po biologii razm-
nozheniya belukhi severnykh morei SSSR (Data on reproduction
biology in beluga in northern Soviet seas). Tr. PINRO, no. 12.

921

Kleinenberg, 5.Е., A.V. Yablokov, У.М. Bel’kovich, and М.М. Тагазе-
vich. 1964. Belukha (Beluga). Moscow.

Kleinenberg, S.E. et al. 1965. Spravochnye pokazateli po kharakteristike
nekotorykh lastonogikh i kitoobranznykh (Standard indices of char-
acteristics for some pinnipeds and cetaceans). In: Morsk. Mlekopit.
Moscow.

Klevezal’, G.A. 1964. Opredelenie tempa rosta i vremeni nastupleniya
polovoi zrelosti lastonogikh (Determining growth tempo and time of
onset of sexual maturity among pinnipeds). In: Opredelenie Vozrasta
Promyslovykh Lastonogikh т Ratsional’noe Ispol’zovanie Morskikh
Mlekopitayushchikh. Nauka, Moscow.

Klevenzal’, G.A. and D.D. Tormosov. 1971. O vydelenii lokal’nykh grup-
pirovok kashalotov po kharakteru sloev v dentine zubov (Isolating
local groups of sperm whales based on type of layers in tooth den-
tine). Tr. AtlantNIRO, no. 39.

Klikh, N. 1933. Grenlandskii tyulen’ (Greenland seal). Sov. Sever., no. 6.

Klumoy, S.K. 1934. К voprosu о raspredelenii i promysle belukhi у Belom
more (Range of beluga and hunting in the White Sea). Rybn. Khoz.
SSSR, no. 4.

Кштох, S.K. 1935. Ostrov Vaigach, ego fauna i promyslovye vozmozh-
nosti (Vaigach Island, its fauna, and hunting possibilities). Sov. Ark-
tika, no. 2.

Klumov, 5.К. 1936a. Raspredelenie belukhi (Delphinapterus leucas Pall.)
na Evropeisko-Aziatskom Severe [Range of beluga (Delphinapterus
leucas Pall.) in the European-Asian north]. 77. Polyar. Komis. AN
SSSR, no. 27.

Klumov, 5.К. 1936b. Dobycha i pervichnaya obrabotka belukhi na severe
i Dal’nem Vostoke (Trapping and Preliminary Processing of Belugas
in the North and Far East). Pishchepromizdat, Moscow.

Klumov, S.K. 1937a. Novye formy belukhi (New beluga forms). Izv. AN
SSSR, Biol. Ser., no. 1.

Klumov, S.K. 1937b. Saika (Boreogadus saida Lepech.) i ее znachenie
dlya nekotorykh zhiznennykh protsessov Arktiki [Polar cod (Bore-
ogadus saida Lepech.) and its significance for some artic life pro-
cesses]. Izv. AN SSSR, Otd. Mat. i Est. Nauk.

Klumov, S.K. 1939. Belukha Sovetskogo Severa (syr’evaya baza i promy-
sel) [Beluga in the Soviet north (raw material base and hunting)].
Tr. VNIRO, vol. 12.

Klumov, S.K. 1954. О tsikle razmnozheniya chernomorskogo del’fina-
belobochki (Reproduction cycle of Black Sea common dolphin). Ту.
In-ta Okeanol. AN SSSR, vol. 7

922

Klumov, S.K. 1955. О lokalnosti kitovykh stad (Localization of whale
herds). Tr. In-ta Okeanol. vol. 18.

Klumov, S.K. 1956. Nekotorye itogi ekspeditsii у Beringovo more i na
Kuril’skie ostrova (Some results of an expedition in the Bering Sea
and on the Kuril Islands). Vest. AN SSSR, vol. 5.

Klumov, S.K. 1957а. Beregovye lezhbishcha kotikov (Callorhinus ursinus)
i mesta obitaniya kalanov (Enhydra lutris) na Kuril’skikh ostrovakh
i orientirovochnoe opredelenie ikh chislennosti [Coastal rookeries
of fur seals (Callorhinus ursinus) and habitats of sea otters (Enhydra
lutris) on the Kuril Islands and their population estimation]. DAN
SSSR, vol. 117, no. 1.

Klumov, 5.К. 1957b. Uchet beregovykh lezhbishch sivuchei (Eumetopias
jubatus) na Kuril’ skikh ostrovakh i orientirovochnoe opredelenie
ikh chislennosti [Counting the coastal rookeries of Steller’s sea lion
(Eumetopias jubatus) on the Kuril Islands and their population esti-
mation]. DAN SSSR, vol. 117, no. 2.

Klumov, S.K. 1959. Promyslovye del’finy Dal’nego Vostoka (Game dol-
phins of the Far East). Izv. TINRO, vol. 47.

Klumov, S.K. 1962. Gladkie (yaponskie) kity Tikhogo okeana [Right
(Japanese) whales of the Pacific Ocean]. Tr. In-ta Okeanol. AN SSSR,
vol. 58.

Klumov, S.K. 1963. Pitanie i gel’mintofauna usatykh kitov (Mystacoceti)
у osnovnykh promyslovykh raionakh Mirovogo okeana [Feeding and
helminth fauna of baleen whales (Mystacoceti) in the main hunting
regions of the World Ocean]. Tr. In-ta Okeanol. AN SSSR, vol. 71.

Klumoy, S.K. and 1.1. Barabash. 1938. Tikhookeanskaya forma belukhi
(Pacific Ocean form of beluga). Rybn. Khoz. SSSR, no. 11.

Klyuge, G. 1936. Biologicheskie raboty na polyarnoi stantsii mysa Zhe-
laniya v 1934/35 g. (Biological studies carried out at Cape Zhelaniya
polar station in 1934/35). Byull. Ат. In-ta, no. 4.

Knipovich, N.M. 1895. Polozhenie morskikh rybnykh i zverinykh
promyslov Arkhangel’skoi gubernii (Status of Marine Fish and
Animal Hunting in Arkhangel’sk Province). St. Petersburg.

Knipovich, N.M.- 1897. О rybnykh i morskikh zverinykh promyslakh
Arkhangel’skoi gubernii (Fishing and Marine Animal Hunting in
Arkhangel’sk Province). St. Petersburg.

Knipovich, N.M. and K.P. Yagodovskii. 1902. Razvedochnaya eksped-
itsiya, ch. 1 (Exploratory expedition, pt. 1). In: Ekspeditsiya dlya
Nauchn.-Prom. Issl. u Beregov Murmana, vol. 1. St. Petersburg.

Knipovich, N.M., K.P. Yagodovskii, and N.S. Zhikharev. 1902. Raboty
ekspeditsii s 10 (22) maya 1899 g. po 31 dekabrya 1899 g. (po n.
st. 12 yanv. 1900 g.) [Expedition activity from May 10 (22), 1899

923"

through December 31, 1899 (according to new calendar January 12,
1900]. In: Ekspeditsiya dlya Nauch.-Prom. Issl. и Beregov Murmana.
St. Petersburg.

Kogai, V.M. 1968. Sovremennoe sostoyanie i dinamika chislennosti stada
kotikov па о. Tyulen’em (Population status and dynamics of fur seal
herds on Tyulen’ Island). Tr. VNIRO, vol. 68; Izv. TINRO, vol. 62.

Kolchev, V.V., Z.V. Marieva, and V.D. Koval’. 1954. Poluchenie zhe-
latiny iz sala kitov (Gelatin production from whale blubber). Ту.
VNIRO, vol. 29.

Kolyushev, М. М. 1933. Materialy po fauna i promyslu Zapadnogo ugla
Taimyrskogo poluostrova (Data on the fauna and hunting in the
western corner of Taimyr Peninsula). Mater. po Izuch. Sibiri, vol.4.

Kondakov, N.N. 1960. К voprosu о sistematicheskom polozhenii
baikal’skogo tyulenya (Taxonomy of Baikal seal). Byull. MOIP, Otd.
Biol., vol. 65 (4).

Korabel’nikov, L.V. 1959. О pitanii kashalotov у antarkticheskikh
moryakh (Feeding of sperm whales in Antarctic seas). Priroda, no. 3.

Korzhuev, P.A. and M.M. Bulatova. 1952. Dykhatel’naya funktsiya krovi
del’finov (Respiratory function of the blood of dolphins). Tr. In-ta
Merf. Zhiv. AN SSSR, no. 6.

Koshkin, V.N. 1937. Nekotorye dannye po promyslovoi fauna ostrova
Begicheva (Some data on game fauna on Begichev Island). Probl.
Arktiki, no. 2.

Koshkin, V.N. 1940. Lezhbishche morzhei na o. Preobrazhen’e (Walrus
rookeries on Preobrazhen’e Island). Probl. Arktiki, no. 5.

Kosygin, С.М. 1966a. Raspredelenie i nekotorye cherty biologii
lastonogikh Beringova morya (vesenne-letnii period 1963 g.)
[Distribution and some biological features of Bering Sea pinnipeds
(spring-summer of 1963)]. Izv. TINRO, vol. 58.

Kosygin, G.M. 1966b. Raspredelenie lakhtaka v Beringovom more v
vesennii period 1962-1964 gg. (Distribution of Bering Sea bearded
seal in 1962 - 1964 spring). Izv. TINRO, vol. 58.

Kosygin, G.M. 1966c. Razmery i ves lakhtaka Beringova morya (Size and
weight of Bering Sea bearded seal). Izv. TINRO, vol. 58.

Kosygin, G.M. 1966d. Nekotorye materialy po pitaniyu lakhtaka v
Beringovom more v vesenne-letnii period (Some data on Bering Sea
bearded seal feeding in spring-summer). Izv. TINRO, vol. 58.

Kosygin, С.М. 1969. К kraniologicheskoi kharakteristike lakhtaka
Beringova morya (Craniological features of Bering Sea bearded seal).
In: Morsk. Mlekop. Moscow. .

Kosygin, С.М. and У.А. Potelov. 1972. Vozrastnaya, polovaya i pop-
ulyatsionnaya izmenchivost’ kraniologicheskikh priznakov morskogo

924

zaitsa (Age-, sex-, and populationwise variations of craniological fea-
tures of bearded seal). Tr. VNIRO, vol. 82; Izv. TINRO, vol. 80.

Kosygin, G.M. and E.A. Tikhomirov. 1969. Larga zaliva Petra Velikogo
(Гагра seal in Peter the Great Bay). IV Vses. Soveshch. po Izuch.
Morsk. Mlekop. (Tez. Dokl.). Moscow.

Kosygin, G.M. and E.A. Tikhomirov. 1970. Larga zaliva Petra Velikogo
(Larga seal in Peter the Great Bay). Izv. TINRO, vol. 70.

Kovalev, K.V. 1938. Migratsii i promysel belukhi v Eniseiskom i Pyasin-
skom zalivakh (Beluga migrations and hunting in the Yenisey and
Pyasinsk straita). Tr. Arkt. In-ta, vol. 123.

Krasheninnikov, S.P. 1755. Opisanie zemli Kamchatki (Description of
Kamchatka Land), vol. 1. St. Petersburg.

Krasheninnikov, S.P. 1755. Opisanie zemli Kamchatki (Description of
Kamchatka Land). St. Petersburg. (republished by AN SSSR in
1949).

Krivosheya, М.У. 1884. Zametki о Novo-Zemel’skoi faune (Notes on
Novaya Zemlya fauna). Priroda i Okhota, November.

Krotov, A.V. 1952. Chernomorskii tyulen’ (Black Sea seal). Priroda, no. 5.

Krylov, V.I. 1965. Pervoe machenie tikhookeanskogo morzha (First
marking of Pacific Ocean walrus). Rybn. Khoz., no. 10.

Krylov, V.I. 1966a. O polovom sozrevanii samok tikhookeanskogo
morzha (Sexual maturity of Pacific Ocean female walrus). Zool.
Zhurn., vol. 45, no. 6.

Krylov, V.I. 1967. Sroki nastupleniya polovoi zrelosti samtsov morzha
(Period of onset of sexual maturity in male walruses). In: Probl. Sev-
era, no. 11.

Krylov, V.I. 1968. О sovremennom sostoyanii zapasov tikhookeanskogo
morzha i perspektivakh ikh ratsional’nogo ispol’zovaniya (Status
of Pacific Ocean walrus reserves and prospects for their rational
exploitation). Tr. VNIRO, vol. 68; Izv. TINRO, vol. 62.

Krylov, V.I. 1969. Sroki sparivaniya i shchenki tikhookeanskogo morzha
(Periods of mating and parturition in Pacific Ocean walruses). In:
Morsk. Mlekop. Nauka, Moscow.

Krylov, V.I. 1971. О pitanii tikhookeanskogo morzha (Feeding of Pacific
Ocean walrus). Tr. AtlantNIRO, no. 39.

Krylov, У.1, С.А. Fedoseev, and А.Р. Shustov. 1964. Lastonogie
Dal’nego Vostoka (Far Eastern Pinnipeds). Moscow.

Kulagin, М.М. 1928. К poznaniyu lin’ki belomorskogo tyulenya (Histrio-
phoca groenlandica Lepechin) [Data on the molting of White Sea
seal (Histriophoca groenlandica Lepechin)]. DAN SSSR, nos. 20 and
2%.

925

Kulagin, М.М. 1929. Vodnye promyslovye mlekopitayushchie SSSR
(Hunting of Mammals in Soviet Waters). Izd-vo Gos. Politekh.
Muzeya.

Kuleshov, P. Ya. 1950. Kamchatskii morskoi lev—sivuch (Kamchatka sea
lion—Steller’s sea lion). Priroda, no. 5.

Kurochkin, Yu.V. 1958. Gel’mintofauna kaspiiskogo tyulenya v period
razmnozheniya. Rab. po gel’mintologii (Helminth fauna of Caspian
seal in the reproduction period. Work on helminthology). In: K 80-
letiyu akad. К. Skryabina. Moscow.

Kurochkin, Yu.V. 1962. Gel’mintofauna kaspiiskogo tyulenya i ee
znachenie v d biotsenozakh del’ty Vol’gi (Helminth fauna of Caspian
seal and its importance in the biocenoses of the Volga delta). 7r.
Soveshch. Ikhtiol. Komis. AN SSSR, по. 12.

Kurochkin, Yu.V. and У.1. Zablotskii. 1958/59. К gel’mintofaune kaspi-
iskogo tyulenya (Helminth fauna of Caspian seal). Tr. Astr. Gos.
Zap-ka, no. 4.

Kurcheva, G. 1955. Lakhtak (Bearded seal). Tr. In-ta Okeanol. AN SSSR,
vol. 14.

Kusto, Zh.I. and Е. Dyuma. 1953. У mire bezmoiviya (In the Silent
World). Moscow.

Kuzin, A.E. 1968. Nekotorye dannye po embriogenezu cherepa morskikh
kotikov (Some data on the embryogeny of the skull of fur seals). Ти.
VNIRO, vol. 68; Izv. TINRO, vol. 62.

Kuzin, A.E. 1972. Morfologicheskaya kartina krovi morskikh kotikov v
ontogeneze (Morphology of the blood of fur seals in ontogeny). Tr.
VNIRO, vol. 82; Izv. TINRO, vol. 80.

Kuzin, A.E., М.К. Maminov, and A.S. Perlov. 1973. Dinamika
chislennosti 1 dal’neishie predpolozhitel’nye регзрекиху rosta stada
morskikh kotikov na Kuril’skikh ostrovakh (Population dynamics
and future prospects of the growth of fur seal herds on Kuril Islands).
Ekologiya, no. 4.

Kuznetsov, 1.1. 1937. Keta i ee vosproizvodstvo (Whale and Its Repro-
duction). Khabarovsk.

Lagovskaya, E.A. 1947. Pechen’ kitov kak syr’e dlya polucheniya vitamina
A (Whale liver as raw material for producing vitamin A). Izv. TINRO,
vol. 23.

Laktionov, A.F. 1946. Severnaya Zemlya (Severnaya Zemlya). Moscow-
Leningrad.

Leis, M. 1960. Kol’chatyi tyulen’ Rizhskogo zaliva (Viiger—hulgest Riia
lahes) [Ringed seal in Riga Bay (Viiger—hulgest Riia lahes)]. Priroda
Estoni (Eesti loodus), no. 1.

926

Lepekhin, I. 1805. Dnevnye zapiski puteshestviya po raznym provintsiyam
Rossiiskogo gosudarstva (Diary of Expeditions in Various Russian
Provinces). St. Petersburg.

Lepin, А.Т. 1936. Biologicheskie raboty na Novoi Zemle za 1934-1935
gg. (Biological work on Novaya Zemlya during 1934-1935). Byull.
Атм. In-ta, no. 2.

Levashova, E.P. 1954. Mikrostruktura shkury kita-kashalota (Microstruc-
ture of sperm whale skin). Legkaya Promyshlennost’, no. 10.

Lindberg, G.U. and G.D. Dul’keit. 1929. Materialy po rybam Shan-
tarskogo morya (Data on Shantar Sea fish). Izv. TINRO, vol. 3, no. 1.

Linko, A.K. 1912. Plankton u beregov Zapadnogo Murmana (Plankton
on west Murman coasts). Tr. Murm. Nauchn.-prom. Eksp. 1905 в.
St. Petersburg.

Lomakin, V.V. 1964. K istorii izucheniya baikal’skoi nerpy (History of
study of the Baikal seal). Byull. MOIP, Otd. Biol., no. 3.

Lun’, S.S. 1936. Lastonogie Zapadnoi Kamchatki (Tigil’skoi r-n) [West-
ern Kamchatka (Tigil’sk region) pinnipeds]. Tr. VNIRO, vol. 3.
Lyubimov, М.Р. 1927. К _ poznaniyu gel’mintofauny zhivotnykh
Moskovskogo zooparka. 1. Filyarii (Key to the helminth fauna of
Moscow zoo animals. 1. Filaria). Tr. Labor. Eksped. Biol. Mosk.

Zooparka, vol. 3.

Maiorova, A.A. and N.I. Danilevskii. 1934. Materialy po biologii cher-
nomorskogo del’fina (Data on the biology of Black Sea dolphin). Tr.
Nauch. Ryb.-khoz. St. Gruzii, vol. 1, no. 1.

Mal’m, E.N. 1932. Del’tiny Chernogo morya (Black Sea Dolphins). Izd-vo
AN SSSR. Leningrad.

Mal’m, E.N. 1936. O vremeni shchenki chernomorskogo del’fina (Del-
phinus delphis L.) [Time of parturition among Black Sea dolphins
(Delphinus delphis L.)]. Tr. Novoros. Biol. St., vol. 2, no. 1.

Manteifel’, B.P. 1943. Saika i ee promysel (Polar Cod and Its Catch).
Arkhangel’sk.

Marakov, S.V. 1965. Iz opyta promysla sivucha i largi na Komandorskikh
ostrovakh (Experience in hunting Steller’s sea lion and larga seal on
the Commander Islands). Sb. Nauch.-tekhn. Inf. VNIIZhAP, no. 12.

Marakov, S.V. 1967a. Materialy po ekologii largi Komandorskikh ostro-
vov (Data on the ecology of larga seal on the Commander Islands).
Tr. PINRO, no. 21.

Marakov, S.V. 1967b. Kitoobraznye vod Komandorskikh ostrovov
(Cetaceans in Commander Island waters). Tr. PINRO, vol. 21.

Marakov, S.V. 1968. Poberezh’ya nekotorykh ostrovnykh sistem Severnoi
Patsifiki kak arena zhizni lastonogikh i kalana (Coasts of some North

927

Pacific island systems as the life arena for pinnipeds and sea otters).
Sb. Nauch.-tekhn. Inf. VNIIZhP, по. 22.

Marakov, S.V. 1969a. Arena zhizni i prisposobitel’nye osobennosti neko-
torykh lastonogikh i kalana (Life arena and adaptation features of
some pinnipeds and sea otters). ГИ Vses. Soveshch. po Izuch. Morsk.
Mlekop. (Tez. Dokl.). Moscow.

Marakov, S.V. 1969b. K otsenke morskikh poberezhii kak okhotnich’ikh
ugodii (Evaluating sea coasts as hunting grounds). Tr. VNIIZhP,
ПО. 22.

Marakov, S.V. 1970. Ostrovnye okhotnich’i ugod’ya Severnoi Patsifiki
i puti povysheniya ikh produktivnosti (North Pacific island hunting
grounds and methods for improving their productivity). Tr. Mezhdun.
Kongr. Biol.-okhotovedov. Moscow.

Marakovy, S.V. and V.G. Voronov. 1969. О zabolevanii zubov i kostei
chelyusti u ostrovnogo tyuleny-antura [Dental and jaw bone diseases
of island (antura) seal]. IV Vses. Soveshch. po Izuch. Morsk. Mlekop.
(Tez. Dokl.). Moscow.

Marakov, S.V. and G.A. Nesterov. 1958. O vrede, nanosimom sivuchami
nakhodyashchimsya na lezhbishche kotikam (Damage caused by
Steller’s sea lions present in fur seal rookeries). Rybn. Khoz., no. 10.

Mchedlidze, G.A. 1970. Nekotorye obshchie cherty istorii kitoobraznykh
(Some Common Features of the History of Cetaceans). Tbilisi.

Middendorff, A.Th. 1869. Puteshestvie na sever i vostok Sibiri (Expedi-
tions into Northern and Eastern Siberia), pt. 2, sect. 5. Sib. Fauna.
St. Petersburg.

Mikeladze, D.A. 1971. O soderzhanii morskikh kotikov i kaspiiskikh tyu-
lenei v Batumskom akvariume (Fur and Caspian seals in the Batum
aquarium). Tr. AtlantNIRO, no. 39.

Mikhailov, S.V. 1958. Rybnye i zverinye promysly Sovetskogo Zapol-
yar’ya (Fishing and animal hunting in the Soviet Trans-polar). Probl.
Severa, no. 2.

Mikhel’, N.M. 1937. Nablyudeniya nad ptitsami i mlekopitayushchimi
raiona Karskoi guby (Observations on birds and mammals of Kara
Gulf region). Izv. Gos. Geogr. Ob-va, no. 3.

Mikhel’, N.M. 1937. Ocherk promyslovoi okhoty v Severo-Vostochnoi
Yakutii (Game hunting in northeastern Yakutia). Sb. Arktika. Vses.
Arktich. In-t, book 5. Leningrad.

Mineev, A.I. 1934. Pyat’ let na ostrove Vrangelya (stenogram. dokl. 9
noyabrya 1934 g.) [Five Years on Wrangel Island (Transcript of
Report of November 9, 1934)]. Khabarovsk.

Mineev, А.1. 1936. Promysel morzha na о. Vrangelya (Walrus hunting
on Wrangel Island). Sov. Arktika, no. 2.

928

Moiseev, Р.А. 1951. Obnaruzhenie morzha у Okhotskom more (Walrus
find in the Sea of Okhotsk). Izv. TINRO, vol. 34.

Mordvinov, Yu.E. 1968. Nablyudeniya nad lokomotsiyami nekotorykh
lastonogikh (Pinnipedia) [Observations on locomotion among pin-
nipeds (Pinnipedia)]. Zool. Zhurn., vol. 17, no. 9.

Mordvinov, Yu.E. 1969a. Dlitel’nost’ prebyvaniya pod vodoi kaspiiskikh
tyulenei (Residence duration of Caspian seals under water). Tez.
Dokl. IV Vses. Soveshch. po Izuch. Morsk. Mlekop. Moscow.

Mordvinov, Yu.E. 1969b. О gidrodinamicheskikh kachestvakh zadnikh
lastov nastoyashchikh tyulenei sem. Phocidae (ош. Pinnipedia)
[Hydrodynamic features of the caudal flukes of true seals of family
Phocidae (order Pinnipedia)]. Biologiya Morya, no. 16.

Morozov, D.A. 1971. Perevozka zhivykh del’finov (Transport of live dol-
phins). Tr. AtlantNIRO, no. 39.

Moskalenko, B.K. 1945. Nerpa Pechorskogo morya i ee promysel
(Pechora Sea ringed seal and its hunting). Rybn. Promysh. SSSR,
sb. 1. Moscow.

Mozgovoi, A.A. 1953. Askaridaty zhivotnykh i cheloveka 1 vyzyvaemye
imi zaboleveniya (Ascaridates of animals and man and the diseases
caused by them). Osnovy Nematodology, vol. 2. Izd-vo AN SSSR.
Moscow.

Mozgovoi, А.А. and K.M. Ryzhikov. 1950. Vopros о proiskhozhdenii
baikal’skogo tyulenya v svete gel’mintologicheskoi nauki (Baikal seal
genesis in the light of helminthology). DAN SSSR, vol. 72, no. 5.

Mrochkov, К.А. 1953a. Pechen’ kitov Antarktiki kak зуге Фуа
polucheniya vitamina A (Antarctic whale liver as raw material for
producing vitamin A). Tr. VNIRO, vol. 25.

Mrochkov, К.А. 1953b. Vesovoi i khimicheskii sostav otdel’nykh chastei
i nekotorykh organov finvalov (Weight and chemical composition of
individual parts and some organs of fin whales). Tr. VNIRO, vol. 25.

Muzhchinkin, V.F. 1964. Raspredelenie sivuchei na yugovostochnom
kotikovom lezhbishche ostrova Mednogo (Range of Steller’s sea lion
in the southeastern fur seal rookery on Mednyi Island). Izv. TINRO,
vol. 54; Tr. VNIRO, vol. 51.

Muzhchinkin, V.F. 1964. Sroki poyavleniya okosteneniya v konechnos-
tyakh morskikh kotikov (Periods of ossification in the limbs of fur
seals). Izv. TINRO, vol. 54; Tr. VNIRO, vol. 51.

Muzhchinkin, V.F. 1968. Vozniknovenie polovykh razlichii v cherepakh
komandorskikh morskikh kotikov (Origin of sex-related differences
in the skulls of Commander fur seals). Tr. VNIRO, vol. 68; Izv.
TINRO, vol. 62.

929.

Muzhchinkin, У.Е. 1972. Myshechnaya sistema 1 sposob peredvizheniya
severnykh morskikh kotikov (Muscular system and locomotion
among northern fur seals). 7r. VNIRO, vol. 82; Izv. TINRO,
vol. 80.

Muzhchinkin, V.F. 1972. Arterial’naya sistema severnykh morskikh
kotikov i varianty ee stroeniya (Arterial system of northern fur
seals and its structural variations). Tr. VNIRO, vol. 82; Izv. TINRO,
vol. 80.

Nansen, F. 1924. Sredi tyulenei i belykh medvedei (Among Seals and
Polar Bears). Leningrad.

Nansen, F. 1939. Grenlanskii tyulen’, ili lysun (Harp seal). Sobr. Soch.,
vol. 5. Leningrad.

Narkhov, A.S. 1937. Morfologiya muskulatury khvostovoi oblasti D. del-
phis i T. truncatus. (Morphology of caudal musculature in D. delphis
and T. truncatus). Zool. Zhurn., vol. 16, no. 4.

Naumoy, S.P. 1930. Nekotorye dannye po biologii morskikh mlekopi-
tayushchikh yugo-zap. chasti Okhotskogo morya (Some data on the
biology of marine mammals in the southeastern Sea of Okhotsk).
Rybn. Khoz. DV, no. 2.

Naumoy, S.P. 1931. Mlekopitayushchie i ptitsy Gydanskogo p-ova (Mam-
mals and birds of Gydansk Peninsula). Tr. Polyarn. Komis. AN SSSR,
no. 4.

Naumov, S.P. 1933. Tyuleni SSSR (Seals of the USSR). Moscow-
Leningrad.

Naumov, 5.Р. 1941. Lastonogie (Pinnipedia) Okhotskogo morya (Pin-
nipeds of the Sea of Okhotsk). Uch. Zap. Mosk. Gos. Ped. In-ta,
vol. 24, no. 2.

Naumov, S.P. and М.А. Smirnov. 1936. Materialy po sistematike
1 geograficheskomu raspredeleniyu Phocidae у severnoi chasti
Tikhogo okeana (Data on taxonomy and geographic range of
Phocidae in the North Pacific Ocean). Tr. VNIRO, vol. 3.

Nazarenko, Yu.I. 1962. К voprosu о sostoyanii stada grenlandskogo tyu-
lenya v Belom more (Status of harp seal herds in the White Sea).
Byull. Tekhn.-ekon. Soveta Arkh. SNKh. Arkhangel’sk.

Nazarenko, Yu.I. 1964. Mechenie belomorskogo lysuna i dannye ро
estestvennoi smertnosti molodnyaka (Marking of White Sea harp
seal and data on the natural mortality of calves). Sessiya Uch. Sov.
Karel. О. GosNIODRKh, no. 1. Petrozavodsk.

Nazarenko, Yu.I. 1965. Materialy К izucheniyu razmnozheniya kol’chatoi
nerpy (Phoca hispida) Cheshskoi guby [Data on the reproduction of
ringed seal (Phoca hispida) in Cheshsk Gulf]. Sb. Morsk. Mlekop.
Moscow.

930

Nazarenko, Yu.I. 1965. Bol’she vnimaniya beregovomu promyslu
morskogo zverya (Greater attention to coastal hunting for marine
animals). Rybn. Khoz., no. 8.

Nazarenko, Yu.I. 1967а. О zakhodakh nerpy у reki Evropeiskogo Severa
(Transgressions of ringed seal into rivers of the European North).
Tr. PINRO, no. 21.

Nazarenko, Yu.I. 1967. Pitanie kol’chatoi nerpy Evropeiskogo Severa
(Feeding of ringed seal in the European North). Tr. PINRO, no. 21.

Nazarenko, Yu.I. 1968. Kol’chataya nerpa (Pusa hispida Schreber, 1775)
Evropeiskogo Severa SSSR, ee morfologicheskaya kharakteristika 1
khozyaistvennoe ispol’zovanie [Ringed seal (Pusa hispida Schreber,
1775) in the European Soviet North, its morphological features, and
commercial exploitation]. Avtoref. diss. Petrozavodsk.

Nazarenko, Yu.I. 1969. Svedeniya о promysle severnoi kol’chatoi nerpy
(Information on the hunting of northern ringed seal). Sb. Morsk.
Mlekop. Moscow.

Nazarenko, Yu.I. and Yu.K. Timoshenko. 1963. Issledovanie biologii
belomorskogo lysuna v 1962 g. (A study of the biology of White
Sea harp seal in 1962). Sessiya Uch. Sov. po Probl. Teoretich. Osnovy
Rats. Ispol’z. Rybn. i Nerybn. Resursov Bel. Morya. Petrozavodsk.

Nazarenko, Уч.[. and Yu.K. Timoshenko. 1964. К biologii razm-
nozheniya grenlandskogo tyulenya belomorskogo stada (Reproduc- —
tion biology of White Sea harp seal). Sessiya Uch. Sov. Karel. Otd.
GosNIORKh, no. 1. Petrozavodsk.

Nazarenko, Уи.[. and A.V. Yablokov. 1962. Otsenka metoda ucheta
chislennosti belomorskogo lysuna 1 soobrazheniya о sostoyanii
zapasov etogo tyulenya (Evaluating the method of calculating the
population of White Sea harp seal and the status of its reserves).
Zool. Zhurn., vol. 41, no. 12.

Nesterov, С.А. 1964. Materialy po biologii i chislennosti sivuchei
Komandorskikh ostrovov (Data on the biology and population of
Steller’s sea lion on Commander Islands). Izv. TINRO, vol. 54; Tr.
VNIRO, vol. 51.

Nikol’skii, A.M. 1889. Ostrov Sakhalin i ego fauna pozvonochnykh zhiv-
otnykh (Sakhalin Island and Its Vertebrate Fauna).

Nikol’skii. A.M. 1891. Pozvonochnye zhivotnye Kryma (Vertebrates of
Crimea). St. Petersburg.

Nikol’skii, V.V. 1927. Byt i promysly naseleniya zapadnogo poberezh’ya
Belogo morya (Soroka-Kandalaksha) [Living conditions and occu-
pations of the people on the western coast of the White Sea (Sorok- |
Kandalaksh)]. Tr. In-ta po Izuch. Severa, no. 36.

931

Nikol’skii, С.Е. 1933. Иов raboty nad biologiei razmnozheniya
grenlandskogo tyulenya (Results of investigations on the reproduc-
tion biology of harp seal). Rybn. Khoz. SSSR, no. 1.

Nikol’skii, G.E. 1936. K biologii razmnozheniya Delphinapterus leucas
Pall. (Reproduction biology of Delphinapterus leucas Pall.). Tr.
VNIRO, vol. 3.

Nikulin, P.G. 1937a. Sivuch Okhotskogo morya i ego promysel (Steller’s
sea lion of the Sea of Okhotsk and its hunting). Izv. TINRO, vol. 10.

Nikulin, P.G. 1937b. Nablyudeniya nad lastonogimi Okhotskogo i
Yaponskogo morei (Observations on pinnipeds of the seas of
Okhotsk and Japan). Izv. TINRO, vol. 10.

Nikulin, Р.С. 1941. Chukotskii morzh (Chukchi walrus). Izv. TINRO,
vol. 20.

Nikulin, P.G. 1946. O raspredelenii kitoobraznykh v moryakh, omy-
vayushchikh Chukotskii poluostrov (Range of cetaceans in the seas
surrounding Chukchi peninsula). Izv. TINRO, vol. 22.

Nikulin, P.G. 1947. Biologicheskaya kharakteristika beregovykh lezh-
bishch morzha (Biological characteristics of coastal walrus rook-
eries). Izv. TINRO, vol. 25.

Nikulin, Р.С. 1951. О zimovke belukh (Wintering of belugas). Izv.
TINRO, vol. 34.

Nikulin, Р.С. 1968. Sovremennoe sostoyanie 1 регзрекиуу rosta populy-
atsii komandorskikh kotikov (Status and growth prospects of Com-
mander fur seal population). Tr. VNIRO, vol. 68; Izv. TINRO, vol. 62.

Nordenshel’d,-A.E. 1881. Puteshestvie vokrug Evropy i Sev. Azii (Expe-
dition around Europe and Northern Asia). St. Petersburg.

Obruchev, S.V. 1949. Massovaya gibel’kitov и beregov Argentiny (Whole-
sale death of whales on the coasts of Argentina). Priroda, no. 3.

Obruchev, S.V. 1951. Gibel’ chernykh del’finov (grind) и beregov Floridy
(Death of pilot whales on the coasts of Florida). Priroda, no. 8.

Ognev, S.I. 1935. Zveri SSSR i prilezhashchikh stran. Khishchnye
i lastonogie (Animals of the USSR and Neighboring Countries.
Carnivores and Pinnipeds). vol. 3. Moscow.

Okuneva, U.B. 1934. Chernomorskii i azovskii del’finy kak promysh-
lennoe syr’e (Black Sea and Azov Sea dolphins as exploitable raw
material). Tr. Vses. NII Rybn. Prom., vol. 3.

Ostroumov, A.G. 1960. Nerpy nakhodyat put’ k moryu (Ringed seals find
a way to the sea.) Priroda, no. 5.

Ostroumov, A.G. 1966. Nekotorye svedeniya 0 povedenii lastonogikh v
pribrezhnykh vodakh Kamchatki (Some information on the behavior
of pinnipeds in Kamchatka coastal waters). Izv. TINRO, vol. 8.

932

Ostroumov, А.С. 1967. Nekotorya svedeniya о chislennosti lastonogikh—
largi i akiby v pribrezhnykh vodakh Kamchatki (Some information on
the population of pinnipeds—large and ringed seals—in Kamchatka
coastal waters). Vopr. Geogr. Kamchatki, no. 5.

Ostroumov, М.А. 1929. K izucheniyu belukhi D. /eucas Pall. i ee promysla
у Eniseiskom zalive (Study of beluga D. Jeucas Pall. and its hunting
in Yenisey Bay). Tr. Sib. Nauch. Rybokhoz. St., vol. 14, no. 1.

Ostroumov, М.А. 1935. К sistematike i razmeshcheniyu belukhi
Evropeiskogo Severa i Karskogo morya (Taxonomy and range of
beluga in the European North and Kara seas). Za Rybn. Industr.
Severa, no. 11.

Ostroumov, М.А. 1936. Eshche о belomorskoi beluge (More about White
Sea beluga). Za Rybn. Industr. Severa, no. 3.

Panin, K.J. 1970. Raspredelenie i migratsii morskikh kotikov raznykh
stad у severnoi chasti Tikhogo okeana po rezul’tatam ikh mecheniya
(Range and migration of fur seals of different herds in the North
Pacific Ocean based on marking results). Izv. TINRO, vol. 70.

Panin, K.I. and С.К. Panina. 1968. Ekologiya 1 migratsii kotikov у zimne-
vesennii period v Yaponskom more (Ecology and migration of fur
seals in winter-spring in the Sea of Japan). Tr. VNIRO, vol. 68; Izv.
TINRO, vol. 62.

Panina, G.K. 1964. Pitanie kotikov v Yaponskom more (Feeding of fur
seals in the Sea of Japan). Izv. TINRO, vol. 54; Tr. TINRO, vol. 51.

Panina, G.K. 1966a. Pitanie kotikov v zapadnoi chasti Tikhogo okeana
(Feeding of western Pacific Ocean fur seals). Izv. TINRO, vol. 58.

Panina, G.K. 1966b. O pitanii sivucha i tyulenei na Kuril’skikh ostrovakh
(Feeding of Steller’s sea lion and seals on the Kuril Islands). Izv.
TINRO, vol. 58.

Panina, G.K. 1970. O pitanii morskikh kotikov v raione Komandorskikh
ostrovov (Feeding of fur seals in the region of Commander Islands).
Izv. TINRO, vol. 70.

Panina, G.K. 1972. Kharakter raspredeleniya kotikov, zimuyushchikh v
Yaponskom more (Nature of distribution of fur seals wintering in
the Sea of Japan). Tr. VNIRO, vol. 82; Izv. TINRO, vol. 80.

Panina, G.K. 1972. Pitanie i kormovaya baza kotikov v Yaponskom more
(Feeding and food base of fur seals in the Sea of Japan). Tr. VNIRO,
vol. 82; Izv. TINRO, vol. 80.

Panina, G.K. 1973. O vliyanii gidrologicheskikh uslovii na raspredelenie
kotikov у mestakh zimovki м vostochnogo poberezh’ya о. Khonsyu
(Effect of hydrological conditions on fur seal distribution in the win-
tering sites on the eastern coast of Honshu Island). In: [ssledovaniya po
Biologii Ryb i Promyslovoi Okeanografi, no. 4. TINRO, Vladivostok.

933

Pastukhov, V.D. 1961. Ob osennem i rannezimnem raspredelenii пегру
na Baikale (Autumn and early winter distribution of ringed seal in
Baikal). Izv. Sib. Otd. AN SSSR, no. 2.

Pastukhov, V.D. 1965a. К metodike kolichestvennogo ucheta baikal’-
skogo tyulenya (Method of counting Baikal seal). Sb. Morsk. Mlekop.
Nauka, Moscow.

Pastukhov, V.D. 1965b. Pitanie baikal’skogo tyulenya (Feeding of Baikal
seal). Tr. Limnol. In-ta, Sib. Otd. AN SSSR, vol. 6 (2b).

Pastukhov, V.D. 1966. Novye dannye o razmnozhenii baikal’skoi nerpy
(New data on the reproduction of Baikal ringed seal). Tez. Dokl. k 3
Vses. Soveshch. po Izuch. Morsk. Mlekop. Nauka, Moscow-Leningrad.

Pastukhov, V.D. 1967. Khozyaistvennoe znachenie baikal’skogo tyulenya
i ratsional’noe ispol’zovanie ego zapasov (Commercial importance
of Baikal seal and rational exploitation of its reserves). Tr. PINRO,
no. 21.

Pastukhov, V.D. 1969a. Nastuplenie polovoi zrelosti u samok baikal’skoi
nerpy (Onset of sexual maturity in female Baikal ringed seals). Sb.
Morsk. Mlekop. Nauka, Moscow.

Pastukhov, V.D. 1969b. O nekotorykh pokazatelyakh sostoyaniya stada
io promysle baikal’skoi nerpy (Some indices of herd condition and
hunting of Baikal ringed seal). Sb. Morsk. Mlekop. Nauka, Moscow.

Pastukhov, V.D. 1971. Uchet priploda baikal’skoi nerpy (Counting the
Baikal ringed seal offspring). Tr. AtlantNIRO, no. 39.

` Pastukhov, V.D. 1971. О pitanii baikal’skoi nerpy у usloviyakh eksper-
imenta (Feeding of Baikal ringed seal under test conditions). Tr.
AtlantNIRO, no. 39.

Perlov, A.S. 1970. Raspredelenie i chislennost’ sivuchei na lezhbischakh
Kuril’skikh ostrovov (Distribution and population of Steller’s sea
lion in Kuril Island rookeries). Izv. TINRO, vol. 70.

Perlov, A.S. 1972. Sroki nastupleniya polovoi zrelosti u sivuchei (Period
of onset of sexual maturity in Steller’s sea lions). Tr. VNIRO, vol. 82;
Izv. TINRO, vol. 80.

Pershin, $.У., A.S. Sokolov, and А.С. Tomilin. 1970. О reguliruemoi
spetsial’nymi sosudistymi organami uprugosti plavnikov del’finov
(Elasticity of dolphin fins controlled by special vascular organs).
DAN SSSR, vol. 190, no. 3.

Pikharev, С.А. 1939. Polosatyi tyulen’ (Banded seal). Vest. Б/у Fil. AN
SSSR, no. 33 (1).

Pikharev, G.A. 1941. Tyuleni yugo-zapadnoi chasti Okhotskogo morya
(Seals in the southwestern Sea of Okhotsk). Izv. TINRO, vol. 20.

Pikharev, С.А. 1946. О pitanii akiby (Feeding of ringed seal). Izv.
TINRO, vol. 22.

934

’РАКВагеу, С.А. 1948. О raspredelenii lakhtaka (Erignathus barbatus
Erxleben) у dal’nevostochnykh moryakh [Range of bearded seal
(Erignathus barbatus Erxleben) in Far East seas]. Izv. TINRO, vol. 28.

Pikharev, G.A. 1948. Promysel lastonogikh v moryakh Dal’nago Vostoka
(Hunting of Pinnipeds in Far East Seas). Vladivostok.

Pirozhnikov, P.L. 1937. Materialy k izucheniyu prois-khozhdeniya sev-
ernykh elementov v faune Kaspiiskogo morya (Data on the origin of
northern elements in Caspian Sea fauna). DAN SSSR, vol. 15.

Plekhanov, Р.Р. 1931. Kustarnyi tyulenii promysel у raione о. Morzhovtsa
(Bush hunting of seals in Morzhovets Island region). Sov. Sever
no. 6.

Plekhanov, Р.Р. 1931. Ostrov Morzhovets (Morzhovets Island). Sov. Sever
no. 6.

Plekhanov, P.P. 1933. Opredelenie vozrasta tyulenei (Determining the
age of seals). Byull. NIT Zool. MGU. Moscow.

Pleske, F.D. 1887. Kriticheskii obzor mlekopitayushchikh i ри
Kol’skogo poluostrova (Critical review of mammals and birds of
Kola Peninsula). Annexure to vol. 56 of Zap. Imp. Ak. Nauk, no. 1.
St. Petersburg.

Plotkin, V.A. 1966. Redkii sluchai mnogoplodiya u seivala (Rare instance
of multiple pregnancy in a sei whale). Zool. Zhurn., vol. 45, no. 2.

Polyakov, I.S. 1884. Otchet ob issledovaniyakh na o. Sakhaline, v
Yuzhno-Ussuriiskom krae i v Yaponii (An account of research
carried out on Sakhalin Island, in southern Ussurii territory, and in —
Japan). Annexure to vol. 48 of Zap. Imp. Ak. Nauk St. Petersburg.

Ponomareva, L.A. 1949. O pitanii planktonoyadnykh kitov Beringova
morya (Feeding of planktotrophic whales of the Bering Sea). DAN
SSSR, vol. 68, no. 2.

Popov, L.A. 1955. Napadenie gornostaya na tyulenya (Ermine attack on
seals). Priroda, no. 6.

Popov, L.A. 1958. Lezhbishcha morzhei па о. Peschanom (Walrus rook-
ery on Peschanyi Island). Priroda, no. 9.

Popov, L.A. 1959. Detnyi period zhizni khokhlacha Vostochnoi Gren-
landii (Juvenile period of hooded seal in eastern Greenland). Inf: Sb.
VNIRO, no. 7. Moscow.

Popov, Г.А. 1959. Osobennosti polovogo dimorfizma tyulenya-
khokhlacha (Features of sexual dimorphism in hooded seals). Tr.
I Konf. Molodykh Nauch. Sotr. Mosk. Morf. Lab. Moscow.

Popov, L.A. 1959. Raspredelenie lezhbisch morzhei v zapadnom sek-
tore Sovetskoi Arktiki v letne-osennii period (Distribution of wal-
rus rookeries in the western sector of the Soviet Arctic in summer-
autumn period). Inf. Sb. VNIRO, no. 7. Moscow.

935

Popov, L.A. 1960a. Materialy po biologii razmnozheniya morzha morya
Laptevykh (Data on reproduction biology of walruses of the Laptev
Sea). Byull. MOIP, Otd. Biol., vol. 65 (2).

Popov, L.A. 1960b. Sostoyanie beregovykh lezhbisch morzha v more
Laptevykh (Status of walrus coastal rookeries in the Laptev Sea).
Sb. Okhrana Poleznykh Zverei, no. 3. Vseros. Ob-vo Sod. Okhrane
Prir. 1 Ozel. Moscow.

Popov, L.A. 1960c. O nastuplenii polovoi zrelosti u khokhlachei (Onset
of sexual maturity in hooded seals). Tr. PINRO, по. 12.

Popov, L.A. 1960d. O regulirovanii promysla tyulenyakhokhlacha v Sev-
ernoi Atlantike (Regulation of hooded seal hunting in the North
Atlantic). Sb. Okhrana Poleznykh Zverei, no. 3.

Popov, L.A. 1960e. O periode razmnozheniya yanmaienskogo stada gren-
landskogo tyulenya (Reproduction period of Jan Mayen herd of harp
seals). Inf. Sb. VNIRO, no. 8. Moscow.

Popov, L.A. 1960f. Khokhlach Grenlandskogo morya (Cystophora cristata
Erxl.) novyi ob” ekt sovetskogo zveroboinogo promysla (obshch. mor-
fologiya, obr. zhizni, raspred. migrats. promysl. znachenie) [Hooded
seal (Cystophora cristata Erxl.) of the Greenland Sea, a new object of
Soviet animal hunting (general morphology, life style, range, migra-
tion, and economic importance)]. Avtoref. kand. diss., Moscow.

Popov, L.A. 1961. Materialy k obshchei morfologii khokhlacha Gren-
landskogo morya (Cystophora cristata Erxl.) [Data on the general
morphology of hooded seal (Cystophora cristata Erxl.) of the Green-
land Sea]. Tr. Soveshch. Ikht. Komis., по. 12.

Popov, L.A. 1963. Mechenye tyuleni (Marked seals). Priroda, no. 10.

Popov, L.A. 1965. Eksperimental’noe issledovanie nekotorykh voprosov
biologii detnogo perioda zhizni belomorskogo stada grenlandskogo
tyulenya (Experimental study of some aspects of the biology of the
juvenile period of White Sea herd of harp seal). Sb. Nauch.-tekhn.
Inf. VNIRO, no. 2.

Popov, L.A. 1966a. Ledovyi dreif biologov (Arctic drifting expedition of
biologists). Rybn. Khoz., nos. 10 and 11.

Popov, L.A. 1966b. Na Гаше $ tyulenyami (On ice with seals). Priroda,
no. 9.

Popov, L.A. 1967а. Snova na I’dine (Again on the ice). Priroda, по. 11.

Popov, L.A. 1967b. Sovremennyi zveroboinyi promysel v moryakh Sever-
noi Atlantiki (Modern hunting in North Atlantic seas). Probl. Severa,
по. №1:

Popov, L.A. 1969. О biologicheskom obosnovanii aerofotos”emok det-
nykh zalezhek grenlandskogo tyulenya v Belom more (Biological

936

rationale of aerial photographs of harp seal nurseries in the White
Sea). Tez. IV Vses. Soveshch. po Izuch. Morsk. Mlekop. Moscow.

Popov, L.A. and L. Pleshakov. 1970. Dreif za grenlandskim tyulenem
(reportazh) [Chasing harp seal (a report)]. Smena, no. 22, Novem-
ber.

Popov, L.A. and V.A. Potelov. 1963. Nekotorya zakonomernosti
raspredeleniya grenlandskogo tyulenya v Grenlandskom more
у period obrazovaniya linnykh zalezhek (Some principles of
distribution of harp seal in the Greenland Sea during the period
of formation of molting rookeries). Sb. Nauch.-issl. Rabot 1962 g.
PINRO. Arkhangel’sk.

Popov, L.N. 1939. Promyslovye mlekopitayushchie poberezh’ya Taimyr-
skogo poluostrova (Game mammals on coasts of Taimyr Peninsula).
Tr. NII Polyar. Zemled. Zhivotn. 1 Promysl. Khoz., no. 8.

Portenko, L.A. 1941. Fauna Anadyrskogo kraya. Ch. 3. Mlekopi-
tayushchie (Fauna of Anadyr territory. Pt. 3. Mammals). Tr. МИ
Polyar. Zemled. Zhivotn. т Promysl. Khoz., Ser. Promysl. Khoz.,
vol. 14.

Potelov, V.A. 1964. Vozrastnoi sostav khokhlacha i lysuna v Grenland-
skom more (Age composition of hooded and harp seals in the Green-
land Sea). Sb. Mater. Rybokhoz. Issl. Sev. Bass., no. 2.

Potelov, V.A. 1969. Sroki beremennosti, shchenki i raiony detorozh-
deniya morskogo zaitsa (Periods of gestation, parturition, and nurs-
ery regions of bearded seal). Tez. ГИ Vses. Soveshch. po Izuch. Morsk.
Mlekop. Moscow.

Potelov, V.A. 1969. Neobkhodimost’ zapreta promysla morskogo zaitsa
у Belom more (Need for banning the hunting of bearded seal in the
White Sea). Tez. Dokl. 8-ya Sessiya Uch. Sov. po Probl. Biologich. Res.
Bel. Morya i Vnutr. Vodoem. Evr. Severa. Petrozavodsk.

Potelov, V.A. 1969. Raspredelenie i migratsii morskikh zaitsev v Belom,
Barentsevom i Karskom moryakh (Range and migration of bearded
seals in the White, Barents, and Kara seas). Sb. Morsk. Mlekop.
Nauka, Moscow.

Potelov, У.А. and O.Ch. Mikhnevich. 1964. Zavisimost’ smeny okraski
volosyanogo pokrova ot vozrasta i dliny tela grenlandskogo tyulenya
(Dependence of hair coat color variation on age and body length
of harp seal). Sesstya Uch. Sov. “Teor. Osnovy Rats. Ispol’z. Rybn. 1
Nerybn. Res. Bel. Morya i Vnutr. Vodoem. Karelii”’ (Tez. Dokl.), no. 1.
Petrozavodsk.

Potelov, V.A. and O.Ch. Mikhnevich. 1967. Ob izmenchivosti okraski
grenlandskogo tyulenya (Color variation of harp seal). Tr. PINRO,
no. 21.

937

Potelov, V.A. and А.С. Beloborodov. 1969. Kharakteristika volosyanogo
pokrova morskogo zaitsa (Hair coat characteristics of bearded seal).
Tez. Dokl. IV Vses. Soveshch. po Izuch. Morsk. Mlekop. Moscow.

Potelov, V.A. and A.V. Yablokov. 1966. Novoe v morfologicheskoi
kharakteristike khokhlacha (New morphological characteristics of
hooded seal). Zool. Zhurn., vol. 45, no. 8.

Poznanskii, B. 1880. Vospominaniya ob okhote v Chernomorskom
okruge (Recollections on hunting in the Black Sea region). Priroda
i Okhota, vol. 3, books 8 and 9.

Pravdin, I.F. 1928. Ocherk zapadnokamchatskogo rybolovstva у svyazi $
obshchimi voprosami dal’nevostochnoi promyshlennosti (Résumé of
western Kamchatka fishing industry in relation to general aspects of
the Far East industry). Izv. Tikh. Nauch.-Prom. St., vol. 1.

Prigorovskii, N.I. and Г.Г. Breitfus. 1906. Deyatel’nosti promyslovogo
telegrafa v 1903 godu (Activity of trade telegrams in 1903). Eksp.
dlya Nauch.-prom. Issl. u Beregov Murmana. St. Petersburg.

Prigorovskii, N.I. and L.L. Breitfus. 1912. Khod promysla na Murmane
v 1905 g. po svedeniyam promyslovogo telegrafa (Industrial trend in
Murman in 1905 from information in trade telegrams). Tr. Murm.
Nauch.-prom. Eksp. 1905 g. St. Petersburg.

Prigorovskii, М.Г. and L.L. Breitfus. 1915. Khod promysla na Murmane
1 1906 g. po svedeniyam “promyslovogo telegrafa” (Industrial trend
in Murman in 1906 from information in trade telegrams). Tr. Murm.
Nauch. prom. Eksp. 1906 g. St. Petersburg.

Probatov, А.М. 1933. Materialy po nauchno-promyslovomu obsle-
dovaniyu Karskoi guby i reki Kary (Data of Scientific-Industrial
Research in Kara Gulf and Kara River). Moscow.

Probatov, А.М. 1934. Itogi rabot Karskoi ekspeditsii 1933 рода 1
perspektivy khozyaistvennogo osvoeniya Karskogo morya (Results
of the work of 1933 Kara expedition and prospects of commercial
exploitation of the Kara Sea). Khozyaistvo Severa, no. 3.

Provorov, N.V. 1957. K voprosu o biologii i promysle belomorskoi
belukhi v Kandalakshskom zalive Belogo morya (Biology and
hunting of White Sea beluga in Kandalakshsk Bay of the White
Sea). Tr. Arkt. In-ta, vol. 205.

Puzanoy, I.J. 1929. Chernoe more (svedeniya о tyulene-monakhe) [Black
Sea (Information on Monk Seal)]. Sevastopol’.

Rakhilin, V.K. 1968. K biologii serogo tyulenya (Halichoerus grypus)
[Biology of Atlantic gray seal (Halichoerus grypus)]. Sb. Tr. Zool.
Muzeys MGU, vol. 10.

Razumovskii, V.I. 1933. Vesennii zveroboinyi promysel (Spring Hunting
of Animals). Vladivostok.

938

Roganov, А.М. 1930. Gagachii boi tyulenya vesnoi 1929 рода (Killing
of eiders by seals in spring 1929). Tr. Astrakh. Nauch. Rybokhoz. St.,
vol. 7, no. 2.

Roganov, A.N. 1931. Kaspiiskii tyulen’ i ego promysel (Caspian seal and
its hunting). Tr. Volgo-Kasp. Nauch. Rybokhoz. St., vol. 7, no. 4.
Roganov, A.N. 1932. Predvaritel’nye itogi issledovanii kaspiiskogo tyu-
lenya i ego promysia (Preliminary results of research on Caspian seal
and its hunting). Byull. Vsekasp. Nauch. Rybokhoz. Eksp., nos. 5 and

6

Romanenko, E.V., A.G. Tomilin, and B.A. Artemenko. 1965. K voprosu
о zvukoobrazovanii 1 napravlennosti zvukov и del’finov (Sound
generation and sound directions in dolphins). Sb. Bionika. Nauka,
Moscow.

Romer, A.Sh. 1939. Paleontologiya pozvonochnykh (Vertebrate Palaeon-
tology). Moscow-Leningrad.

Rovnin, A.A. 1969. Raspredelenie krupnykh kitoobraznykh v tropich-
eskoi chasti Tikhogo okeana (Range of large cetaceans in the trop-
ical Pacific Ocean). Sb. Morsk. Mlekop. Nauka, Moscow.

Rozanov, M.P. 1931. Promysel morskogo zverya na Chukotskom polu-
ostrove (Hunting of marine animals on Chukchi Peninsula). Sov.
Sever, no. 6.

Rudakov, P.A. 1936. Predvaritel’nye itogi zimne-vesennei ekspeditsii po
izucheniyu belomorskogo stada grenlandskogo tyulenya v 1936 godu
(Preliminary results of winter-spring expedition in 1936 to study the
White Sea herd of harp seal). Byull. Arkt. In-ta, nos. 8 and 9.

Rundan, А.М. 1936. Kratkie biologicheskie nablyudeniya у yugo-
zapadnoi chasti Karskogo morya (Brief biological observations in
the southwestern Kara Sea). Byull. Arkt. In-ta, no. 12.

Rutilevskii, L.G. 1939. Promyslovye mlekopitayushchie poluostrova
Chelyuskin 1 proliva Vil’kitskogo (Game mammals of Chelyuskin
Peninsula and Vil’kits Strait). Tr. In-ta Polyar. Zemled., Zhivot. i
Promysl. Khoz., no. 8.

Rutilevskii, G.A. 1962. Zhivotnyi mir (Animal world). In: Severnaya
Yakutiya. Tr. Arkt. 1 Antarkt. In-ta, vol. 236.

Rutilevskdi, G.L. and S.M. Uspenskii. 1957. Fauna mlekopitayushchikh
i ptits Tsentral’noi Arktiki (Fauna of central arctic mammals and
birds). Tr. Arkt. In-ta, vol. 205.

Sal’nikov, N.E. 1959. Novi dani pro tyulenya-monakha v Chernomu mori
(New data on the monk seal in the Black Sea). Naukovi Zap. Odes’koi
Biol. St., no. 1.

Sal’nikov, N.E. 1953. Pitanie finvala i sinego kita v Antarktike (Feeding
of fin whale and blue whale in Antarctica). Tr. VNIRO, vol. 25.

939

Sal’nikov, М.Е. 1956. Nekotorye уоргозу 6101021 razmnozheniya finvala
у sinego kita Antarktiki (Some aspects of reproduction biology in
fin whale and blue whale in Antarctica). Tr. Balt. In-ta Ryb. Khoz. 1
Okean. (BaltNIRO), no. 2.

Sal’nikov, М.Е. 1967. Lastonogie (Pinnipedia) [Pinnipeds (Pinnipedia)].
In: Biologiya Severo-zapadnoi Chasti Chernogo Morya. Kiev.

Samofalov, P.E. 1930. Promysel tyulenya v Mangyshlakskom raione v
1928 - 1929 godakh (Seal hunting in Mangyshlak region in 1928 and
1929). Tr. Astr. Nauch. Rybkhoz. St., vol. 7, no. 2.

Samorodov, A.V. 1939. K faune mlekopitayushchikh zemli olyutorskikh
koryakov (Mammalian fauna of Olyutorsk Koryaks). Sb. Tr. Gos.
Zool. Muzeya pri MGU, vol. 5.

Sazhinov, E.G. 1970. Nastuplenie polovoi i fizicheskoi zrelosti u sinikh
kitov-pigmeev (Balaenoptera musculus brevicauda) [Onset of sexual
and physical maturity in great blue whales (Balaenoptera musculatus
brevicauda)]|. Tr. AtlantNIRO, no. 25.

Sdobnikov, V. 1932. O setnom promysle grenlandskogo tyulenya na Mur-
manskom beregu (Catching of Greenland seal in nets off the coast
of Murman). Tr. Ат. In-ta, vol. 3, no. 2.

Shastin, L. 1936. Rekognostsirovochnye raboty v Ust’-Taimyrskoi gube
i po reke Nizhnei Taimyre (Exploratory work in Ust’-Taimyr Gulf
and along the lower Taimyr River). Byull. Arkt. In-ta, no. 12.

Shchupakov, [.С. 1936. Parazitofauna kaspiiskogo tyulenya. К vopr. о
parazitofaune reliktov (Parasite fauna of Caspian seal. Parasite fauna
of relicts). Uch. Zap. Leningr. Gos. Un-ta, Ser. Biol., vol. 7, no. 3.

Shepeleva, V.K. 1963. O sekretornoi funktsii zheludka u tyulenei (Secre-
tory function of the stomach in seals). Sb. Nauch.-issl. Rabot 1962 g.
(Grenlandskii Tyulen’ i Khokhlach). Arkhangel’sk.

Shepeleva, V.K. 1971. Prisposoblenie tyulenei k obitaniyu v Arktike
(Adaptation of Seals to Arctic Life). Novosibirsk.

Shevchenko, V.I. 1970. Zagadka “belykh shramov” na tele kitov (Con-
jecture on “white scars” on whale bodies). Priroda, no. 6.

Shevchenko, V.I. 1971. К voprosu о proiskhozhdenii “belykh shramov”
na tele kitov (On the origin of “white scars” on whale bodies). Tr.
AtlantNIRO, no. 39.

Shuleikin, V., V. Luk’yanova, and I. Stas’. 1937. Gidrodinamicheskie
kachestva губ 1 del’finov (Hydrodynamic qualities of fish and dol-
phins). Izv. AN SSSR, Ser. Geofiz., vol. 14.

Shustov, А.Р. 1965. О vliyanii promysla na sostoyanie beringovomorskoi
populyatsii krylatki (Effect of hunting on the population status of
ribbon seal in the Bering Sea). Izv. TINRO, vol. 59.

940

Shustov, А.Р. 1965a. Pitanie krylatki у Beringovom more (Feeding of
ribbon seal in the Bering Sea). Izv. TINRO, vol. 59.

Shustov, A.P. 1965b. Nekotorye cherty biologii i temp razmnozheniya
krylatki (Histriophoca fasciata) [Some biological features and growth
tempo of ribbon seal (Histriophoca fasciata)}. Izv. TINRO, vol. 59.

Shustov, A.P. 1965c. Nekotorye osobennosti gel’mintofauny krylatki v
Beringovom more (Some characteristics of helminth fauna of ribbon
seal in the Bering Sea). Izv. TINRO, vol. 59.

Shustov, A.P. 1967. K voprosu o ratsional’nom ispol’zovanii zapasov
lastonogikh Beringova morya (Rational exploitation of pinniped
reserves of the Bering Sea). Probl. Severa, no. 11.

Shustov, А.Р. 1969а. K voprosu о sutochnoi dinamike zalezhek tyulenei
v Okhotskom more (Diurnal dynamics of seal rookeries in the Sea of
Okhotsk). Tez. Dokl. IV Vses. Soveshch. po Izucheniyu Morsk. Mlekop.
Moscow.

Shustov, A.P. 1969b. Opyt kolichestvennogo aerovizual’nogo ucheta tyu-
lenei v severo-zapadnoi chasti Beringova morya (Quantitative aerial
census of seals in the northwestern Bering Sea). Sb. Morsk. Mlekop.
Nauka, Moscow.

Shustov, А.Р. 1969c. О vliyanii promysla na sostoyanie zapasov
beringovomorskoi populyatsii krylatki (Effect of hunting on the
reserves Status of Bering Sea ribbon seal population). Sb. Morsk.
Mlekop. Nauka, Moscow.

Shustov, A.P. 1969. Otnositel’nyi pokazatel’ i vozmozhnye faktory smert-
nosti beringovomorskoi krylatki (Relative index and possible reasons
for the mortality of Bering Sea ribbon seal). Sb. Morsk. Mlekop.
Nauka, Moscow. ;

Shustov, A.P. and A.V. Yablokov. 1967. Sravnitel’no morfologicheskaya
kharakteristika grenlandskogo 1 polosatogo tyulenei (Comparative
morphological characteristics of harp and ribbon seals). Tr. PINRO,
ПО: 121.

Sklabinskii, М. 1891. Svedeniya о tyulen’ei promyshlennosti na Kaspi-
iskom more za poslednie 25 let (Information on seal industry in the
Caspian Sea for the last 25 years). Rybn. Delo., no. 6.

Skryabin, A.S. 1958. Zoogeograficheskaya kharakteristika gel’mintofauny
morskikh mlekopitayushchikh Boreo-Patsificheskoi podoblasti (Zoo-
geographic characteristics of the helminth fauna of marine mammals
of the Boreo-Pacific subregion). Гу. Krym. Ped. In-ta, vol. 31.

Skryabin, A.S. 1959. Novye vidy gel’mintov ot morskikh mlekopi-
tayushchikh Tikhogo okeana i dal’nevostochnykh morei (New types
of helminths from marine mammals of the Pacific Ocean and Far
East seas). Izv. Krym. Ped. In-ta, vol. 34.

941

Skryabin, A.S. 1960. Gel’mintofauna morskikh mlekopitayushchikh
Tikhogo okeana i dal’nevostochnykh morei (Helminth fauna of
marine mammals of the Pacific Ocean and Far East seas). Avtoref.
diss., Simferopol’.

Skryabin, A.S. 1969. Novyi vid trematod roda Odmogaster (Notocoty-
lidae)—parazit usatykh kitov [New species of trematode of the
genus Odmogaster (Notocotylidae) parasitizing baleen whales]. Zool.
Zhurn., vol. 48, no. 2.

Skryabin, A.S. 1970. Novyi vid skrebnei Bolbosoma tuberculata sp. n.
(sem. Polymorphydae Meyer, 1931)—parazit kitov [New species of
acanthocephalans, Bolbosoma tuberculata sp. n. (fam. Polymorphy-
dae Meyer, 1931), parasitizing whales]. Parazitologtya, vol. 4, по. 4.

Skryabin, К.Г. 1950. Trematody zhivotnykh i cheloveka (Trematodes of
Animals and Man). vol. 4. Moscow-Leningrad.

Skvortsov, I. 1927. Setnoi promysel morskogo zverya na Murmane
(Catching of marine animals in nets in Murman). ВуиП. Rybn. Khoz.,
nos. 11 and 12.

Sleptsov, M.M. 1940a. Opredelenie vozrasta del’fina (Determining the
age of dolphins). Byull. MOIP, Otd. Biol., vol. 49 (2).

Sleptsov, М.М. 19406. Ob osobennostyakh rozhdeniya i pitaniya
detenyshei chernomorskogo del’fina (Birth and feeding character-
istics of Black Sea dolphin calves). Zool Zhurn., vol. 19, no. 2.

Sleptsov, M.M. 1940c. Razvitie kostnogo cherepa v ontogeneze i filo-
geneze (Growth of bony skull in ontogeny and phylogeny). DAN
SSSR, vol. 28, no. 4.

Sleptsov, M.M. 1940d. O prisposoblenii k plavaniyu lastonogikh (Adap-
tation of pinnipeds to swimming). Zool. Zhurn., vol. 19, no. 3.

Sleptsov, М.М. 1941. О biologii razmnozheniya chernomorskogo
(Reproduction biology of Black Sea dolphin). Zool. Zhurn., vol. 20,
nos. 4 and 5.

Sleptsov, М.М. 1943. О biologii razmnozheniya lastonogikh Dal’nego
Vostoka (Reproduction biology of pinnipeds in the Far East). Zool.
Zhurn., vol. 2.

Sleptsov, M.M. 1948. Giganty okeana (Ocean Giants). Vladivostok.

Sleptsov, М.М. 1949. Novye dannye po razmnozheniyu dal’nevostoch-
nykh lastonogikh (New data on the reproduction of Far East
pinnipeds). Izv. TINRO, vol. 31.

Sleptsov, M.M. 1950. O biologii dal’nevostochnogo sivucha (Biology of
Far East Steller’s sea lion). Izv. TINRO, vol. 32.

Sleptsov, M.M. 1950. Novye dannye o raspredelenii samok kashalotov
(New data on the distribution of female sperm whales). Izv. TINRO,
vol. 32.

942

Sleptsov, М.М. 1952. Kitoobraznye dal’nevostochnykh тоге! (Cetaceans
of the Far East seas). Izv. TINRO, vol. 38.

Sleptsov, M.M. 1955a. Kitoobraznye dal’nevostochnykh morei (Cetaceans
of the Far East seas). Vladivostok.

Sleptsov, M.M. 1955b. Osobennosti razvitiya kitoobraznykh na rannikh
embrional’nykh stadiyakh (Growth characteristics of cetaceans in the
early embryonal stages). Tr. In-ta Okeanol. AN SSSR, vol. 18.

Sleptsov, М.М. 1961. Nablyudeniya nad melkimi kitoobrazhymi у
dal’nevostochnykh moryakh i severo-zapadnoi chasti Tikhogo
okeana (Observations on small cetaceans in the Far East seas and
northwestern Pacific Ocean). Tr. In-ta Okeanol. AN SSSR, no. 34.

Smirnov, A.F. 1954. Rybokhozyaistvennoe znachenie vnutrennikh vodoe-
mov Karelo-Finskoi SSR (Importance of the water bodies of Karelo-
Finsk SSR for fishing). Mat. Soveshch. po Probl. Povysh. Rybn. Prod.
Vnutr. Vodoemov K.-F. SSR. Petrozavodsk.

Smirnov, М.А. 1903. О morskom zverinom promysle na russkikh sudakh
(Animal hunting at sea by Soviet craft). Eksp. dlya Nauch.-prom. Issl.
u Beregov Murmana. St. Petersburg.

Smirnov, N.A. 1904. Otchet o deyatel’nosti Mangyshlakskoi stantsii
Казризко! ekspeditsii (An account of the activity of Mangyshlak
station of the Caspian expedition). Tr. Kasp. Eksp. 1904 g., vol. 1.

Smirnov, М.А. 1907. О zimnem tyulen’em promysle па Kaspiiskom more
(Winter Hunting of Seals in the Caspian Sea). Department Zem-
ledeliya. St. Petersburg.

Smirnov, N.A. 1908. Ocherk russkikh lastonogikh (Résumé of Russian
pinnipeds). Zap. Imp. AN, vol. 23, no. 4.

Smirnov, N.A. 1911. Materialy k izucheniyu rybnoi promyshlennosti v
Amakhtonskom zalive (Data on the fishing industry in Amakhtonsk
Bay). Rybn. Prom. DV, no. 4.

Smirnov, N.A. 1912. O rasprostranenii Pinnipedia v Severnom polusharii
(Distribution of Pinnipedia in the Northern Hemisphere). Zap.
Novoros. Ob-va Estestv., vol. 39.

Smirnov, М.А. 1924. Tyulen’ (Phoca groenlandica) i lov treski na Murmane
i Finmarkene [Seal (Phoca groenlandica) and cod fishing in Murman
and Finmark]. Izv. Ord. Prikl. Ikht. Gos. In-ta Opytn. Agr., vol. 2.

Smirnov, N.A. 1927b. Issledovaniya nad promyslom belomorskogo tyu-
lenya. Noveishie nablyudeniya nad belomorskim lysunom (Study of
White Sea seal hunting. Latest observations on White Sea harp seal).
Izv. О. Prikl. Ikht. 1 Nauchn.-prom. Issl., vol. 6, no. 1.

Smirnov, N.A. 1927c. Grenlandskii tyulen’, ili lysun (Harp seal). Priroda,
no. 5.

943

Smirnov, М.А. 19274. Biologicheskie otnosheniya nekotorykh lastonogikh
ko I’dam (Biological affinity of some pinnipeds for ice). Sb. V Chest’
Knipovicha. Moscow-Leningrad.

Smirnov, N.A. 1928. Zamechaniya ob osnovnykh printsipakh ischisleniya
normy uboya lastonogikh (Notes on the basic principles of determin-
ing the killing norms of pinnipeds). Гу. Оша. Ikhtiol. 1 Nauch.-prom.
Issl. Gos. In-ta Opytn. Agr., vol. 8.

Smirnov, М.А. 1929. Opredelitel’ lastonogikh (Pinnipedia) Evropy i Sev-
ernoi Azii [Key to pinnipeds (Pinnipedia) of Europe and northern
Asia]. Izv. Otd. Prikl. Ikht. г Nauch.-prom. Issl., vol. 9, no. 3.

Smirnov, N.A. 1931. Itogi issledovanii kaspiiskogo tyulenya i ego
promysla v 1929 g. (Results of studying Caspian seal and its hunting
in 1929). Izv. Len. Nauch.-issl. Ikht. In-ta, vol. 12, no. 1.

Smirnov, М.А. 1935. Lastonogie (Pinnipeds). In: Zveri Arktiki. Moscow.

Smirnov, N.A. 1935. Morskie zveri arkticheskikh morei (Marine animals
of Arctic seas). In: Zvert Arktiki. Moscow.

Smirnov, N.A. and K.K. Chapskii. 1932. K izucheniyu vozrastnykh priz-
nakov na cherepe tyulenei podroda Pusa (A study of the age-related
features of the skull of seals of the subgenus Pusa). Izv. Leningr. N.-I.
Ikht. In-ta, vol: 13, no. 2.

Sokolov, A.S. 1956. Materialy po vozrastnoi izmenchivosti razmerov tela
i cherepa ladozhskogo tyulenya (Data on age-related variation of
body and skull dimensions of Ladoga seal). Uch. Zap. Len. Gos. Ped.
In-ta, vol. 19.

Sokolov, A.S. 1958. Materialy po biologii ladozhskogo tyulenya (Data on
the biology of Ladoga seal). Uch. Zap. Len. Gos. Ped. In-ta, vol. 179.

Sokolov, A.S. 1972. Razmernaya kharaKteristika skeleta. konechnostei
severnogo morskogo kotika 1 sivucha (Size features of the limb skele-
ton of northern fur seal and Steller’s sea Поп). Tr. VNIRO, vol. 82;
Izv. TINRO, vol. 80.

Sokolov, A.S., G.M. Kosygin, and E.A. Tikhomirov. 1966. Nekotorye
dannye o vese vnutrennikh organov lastonogikh Beringova morya
(Some data on the visceral weight of pinnipeds of the Bering Sea).
Izv. Tikh. NIT Rybn. Khoz. i Okean., vol. 58.

Sokolov, A.S. et al. 1969. Spravochnye pokazateli po kharakteristike
vnutrennikh organov tikhookeanskikh lastonogikh (Standard indices
of visceral characteristics of Pacific pinnipeds). Sb. Morsk. Mlekop.
Nauka, Moscow.

Sokolov, V.E. 1953. K biologii razmnozheniya obyknovennogo Cher-
nomorskogo del’fina (Reproduction biology of common Black Sea
dolphin). Tr. Mosk. Pushno-Mekh. In-ta, vol. 4.

944

Sokolov, У.Е. 1954. Materialy ро biologii га’тпо7вешуа cher-
nomorskogo del’fina (Data on the reproduction biology of Black
Sea dolphin). Byull. MOIP, Nov. Ser., Ош. Biol., vol. 59, no. 1.

Sokolov, V.E. 1955. Struktura kozhnogo pokrova nekotorykh kito-
obraznykh (Structure of the integument of some cetaceans). Byull.
MOIP, vol. 60, no. 6.

Sokolov, V.E. 1961. Opredelenie stadii polovogo tsikla samok del’fina-
belobochki Delphinus delphis L. metodom vlagalishchnykh mazkov
(Determining the stage of sexual cycle of female common dolphin
Delphinus delphis L. by the vaginal smear method). [kht. Komis., Tr.
Soveshch., AN SSSR, no. 12.

Sokolov, V.E. and T. Kasuya. 1969. O promysle del’finov v Yaponii (Dol-
phin hunting in Japan). Nauch. Dokl. Vyssh. Shk., Biol. Nauki, no. 1.

Sokolov, V.E. and N.S. Stepanov. 1971. Opyt radioproslezhivaniya
morskikh kotikov (Radio location of fur seals). Tr. AtlantNIRO,
no. 39.

Soldatov, V.K. 1902. Otchet o rabotakh, proizvedennykh v ekspeditsii
nauchno-promyslovykh issledovanii Murmana $ 14 dekabrya 1901
g. do 19 maya 1902 g. (An account of the investigations carried
out in the scientific research expedition in Murman from December
14, 1901 to May 19, 1902). Eksp. dlya Nauch.-prom. 1551. и Beregov
Murmana. St. Petersburg.

Sorokin, 5.М. 1957. О zimnikh skopleniyakh ladozhskogo tyulenya (Win-
ter collections of Ladoga seals). Nauch.-tekh. Byull. VNIORKA, по. 5.

Sorokin, S.M. 1970. Nekotorye dannye 0 popadanii ladozhskogo tyulenya
v rybolovnye orudiya (Some data on Ladoga seals trapped in fishing
gear). Sb. Rybokhoz. Izuch. Vnutr. Vodoemov, по. 5.

Sosnovskii (ed.). 1911. Materialy po issledovaniyu Novoi Zemli (Data
on Research on Novaya Zemlya), no. 2. St. Petersburg.

Suvorov, E.K. 1912. Komandorskie ostrova i pushnoi promysel na nikh
(Commander Islands and Fur Seal Hunting on Them). St. Peters-
burg.

Suvorov, Е.К. 1914. О promysle morzha i kita na Chukotskoi zemle
(Walrus and whale hunting on Chukchi land). Mater. k Pozn. Russk.
Rybol., vol. 3, no. 5.

Suvorov, E.K. 1927. Predvaritel’nyi otchet cheshskoi ekspeditsii, 1925
g. (Preliminary report of Cheshsk expedition in 1925). Tr. In-ta po
Izuch. Severa, no. 34.

Surkov, S.S. 1957. Raspredelenie i zapasy lysuna v Belom more (Range
and reserves of harp seal in the White Sea). Murman.

Surkov, 5.5. 1959. О rezul’tatakh zveroboinogo promysla i provedenie
aerofotos”emok detnykh zalezhek belomorskogo lysuna у 1959 р.

945

(Results of animal hunting and aerial photography of White Sea
harp seal nurseries in 1959). Nauch.-tekhn. Byull. PINRO, no. 2 (10).

Surkov, S.S. 1960. Dannye po biologii i promyslu yanmaienskogo lysuna
i khokhlacha у zapadnykh I’dakh (Data on the biology and hunting
of Jan Mayen harp seal and hooded seal on western ice). Tr. PINRO,
mo. 12.

Surkov, S.S. 1963. К voprosu о metodike opredeleniya chislennosti 1
normakh vyboya belomorskogo lysuna Method of determining the
population and killing norms of White Sea harp seal). 77. PINRO,
ПО. 15.

Surkov, 5.5. 1966. Morskie mlekopitayushchie, vstrechayushchiesya и
beregov Kol’skogo poluostrova (Marine mammals encountered off
the coasts of Kola Peninsula). Sb. Кубу Murmanskoi Obl. Murman.

Surkov, S.S. and R.Sh. Khuzin. 1959. Itogi belomorskoi zveroboinoi eks-
peditsii i nekotorye rezul’taty issledovanii lysuna v 1959 g. (Results
of White Sea animal hunting expedition and some results of studying
harp seal in 1959). Nauchn.-tekhn. Byull. PINRO, no. 4 (8).

Sushkina, N.N. 1956. Narvaly (Narwhals). Priroda, no. 5.

Sverdrup, G.U. 1930. Plavanie na sudne “Mod” v vodakh morei
Laptevykh i Vostochno-Sibirskom (Sailing on the ship “Mod” in
Laptev and East Siberian waters). Mater. Komis. po Izuch. Yakut.
ASSR, no. 30.

Sviridov, N.S. 1954. Opyt ucheta chislennosti i raspredeleniya nerpy na
ozere Baikal s pomoshch’yu samoleta (Aerial census and distribution
of ringed seal in Lake Baikal). Izv. Irk. S/kh In-ta, no. 6.

Tambovtsev, B.M. 1966. Massovye podkhody detenyshei grenlandskogo
tyulenya k beregam Belogo morya (En masse arrival of harp seal
calves on coasts of the White Sea). Rybn. Khoz., no. 12.

Tarasevich, M.N. 1951. Vozrastno-polovaya struktura kosyakov del’fina-
belobochki (Age and sex structure of common dolphin herds). Tr.
Vses. Gidrobiol. Ob-va, vol. 3.

Tarasevich, M.N. 1958. Raspredelenie del’fina-belobochki v Chernom
more v teplyi period vremeni (Distribution of common dolphin in
the Black Sea in the warm season). Inf. Sb. VNIRO, по. 2.

Tarasevich, M.N. 1958b. Pitanie del’fina-belobochki v Chernom more v
teplyi period vremeni (Feeding of common dolphin in the Black Sea
in the warm season). Inf. Sb. VNIRO, no. 3.

Tarasevich, M.N. 1960a. Zakonomernosti podkhoda belukhi k beregam
severnykh morei (Beluga arrival patterns on northern sea coasts).
Inf. Sb. VNIRO, no. 8.

Tarasevich, M.N. 1960b. Kharakteristika podkhoda belukhi k beregam
(Beluga arrival characteristics on coasts). Tr. PINRO, по. 12. .

946

Tarasevich, М.М. 1963. К biologii morskogo zaitsa (Biology of bearded
seal). Tr. In-ta Okeanol. AN SSSR, vol. 71.

Tarasevich, M.N. 1963. Materialy po pitaniyu kashalotov severnoi chasti
Kuril’skikh vod (Data on feeding of sperm whales in northern Kurill
waters). Tr. In-ta Okeanol. AN SSSR, vol. 71.

Tarasevich, M.N. 1965. Raspredelenie kashalotov v severnoi chasti
Kuril’skikh vod v 1959-1961 gg. (Sperm whale distribution in
northern Kuril waters in 1959-1961). Sb. Morsk. Mlekop. Nauka,
Moscow.

Tarasevich, M.N. 1967a. O strukture gruppirovok kitoobraznykh. Soob-
shch. 1. Struktura gruppirovok samtsov kashalotov (Group structure
of cetaceans. Report 1. Structure of groups of male sperm whales).
Zool. Zhurn., vol. 46, no. 1.

Tarasevich, М.М. 19676. О strukture gruppirovok kitoobraznykh.
Soobshch. 2. Struktura gruppirovok finvalov (Group structure of
cetaceans. Report 2. Structure of groups of fin whales). Zool. Zhurn.,
vol. 46, no. 3.

Tarasevich, M.N. 1968. Pishchevye svyazi kashalotov v severnoi chasti
Tikhogo okeana (Trophic relations of sperm whales in the North
Pacific Ocean). Zool. Zhurn., vol. 47, no. 4.

Terebenin, У.Е. 1930. Zimnii boi $ promysla (Winter hunting). Gi.
Lokatin. Soveshch. po Issled. Kasp. Туш. 1 Ego Prom. Astrakhan.
Tikhenko, T.A. 1914. Ob ostrovakh Tyulen’em i Kuril’skikh (Tyulen’ and

Kuril Islands). Mat. k Pozn. Rus. Rybol., vol. 3, no. 5.

Tikhomirov, Е.А. 1959. К voprosu о pitanii sivucha teplokrovnymi zhiv-
otnymi (Intake of warm-blooded animals by Steller’s sea lion). Izv.
TINRO, vol. 47.

Tikhomirov, E.A. 1961. Rasshiryat’ dobychu largi (Enlarging the catch
of larga seal). Rybn. Prom. DV, no. 8.

Tikhomirov, Е.А. 1961. Raspredelenie 1 migratsii tyulenei у vodakh Dal’-
nego Vostoka (Seal distribution and migration in Far East waters).
Tr. Soveshch. Ikht. Komis., no. 12.

Tikhomirov, E.A. 1964a. O raspredelenii i promysle sivucha v
Beringovom more i sopredel’nykh raionakh Tikhogo okeana
(Steller’s sea lion distribution and hunting in the Bering Sea and
adjoining Pacific Ocean regions). Tr. VNIRO, vol. 53; Izv. TINRO,
vol. 52.

Tikhomirov, E.A. 1964b. Vstrechaemost’ kotikov v Okhotskom more
(Fur seals encountered in the Sea of Okhotsk). Izv. TINRO, vol. 51.

Tikhomirov, E.A. 1964c. Promysel morzha i puti okhrany ego populyat-
sii (Walrus hunting and methods of conserving its population). Sb.
Okhrana Prirody na Dal’nem Vostoke, по. 2. Vladivostok.

947

Tikhomirov, Е.А. 1964. О raspredelenii i biologii lastonogikh Beringova
morya (Pinniped distribution and biology in the Bering Sea). Izv.
TINRO, vol. 52; Tr. VNIRO, vol. 53.

Tikhomirov, Е.А. 1966. Lastonogie—ob’ekt sudovogo zveroboinogo
promysla (Pinnipeds—targets of hunting using craft). In: Dal’nevost.
Laston. Vladivostok.

Tikhomirov, E.A. 1966a. Nekotorye dannye o raspredelenii i biologii
largi v Okhotskom more v letne-osennii period i organizatsiya ee
promysla (Some data on larga seal distribution in the Sea of Okhotsk
in summer-autumn and organization of its hunting). Ру. TINRO,
vol. 58.

Tikhomirov, E.A. 1966b. Opredelenie vidov dal’nevostochnykh tyulenei
s samoleta (Aerial identification of species of Far East seals). Izv.
TINRO, vol. 58.

Tikhomirov, Е.А. 1966c. О razmnozhenii tyulenei semeistva Phocidae
severnoi chasti Tikhogo okeana (Reproduction of seals of the family
Phocidae in the North Pacific Ocean). Zool. Zhurn., vol. 15, no. 2.

Tikhomirov, E.A. 1968. Rost tela i razvitie organov razmnozheniya
severotikhookeanskikh nastoyashchikh tyulenei (Body growth and
development of reproductive organs of North Pacific true seals). Tr.
VNIRO, vol. 68; Izv. TINRO, vol. 62.

Tikhomirov, E.A. 1970. K voprosu razmnozheniya tyulenei sem. Pho-
cidae (Reproduction of seals of the family Phocidae). Гу. TINRO,
vol. 70.

Tikhomirov, E.A. 1970. Sluchai poedaniya largoi ptits (A case of larga
seal consuming birds). Izv. TINRO, vol. 70.

Tikhomirov, Е.А. and С. Козурш. 1966. Perspektivy promysla
lastonogikh v Beringovom more (Prospects of hunting pinnipeds in
the Bering Sea). Rybn. Khoz. SSSR, no. 9.

Timoshenko, Yu.K. 1963. K voprosu o pitanii grenlandskogo tyulenya
(Food of harp seal). Sb. Nauch.-issl. Rabot 1962 g. (Grenlandskii
Tyulen’ i Khokhlach). Arkhangel’sk.

Timoshenko, Yu.K. 1964. K razlichiyam v biologii grenlandskogo tyu-
lenya belomorskogo in n’yufaundlendskogo stad (Biological differ-
ences between harp seals of White Sea and Newfoundland herds).
Tez. Dokl. Sessii Uch. Sov. Karel. О. GosNIORKh, по. 1. Petroza-
vodsk.

Timoshenko, Yu.K. 1968. Novye dannye k biologii razmnozheniya gren-
landskogo tyulenya belomorskogo stada (New data on the repro-
duction biology of harp seal herd of the‘White Sea). Tez. Dokl. VII
5ез5й Uch. Sov. Sb. Biol. Кез. Bel. Morya i Vnutr. Vodoem. Karelii.
Petrozavodsk.

948

Timoshenko, Yu.K. 1969. Voprosi ratsional’nogo ispol’zovaniya zapasov
kaspiiskogo tyulenya (Rational exploitation of Caspian seal
reserves). Sb. Morsk. Mlekop. Moscow.

Timoshenko, Yu.K. 1969. O srokakh nastupleniya polovoi zrelosti u
samtsov kaspiiskogo tyulenya (Onset of sexual maturity in Caspian
seal males). Sb. Morsk. Mlekop. Moscow.

Timoshenko, Yu.K. and V.A. Potelov. 1969. Sravnitel’naya morfologich-
eskaya kharakteristika kaspiiskogo tyulenya i kol’chatoi nerpy (Com-
parative morphological characteristics of Caspian seal and ringed
seal). Tez. Dokl. IV Vses. Soveshch. po Izuch. Morsk. Mlekop. Moscow.

Tomilin, A.G. 1936a. Kashalot Kamchatskogo morya (Sperm whale of
the Kamchatka Sea). Zool. Zhurn., vol. 15, no. 3.

Tomilin, A.G. 1936b. Ambra i ee proiskhozhdenie (Ambergris and its
origin). Priroda, no. 5. |

Tomilin, А.С. 1937. Киу Dal’nego Vostoka (Whales of the Far East).
Uch. Zap. MGU, Zool., no. 17.

Tomilin, A.G. 1937. Seryi kit v lagunakh vostochnogo poberezh’ya sred-
nei chasti Beringova morya (Gray whale in the lagoons on the eastern
coast of the central Bering Sea). Priroda, no. 7.

Tomilin, A.G. 1938. Butylkonos i melkie polosatiki Dal’nego Vostoka
(Bottlenose whale and Minke fin whale of the Far East). Byull.
MOIP, vol. 47, no. 3.

Tomilin, A.G. 1945. Opredelenie vozrasta kitov po usovomu apparatu
(Age determination of whales from baleen plates). DAN SSSR, Nov.
Ser., vol. 49, no. 6.

Tomilin, A.G. 1946a. Termoregulyatsiya i geograficheskie rasy kito-
obraznykh (Thermal regulation and geographic races of cetaceans).
DAN SSSR, Nov. Ser., vol. 54, no. 5.

Tomilin, A.G. 1946b. K voprosu o laktatsii i pitanii kitoobraznykh (Lac-
tation and feeding of cetaceans). Byull. MOIP, vol. 51, nos. 4 and 5.

Tomilin, А.С. 1951. Opredelitel’ kitoobraznykh po povedeniyu i vnesh-
nim priznakam (Key to Cetaceans Based on Behavior and External
Features). Izd-vo MOIP, Moscow.

Tomilin, А.О. 1952. О severnom berardiuse ili tak nazyvaemom
dal’nevostochnom “butylkonose” (Northern Baird’s whale or so-
called Far East “БоШепозе”). Byull. MOIP, vol. 57, no. 2.

Tomilin, А.С. 1953. К voprosu о sistematicheskikh vzaimootnosheniyakh
severnykh i yuzhnykh polosatikov (Taxonomic relations between
northern and southern fin whales). Byull. MOIP, vol. 58, no. 6.

Tomilin, A.G. 1955. O povedenii i zvukovoi signalizatsii kitoobraznykh
(Behavior and sound signals of cetaceans). Tr. In-ta Okeanol. AN
SSSR, vol. 18.

949

Tomilin, А.С. 1957. Zveri SSSR i prilezhashchikh stran, t. 9.
Kitoobraznye (Animals of the USSR and Neighboring Countries,
vol. 9. Cetaceans). Izd-vo AN SSSR, Moscow.

Tomilin, A.G. 1962. Kitoobraznye fauny morei SSSR (Cetacean Fauna
of Soviet seas). Izd-vo AN SSSR, Moscow.

Tomilin, A.G. 1965. O migratsiyakh, geograficheskikh rasakh, termoreg-
ulyatsii i vliyanii temperatury sredy na rasprostranenie kitoobraznykh
(Migrations, geographic races, thermal regulation, and effect of envi-
ronmental temperature on the distribution of cetaceans). Sb. Migr.
Zhiv., no. 25. Moscow.

Tomilin, А.С. and О.А. Morozov. 1969. Vsasyvanie korma—ranee
neizvestnyi sposob pitaniya Phocaena phocaena (Food intake—a
hitherto unknown method of feeding in Phocaena phocaena). Tr.
VSKhIZO, vol. 31.

Tormosov, D.D. 1970. Geograficheskoe rasprostranenie razlichnykh
fiziologicheskikh i vozrastnykh grupp kashalotov у yuzhnom
polusharii (Geographic range of different physiological. and age
groups of sperm whales in the Southern Hemisphere). Ти.
AtlantNIRO, no. 25. |

Treshchev, V.V. and А.М. Serdyukov. 1965. Materialy Chukotskoi eks-
peditsii 1964 g. (Data of Chuchi expedition of 1964). Itogov. Nauch.
Konf. Profes.-prep. Sostava Krym. Ped. In-ta. Simferopol’.

Treshchev, V.V., V.A. Potelov, and D.D. Zavaleeva. 1967. K izucheniyu
gel’mintofauny kol’chatoi nerpy i khokhlacha (A study of the
helminth fauna of ringed and hooded seals). Tr. PINRO, no. 21.

Tryuber, I.F. 1937. Izmenchivost’ cherepa chernomorskogo del’fina Del-
phinus delphis ponticus Barabash (Skull variation in Black Sea dol-
phin Delphinus delphis ponticus Barabash). Byull. MOIP, Otd. Biol.,
46 (1).

Tyulin, A. 1938. Promyslovaya fauna о. Belogo (Game fauna of Belhi
Island). Tr. МИ Polyar. Zemled., Zhivotn. i Promysl. Khoz., vol. 1.
Ural’skaya, I.V. 1957. Khimicheskii sostav moloka del’finov Chernogo
morya (Chemical composition of the milk of Black Sea dolphins).

Uch. Zap. Rost. Gos. Un-ta, vol. 57, no. 1.

Urvantsev, N.N. 1935. Dva goda na Severnoi zemle (Two Years in the
Soviet North). Leningrad.

Usachev, P.I. 1940. Obrastanie kitov diatomovymi vodoroslyami (Chok-
ing of whales due to diatomaceous algae). Zool. Zhurn., vol. 19, no. 2.

Ushakov, G.A. 1953. Po nekhozhennoi zemle (On Virgin Soil). Moscow.

Uspenskii, S.M. 1941. К nakhozhdeniyu tevyaka (Halichoerus grypus) у
zapovednike “Зет? ostrovov” [Find of Atlantic gray seal (Halichoerus

950

grypus) in Sem’ Ostrovov sanctuary]. Nauch.-Met. Zap. Gl. Орг. ро
Zapov., Zoop., Zoosad., no. 8.

Uspenskii, S.M. 1955. Proshloe rasprostranenie kotika (Callorhinus игя-
nus L.) i kalana (Enhydra lutris L.) na ostrovakh Kuril’skoi gryady
[Past distribution of northern fur seal (Callorhinus ursinus L.) and
sea otter (Enhydra lutris Г.) on the Kuril Island range]. Byull. MOIP,
vol. 60, no. 4.

Uspenskii, S.M. 1956. Pozvonochnye zhivotnye Tsentral’noi Arktiki
(Central arctic vertebrates). Priroda, no. 8.

Uspenskii, S.M. 1958. Narvaly у Tsentral’noi Arktike (Central arctic nar-
whals). Priroda, no. 3.

Uspenskii, S.M. 1958. Raspredelenie morzha v more Laptevykh i
zapadnoi chasti Vostochno-Sibirskogo morya у zimne-vesennii
period (Walrus distribution in the Laptev Sea and western East
Siberian Sea in winter-spring). Probl. Severa, no. 2.

Vaigachev, A.A. 1958. Sluchai v Arktike za 81 parallel’yu (An incident
in the Arctic beyond the 81st parallel). Priroda, no. 6.

Valikov, М. 1936. Promysel belukhi у 1935 р. na Obskom severe (Beluga
hunting in 1935 in the Northern Ob’). Za Rybn. Industriyu Severa,
no. 3.

Varpakhovskii, N. 1891. Neobkhodimost’ issledovaniya kaspiiakogo tyu-
lenya (Need for studying Caspian seal). Rybn. Delo, no. 2.

Vedenskii, А.Р. 1950. Nakhozhdenie krylatki (Histriofoca fasciata) у
Tsusimskom prolive [Find of ribbon seal (Histriophoca fasciata) in
Tsusimsk Strait]. Izv. TINRO, vol. 32.

Velizhanin, A.G. 1967. Novyi vid tyulenya (New species of seal). Priroda,
no. 7.

Vinogradov, M.P. 1949. Morskie mlekopitayushchie Arktiki (Marine
mammals of the Arctic). Tr. Arkt. In-ta, vol. 202.

Vishnyakov, A.P. 1961. Tyulen’-peshekhod (A stray seal). Priroda, no. 7.

Viladimirov, У.А. 1972. О dinamike chislennosti morskikh kotikov
ostrova Tyulenii (Population dynamics of fur seals on Tyulen’
Island). Tr. VNIRO, vol. 82; Izv. TINRO, vol. 80.

Vladimirov, V.A. and F.G. Chelnokov. 1972. Lezhbishcha morskikh
kotikov na Komandorskikh ostrovakh (Fur seal rookeries on the
Commander Islands). Tr. VNIRO, vol. 82; Izv. TINRO, vol. 80.

Voloshinov, N. 1889. Morskie kotiki (Fur Seals), pt. 1. St. Petersburg.

Voronin, L.G. 1970. Povedenie “primata” morya (Behavior of a sea “pri-
mate”). Usp. Sovr. Biol., vol. 69, no. 2.

Vrublevskii, V.K. 1959. Narpa v lesu (Ringed seal in a forest). Priroda,
no. 3.

95

Vylegzhanin, А.Е. 1965. Diplokokkovoe zabolevanie kaspiiskikh tyulenei
(Diplococcus disease in Caspian seals). Rybn. Khoz., no. 4.

Vylegzhanin, А.Е. 1968. Vibratsionnoe zabolevanie kaspiiskikh tyulenei
(Disease caused by vibration in Caspian seals). Rybn. Khoz., no. 3.

Yablokov, A.V. 1956. Ob interesnoi funktsii beloi okraski (An interest-
ing function of white coloration). Priroda, no. 6.

Yablokov, A.V. 1957. Ob organskh khimicheskogo vospriyatiya i
zhelezakh spetsial’nogo naznacheniya и zubatykh kitoobraznykh
(Organs of chemical perception and special purpose glands in
toothed cetaceans). Sb. Nauch. Stud. Rabot. MGU. Moscow.

Yablokov, A.V. 1958a. O streenii zubnoi sistemy i tipakh zubov u kitoo-
braznykh (Dental structure and type of teeth in cetaceans). Byull.
MOIP, Otd. Biol., vol. 63.

Yablokov, A.V. 1958b. K morfologii pishchevaritel’nogo trakta zubatykh
kitoobraznykh (Morphology of the alimentary canal in toothed
cetaceans). Zool. Zhurn., vol. 37, no. 4.

Yablokov, A.V. 1959. Nekotorye osobennosti stroeniya skeleta belukhi
kak predstavitelya zubatykh kitoobraznykh (Some structural features
of the skeleton of beluga as a member of toothed cetaceans). Tr. I
Konf. Mol. Nauch. Sotr. Mosk. Morf. Laboratorii. Moscow.

Yablokov, A.V. 1961a. Ob “obonyanii” morskikh mlekopitayushchikh
(Sense of smell in marine mammals). Tr. Soveshch. Ikht. Komis. AN
SSSR, no. 12.

Yablokov, A.V. 1961b. Funktsional’naya morfologiya organov dykhaniya
zubatykh kitoobraznykh (Functional morphology of respiratory
organs in toothed cetaceans). Tr. Soveshch. Ikht. Komis. AN SSSR,
no. 12.

Yablokov, A.V. 1962. Klyuch k biologicheskoi zagadke (Key to biological
enigma). Priroda, no. 4.

Yablokov, A.V. 1962. Sud’ba grenlandskogo tyulenye (Fate of harp seal).
Priroda, no. 2.

Yablokov, A.V. 1963. Nekotorye predvaritel’nye materialy k morfo-
logicheskoi kharakteristike grenlandskogo tyulenya Belogo i Gren-
landskogo morei (Some preliminary data on the morphological char-
acteristics of harp seal of the White and Greenland seas). Sb. Rabot
PINRO za 1962 g. Arkhangel’sk.

Yablokov, A.V. 1964. Konvergentsiya ili parallelizm у razvitii
kitoobraznykh (Convergence or parallelism in the development of
cetaceans). Paleont. Zhurn., no. 1.

Yablokov, A.V., V.M. Bel’kovich, and V.I. Borisov. 1972. Kity i del’finy
(Whales and Dolphins). Nauka, Moscow.

952

Yablokov, A.V. and С.А. Klevezal’. 1964. Vibrissy kitoobraznykh i
lastonogikh, ikh raspredelenie, stroenie, znachenie (Whiskers of
cetaceans and pinnipeds and their distribution, structure, and
importance). Sb. Morf. Osob. Vodn. Mlekop. Moscow.

Yablokov, A.V. and Р.Е. Sergeant. 1963. Izmenchivost’ kraniologich-
eskikh priznakov grenlandskogo tyulenya (Pagophilus groenlandicus
Erxl., 1777) [Variability of craniological characteristics of harp seal
(Pagophilus groenlandicus Erxl., 1777)]. Zool. Zhurn., vol. 42, no. 12.

Yablokov, A.V. and V.Ya. Etin. 1965. Analiz populyatsionnykh razlichii
у Okraske tela mlekopitayushchikh (na ритеге grenlandskogo tyu-
lenya) [An analysis of body color variations of different populations
of mammals (on the example of harp seal)]. Zool. Zhurn., vol. 44,
no. 7.

Yakovenko, M.Ya. 1957. О sovetskom promysle melkikh polosatikov
(minke) v moryakh Evropeiskogo Severa [Soviet hunting of lesser
finback (Minke) whale in seas of the European North]. Nauch.-tekhn.
Byull. PINRO, nos. 1 and 2.

Yakovenko, M.Ya. 1958. Malye polosatiki sredi Г4оу (Lesser finback
whales on ice). Priroda, no. 2.

Yakovenko, M. Ya. 1959. О metodike opredeleniya vozrasta i nekotorykh
voprosakh biologii khokhlacha (Method of determining the age and
some biological aspects of hooded seal). Byull. PINRO, no. 4 (8).

Yakovenko, M.Ya. 1960. Nablyudeniya za vozrastno-polovym sostavom
belomorskogo lysuna v 1959 g. (Observations on the age and sex
composition of White Sea harp seal in 1959). Nauch.-tekhn. Byull.
PINRO, no. 3 (13).

Yakovenko, M.Ya. 1960. Opredelenie vozrasta i srokov nastupleniya
polovoi zrelosti u belomorskogo lysuna (Determining the age and the
onset of sexual maturity in White Sea harp seals). Tr. PINRO, no. 12.

Yakovenko, М.Уа. 1961а. Metodika opredeleniya vozrasta belo-
morskogo lysuna v svyazi s opredeleniem norm ego promysla
(Method of determining the age of White Sea harp seal in the context
of fixing the hunting norms). Ту. Soveshch. Ikht. Komis. AN SSSR,
no. 12.

Yakovenko, M.Ya. 1961b. Issledovanie sostoyaniya zapasov i promysla
belomorskogo lysuna (A study of the reserves status and hunting of
White Sea harp seal). Tez. Dokl. Sessii Uch. Sov. Karel. Fil. AN SSSR.
Petrozavodsk.

Yakovenko, M.Ya. 1963. Razvitie promysla i problemy chislennosti belo-
morskogo lysuna (Development of hunting and population problems
of White Sea harp seal). Tr. PINRO, по. 15.

953

Yakovenko, М.Уа. 1967. Belomorskaya populyatsiya grenlandskogo
tyulenya i perspektivy ee ekspluatatsii (White Sea population of harp
seal and prospects of its exploitation). Tr. PINRO, no. 21.

Yakovenko, M.Ya. 1970. Rezul’taty aerofotoucheta chislennosti belo-
morskogo lysuna v 1968 g. (Results of aerial photographic census
of White Sea harp seal in 1968). Mater. Rybokhoz. 1551. Sever. Bass.,
no. 16 (pt. 2). PINRO, Murman.

Yakovenko, M.Ya. and Yu.I. Nazarenko. 1962. Rezul’taty issledovaniya
promysla i vozrastnogo sostava belomorskogo lysuna v 1960-1961
gg. (Results of studies on hunting and age composition of White Sea
harp seal in 1960-1961). Nauch.-tekhn. Byull. PINRO, vol. 1 (19).

Yakovenko, M.Ya. and Yu.J. Nazarenko. 1967. Materialy ро
razmnozheniyu grenlandskogo tyulenya belomorskoi populyatsii
(Data on the reproduction of White Sea population of harp seal).
Tr. PINRO, no. 21.

Yakovenko, М.Уа., Уи.Г. Nazarenko, and Yu.K. Timoshenko.
1963. Kratkie itogi promysla i nekotorye rezul’taty issledovanii
belomorskogo lysuna v 1962 g. (Brief data on hunting and some
results of studies on White Sea harp seal in 1962). Sb. Nauch.-issl.
Rabot 1962 g. Arkhangel’sk.

Yakovenko, M.Ya. and R.Sh. Khuzin. 1963. O raspredelenii i biologii
khokhlacha (Cystophora cristata) Grenlandskogo morya [Distribu-
tion and biology of hooded seal (Cystophora cristata) in the Green-
land Sea]. Tr. PINRO, по. 15.

Yakovlev, V. 1875. Tyulenii promysel na Kaspiiskom more (Seal hunting
in the Caspian Sea). Priroda, bk. 2.

Zaikin, V.V. 1953. Razrabotka sposoba obezzhirivaniya sala kashalota
5 sokhraneniem kozhevennogo syr’ya (Developing a method for
degreasing sperm whale blubber while preserving the skin). 7r.
VNIRO, vol. 25.

Zaikov, M.F. 1934. Promysel belukhi (Delphinapterus leucas Pall.) v
Obskoi gube latom 1932 g. [Beluga (Delphinapterus leucas Pall.)
hunting in Obsk Gulf in summer of 1932]. Raboty Obsko-Tazovskoi
Nauch. Rybkhoz. St., VNIRO, vol. 2, no. 1.

Zasosov, A.V. 1971. Sposob opredeleniya velichiny kotikovogo stada i
maksimal’noi vozmozhnoi stabil’noi kvoty ikh vyboya (Method for
determining the size of fur seal herds and the maximum stable quota
permissible in killing them). Tr. AtlantNIRO, no. 39.

Zakharov, У.Е. 1958. Lezhbishche morzhei na о. Vil’kitskogo (Walrus
rookeries on Vil’kits Island). Probl. Arktiki, no. 5.

Zemskii, У.А. 1950а. К biologii razmnozheniya nekotorykh vidov
usatykh kitov Antarktiki (Reproduction biology of some species of
Antarctic baleen whales). Byull. MOIP, vol. 55, no. 2.

954

Zemskii, У.А. 19506. Materialy К izucheniyu embrional’nogo razvitiya
Balaenoptera physalus L. Antarktiki (Data on embryonal growth of
Balaenoptera physalus L. of Antarctica). Byull. MOIP, vol. 55, no. 6.

Zemskii, V.A. 1953. Kharakteristika antarkticheskogo stada finvalov
(Characteristics of Antarctic fin whale herds). Tr. VNIRO, vol. 25

Zemskii, V.A. 1958a. Opredelenie sledov zheltykh tel и finvala
Antarktiki (Determining traces of the corpus luteum in Antarctic
fin whales). Tr. VNIRO, vol. 33.

Zemskii, У.А. 1958b. Rost novorozhdennykh finvalov у period
molochnogo pitaniya (Growth of newborn fin whales during the
suckling period). Tr. VNIRO, vol. 33.

Zemskii, V.A. 1958c. Morfobiologicheskie prisposobleniya novorozhden-
nykh finvalov Antarktiki (Morphobiological adaptations of newborn
fin whales of Antarctica). Tr. VNIRO, vol. 33.

Zemskii, У.А. 1961. О prodolzhitel’nosti zhizni finvalov i periodike
ikh razmnozheniya (Longevity of fin whales and their reproduction
cycle). [khtiol. Komis. AN SSSR. Tr. Soveshch., no. 12.

Zemskii, V.A. 1962. Kity Antarktiki (Whales of Antarctica). Kaliningrad.

Zemskii, V.A. and G.A. Budylenko. 1970. O kosatkakh Severnogo i
Yuzhnogo polusharii (Killer whales of the Northern and Southern
Hemispheres). Tr. AtlantNIRO, no. 25.

Zemskii, V.A. and A.A. Berzin. 1959. Nakhodka ambry (Find of amber-
gris). Priroda, no. 3.

Zenkovich, B.A. 1934. Materialy К poznaniyu kitoobraznykh dal’nevosto-
chnykh morei. Seryi kaliforniiskii kit (R. glaucus Cope) [Key to
cetaceans in the Far East seas. Gray Californian whale (R. glaucus
Cope)]. Vest. D/v Fil. AN SSSR, vol. 10.

Zenkovich, B.A. 1935. O zarodyshakh kitov (Whale embryos). Dokl. AN
SSSR, vol. 2, nos. 3 and 4. ;

Zenkovich, B.A. 1936a. Nablyudeniya nad kitami dal’nevostochnykh
morei v 1933 g. (Observations on Far East whales in 1933). Vest.
Dj Fil. AN SSSR, vol. 1.

Zenkovich, B.A. 1936b. O migratsii kitov v severnoi chasti Tikhogo
okeana (Whale migrations in the North Pacific Ocean). Izv. TINRO,
vol. 10.

Zenkovich, B.A. 1937a. Gorbatyi, ili dlinnorukii kit (Megaptera nodosa
Bonaterre, 1789) [Humpbak whale (Megaptera nodosa Bonaterre,
1789)]. Vest. D/v Fil. AN SSSR, vol. 27.

Zenkovich, B.A. 1937b. Vzveshivanie kitov (Weighing whales). DAN
SSSR, Nov. Ser., vol. 16, no. 3.

Zenkovich, B.A. 1937c. Pishcha dal’nevostochnykh kitov (Feeding of Far
East whales). DAN SSSR, vol. 16, no. 4.

955

Zenkovich, В.А. 19374. Eshche о serom kaliforniiskom kite (Rachia-
nectes glaucus Cope) [More about gray Californian whale
(Rachianectes glaucus Соре)]. Vest. D/v Fil. AN SSSR, no. 23.

Zenkovich, B.A. 1938a. Temperatura tela kitov (Body temperature of
whales). DAN SSSR, vol. 19, no. 9.

Zenkovich, B.A. 1938b. O kosatke, ili kite-ubiitse (Killer whale). Priroda,
no. 4.

Zenkovich, B.A. 1938c. Razvitie promysla morskikh mlekopitayushchikh
na Chukotke (Hunting of marine mammals in the Chukchi). Priroda,
nos. 11 and 12.

Zenkovich, В.А. 19384. О зеГдуапот Kite, ili finvale dal’nevostochnykh
morei (Balaenoptera physalus L.) [Finwhale (Balaenoptera physalus
L.) in the Far East seas]. Priroda, no. 6.

Zenkovich, В.А. 1939. Novyi ob”ekt kitoboinogo promysla (New target
of whale hunting). Priroda, no. 2.

Zenkovich, B.A. 1951. Ambra (seraya ambra) [Ambergris (gray amber-
gris)]. Rybn. Khoz., no. 4.

Zenkovich, B.A. 1952. Kity i kitoboinyi promysel (Whales and Whale
Hunting). Pishchepromizdat.

Zenkovich, B.A. 1955. Promysel malogo polosatika (Hunting of lesser
fin whale). Rybn. Khoz., no. 7.

Zenkovich, B.A. 1958. Promysel kitov v Antarktike v 1931-1955 gg. i
soobrazheniya о sostoyanii ikh zapasov (Whale hunting in Antarctica
in 1931-1955 and an estimate of their reserves). Tr. VNIRO, vol. 33.

Zenkovich, B.A. 1971. Uchast’ kitov (Density of whales). Tr. AtlantNIRO,
no. 39.

Гегпоу, S.A. 1913. К voprosu ob izuchenii zhizni Chernogo morya (о
tyulene-monakhe) [A study of Black Sea life (monk seals)]. Zap. AN,
vol. 32. St. Petersburg.

Zernov, S.A. 1913. Kratkii otchet po komandirovke ot Zoologicheskogo
muzeya Imp. Akademii nauk dlya sobiraniya kollektsii v Chernom
more u beregov Turtsii (Anatolii) v 1912 g. [Résumé of the mission
of the Zoological Museum of the Imperial Academy of Sciences for
making collections in the Black Sea on Turkish coasts (Anatolia) in
1912]. Ezhegodnik Zool. Muzeya Imp. Akad. Nauk, vol. 18.

Zhemkova, Z.P. 1965. О proiskhozhdenii kitoobraznykh (Origin of
cetaceans). Zool. Zhurn., vol. 44, no. 10.

Zhitkov, B.M. 1901. Po severu Rossii (Along northern Russia). Otchet
Imp. Ob-vu Lyubit. Est., Antrop. 1 Etnogr. po Kom-ke Letom 1910 р.
na o. Kolguev i Novuyu Zemlyu.

Zhitkov, B.M. 1904. Otchety ekspeditsii Russkogo geograficheskogo
obshchestva na Kanin poluostrov v 1902 g. (An account of the

956

Russian Geographic Society expedition into Kanin peninsula т
1902). Zap. Russk. Geogr. Ob-va po Obshch. Geogr., vol. 41, no. 1.

Zhitkov, B.M. 1904. Po Kaninskoi tundre (On Kanin tundra). Zap. Russk.
Geogr. Ob-va, vol. 41, no. 1.

Zhitkov, B.M. 1913a. O kollektsii mlekopitayushchikh, sobrannykh Koly-
makoi okspeditsiei 1905 g. (Mammals collected by the 1905 Kolyma
expedition). Dnevnik Zool. Otd. Ob-va Lyubit. Estastvozn., Antropol.
1 Etnogr., Nov. Ser., vol. 1, no. 5.

Zhitkov, В.М. 1913b. Poluostrov Yamal (Yamal Peninsula). St. Peters-
burg.

Zhitkov, B.M. 1924. Morskie zveri i morskie promysly (Sea Animals and
Hunting at Sea). Moscow.

Zhitkov, B.M. and S.A. Buturlin. 1901. Po severy Rossii (In northern
Russia). Zemlevedenie, vol. 8, books 3 and 4.

Zimushko, V.V. 1969. Nekotorye dannye po biologii serogo kita (Some
data on the biology of gray whale). Sb. Morsk. Mlekop. Nauka,
Moscow.

Zimushko, V.V. 1971. Materialy po razmnozheniyu serykh kitov (Data
on the reproduction of gray whales). Tr. AtlantNIRO, no. 39.

683

OTHER SOURCES

Adams L. 1968. Census of the Gray Whale, 1966-1967. Norsk
Hvalfangst—Tidende, N. 2, 41-43.

Allen J.A. 1880. History of North American Pinnipeds. Washington.

Allen J.A. 1902. List of Mammals, collected in Alasca by the Andrew J.
Stone—Expedition of 1901. Bull. Amer. Mus. Nat. Hist., 16.

Allen J.A. 1908. The North Atlantic Righ Whale and its allies. Bull.
Amer. Mus. Nat. Hist., 24, 277-329.

Allen G.M. 1930. The Walrus in New England. Journ. Mammal., v. 11,
№2.

Allen С.М. 1938. Order Pinnipedia. The Mammals of China and Моп-
golia, у. II, pt. I, Amer. Mus. Nat. Hist.

Amoroso E.C. and Mathews J.H. 1951. The growth of the grey seal (Hali-
choerus grypus Fabricius) from birth to weaning. Journ. of Anatomy,
у. 85, pt. 4, 427-428.

Anderson R. 1934. Mammals of the eastern Arctic and Hudson Bay.
Canada’s eastern Arctic. Bull. Canad. Dept. Inter.

Anderson S. and Jones J.K. 1967. Recent mammals of the world. The
Ronald Press Company, N. Y.

Andersson R.M. 1942. Two new seals from arctic Canada with key to
the canadian forms of hair seals (family Phocidae). Annal. Rep.
Provancher Soc. Nat. Hist. Quebec.

Andrews R.C. 1909. Observation on the habits of the Finback and Hump-
back Whales of the North Pacific. Bull. Amer. Mus. Nat. Hist., v. 26,
222226:

Andrews В.С. 1914. The California Gray Whale (Rachianectes glaucus
Cope). Memories Amer. Mus. Nat. Hist.

Aul J., Ling H. and Paaver K. 1957. Eesti NSV imetajad. Tallin.

Backhouse K.M. 1960. The grey seal (Halichoerus grypus) outside the
breeding season. A preliminary report. Mammal., v. 24, N. 3.

Backhouse K.M. and Hewer H.R. 1956. Delayed implantation in the grey
seal, Halichoerus grypus (F.). Nature, v. 178.

686

958

Backhouse K.M. and Hewer H.R. 1957. A note on spring pupping in
the grey seal (Halichoerus grypus Fab.). Proc. Zool. Soc. London,
v. 128.

Backhouse K.M. and Hewer H.R. 1957. Behaviour of the grey seal (Hali-
choerus grypus Fab.) in the spring. Proc. Zool. Soc. London, v. 129.

Backhouse H.R. and Hewer H.R. 1960. Unusual colouring in the grey
seal Halichoerus grypus (Fabricius). Proc. Zool. Soc. London, v. 134,
pt. 3, 497 - 499.

Backus R.H. and Schevill W.E. 1961. The stranding of a Cuvier’s
Beaked Whale (Ziphius cavirostris) in Rhode Island, U.S.A. Norsk
Hvalfangst—Tidende, М. 5, 177 - 181.

Bailey A.M. and Hendee R.W. 1926. Notes on the Mammals of North
western Alasca. Journ. Mammal., v. 7.

Baker R.C., Wilke F. and Baltzo C.H. 1963. The Northern Fur Seal. Fish
and Wildlife Service, Bureau of Commercial Fisheries, Washington,
20

Bartholomew С. and Hoel Р. 1953. Reproductive behavior of the Alaska
Fur Seal. Journ. Mammal., у. 34. М. 4, 417 - 436.

Bartlett R.A. 1929. The seling saga of Newfoundland. Nat. Georg. Мар.,
у. 61; №. 1.

Baylis Н.А. and Daubney К. 1925. A revision of the lung-worms of
Cetacea. Parasitol., v. 17, N. 2, 201-216. Cambridge.

Beddard F.W. 1900. A book of Whales. London.

Bee J.W. and Hall E.R. 1956. Mammals of northern Alaska. Univ. Kansas
Misc. Publ. N. 8.

Bee J.W., Hall E.R. 1958. Mammals of Northern Alaska. Univ. Kansas
Mus. Nat. Hist. Misc. Publ. N. 8. p. p. 1-310.

Bemmel A. 1956. Planning a census of the harbour seal (Phoca vit-
ulina L.) on the coasts of the Netherlands. Beaufortia. Ser. of mis-
celaneous publications. v. 5, N. 54. Amsterdam.

Bemmel A. 1968. Eine merwurdige Farbvariante beim gewohnlichen See-
hund. Natur und Museum, у. 98, М. 4, 167-169.

Benson S. and Grood T. 1942. Notes on the Dall porpoise (Phocaenoides
dalli), Journ. Mammal, v. 23, 41-51.

Bergman G. 1956. Rannikoi Hemme Nyljekannajta. Suomen tutkija, 60,
N. 3.

Bergman G. 1956. Salbestandet vid vora kuster. Nordenskiold samlundets
tidskrift.

Bergman G. 1958. Suomen Hyljekannoista. Suomen Riista Helsinki.

Berland B. 1958. The hood of the hooded seal, Cystophora cristata Erxl.
Nature, v. 182, 408 - 409.

959

Berland В. 1963. Phocascaris cystophorae sp. nov. (Nematoda) from the
hooded seal, with an emendation of the genus. Arbok for univer-
sitetet i Bergen. Nat.—Naturv. serie, N. 17. Norvegian Universities
press. 21 p. Bergen—Oslo.

Berland B. 1966. The Hood and its extrusible ballon in the Hooded seal
Cystophora cristata Erxl. Norsk Polarinstitutt—Arbok, Oslo.

Bigg M. 1969. The harbour seal in British Columbia. Fish. Res. Bd.
Canada. Bull. 172.

Boettger C.R. 1951. Notizen zur Verbreitung und uber die Verwand-
schaftsbeziehungen der Monchsrobbe (Monachus albiventer Bod-
daert). Zool. Anz., Bd. 147, II. 11-12, 303—310.

Boyd J.M. 1962. Seasonal occurrence and movements of seals in north-
west Britain. Proc. Zool. Soc. London, v. 138.

Boyden A. and Gemery D. 1950. The relative position of the Cetacea
among the orders of Mammalia as indicated by precipitating tests.
Zoologica, v. 35, 145-151.

Braun M. 1893. Uber die Distomen in Leben der Hauskatzen. Zool.
Anz., Bd. 16, 347-355.

Brown D.N. 1960. Behavior of a captive Pacific Pilot Whale. Journ. Mam-
mal., у. 41, М. 3, 342-349.

Brown R. 1868. Notes on the history and geographic relations of the
Pinnipedia frequenting the Spitzbergen and Greenland Seas. Proc.
Zool. Soc. London.

Brown S.G. 1956. Whale Marks recently recovered. Norsk Hvalfangst
Tidende, М. 12, 661-664.

Brown S.G. 1957. Whale marks recovered during the Antarctic Whaling
season 1956-57. Norsk Hvalfangst—Tidende, N. 10, 555-559.
Brown S.G. 1958. Whales and their distribution. Part I. The Marine

Observer, v. 2, N. 184.

Brown S.G. 1959. Whales observed in the Atlantic Ocean. Notes on their
distribution. Norsk Hvalfangst—Tidende, М. 6, 289-308.

Brown S.G. 1961. Observations of Pilot Whales (Globicephala) in the
North Atlantic Ocean. Norsk Hvalfangst—Tidende, N. 6, 225 -248.

Brondsted H.V. 1931. Bygningen af Snuden og Ansigtsmuskulaturen hos
nogle Pinnipedier med saerligt Hensyn til Oppustningssaekken hos
Klapmydesen. K. danske vidensk. Selsk. naturv. math., 41-85.

Brink van den Е.Н. 1958. Die Saugetiere Europas. Р. Parey.
Hamburg—Berlin, 1-225.

Brink van den F.H. 1967. A field guide to the mammals of Britain and
Europe. London, 1-221.

Buckley J.L. 1958. The Pacific Walrus. Fish and Wildlife Service. Special
Scientific Report N. 41. Washington.

687

960

Bullock T.H., Grinnel A.D., [kezono E., Katsuki J., Nomoto M., Saito O.,
Suga N., Janagisava K. 1968. Electrophysiological studies of central
auditory mechanism in Cetaceans. Zetschr. Verl. Physiol., Bd. 59,
М2.

Bunge A., and Schrenck Г. 1887. Expedition nach den Neusibirischen
Inseln und dem Jana—Lande. Petersb.

Burns J.J. and Fay F.H. 1970. Comparative morphology of the skull of
the Ribbon seal, Histriophoca fasciata, with remarks on systematics
of Phocidae. Journ. of Zoology, Proc. Zool. Soc. London, v. 161,
pt. 3, 363 -394.

Caldwell D.K. 1960. Notes on the Spotted Dolphin in the Gulf of Mexico.
Journ. Mammal., v. 41, М. 1, 134-136.

Caldwell D.K. and Caldwell M.C. 1969. The Harbour seal, Phoca vitulina
concolor in Floridae, Journ. Mammal., v. 50, N. 2.

Caldwell D.K., Inglis A. and Siebanaler J.B. 1960. Sperm and Pigmy
sperm Whales stranded in the Gulf of Mexico. Journ. Mammal.,
у. 41, М. 1, 136-138.

Caldwell D.K. 1964. Tusk twinning in the Pacific Walrus. Journ. Mam-
mal., у. 45, М. 3, 490-491.

Calinescu R.J. 1937. Focele dela Сару! Caliacra. Bull. Soc. Nat. 4. Roma-
nia, N. 8, 6-10.

Cameron A. 1970. Seasonal movements and diurnal activity rhytmus
of the Grey Seal (Halichoerus grypus). Journ. Zool., v. 161, pt. 1,
id= 23:

Cameron A.W. 1967. Breeding behavior in a colony of western Atlantic

gray seals. Canadian Journ. Zool., у. 45, 161-173.

Carrucio A. 1893. Su di un Pelagius monachus aduel e del suo feto presi
a Cap Teulado nel Mediterraneo. Boll. Soc. Romania Zool., v. 2.

Caspers Н. 1950. Beobachtungen uberdas Vorkommen der Monchsrobbe
(Monachus albiventer Bodd) im Schwarren Meer, Neune Ergebnisse
und Probleme der Zoologie. Zool. Anz. Erganz. Bd. 145,
91-105.

Chafe L.G. 1923. A history of the Newfoundland seal fishery from the
earliest available records down to and including the voyage of 1923.
St. John.

Chapman D.G. 1961. Population dynamics of the Alaska fur seal
heard. Wildlife Management Institute Transaction of the Twenty-
Sixth North American Wild-life and Natural Resources Conference.
Washington. 356-369.

Chapskij K.K. 1960. Morphologie, systematique, differentiation intra-
specifique et phylogenese du sousgenre Phoca sensu stricto.
“Mammalia”, Т. 24, М. 3, 343-360.

961

Chittleborough К.С. 1953. Aerial observation of the Humpback Whale,
Megaptera nodosa (Bonnaterr.), with notes on other species. Aus-
tral. J. Mar. Freshwater Res., v. 4, 219-226.

Chittleborough R.G. 1954. Studies on the ovaries of the Humpback
Whale, Megaptera nodosa (Bonnaterre) on Western Australian
Coast, Austral. J. Mar. u Freshwater Res., v. 5, N. 1, 35-63.

Chittleborough R.G. 1955a. Aspects of reproduction in the male Hump-
back Whale, Megaptera nodosa (Bonnaterre). Austral. J. Mar. u
Freshwater Res., v. 6, N. 1, 1-29.

Chittleborough R.G. 1955b. Puberty, physical maturity and relative
growth of the female Humpback Whale, Megaptera nodosa
(Bonnaterre) on the Western Australian coast. Austral. J. Mar. u
Freshwater Res., v. 6, N. 1, 315-327.

Chittleborough R.G. 1956. Southern right Whale in Australian waters.
Journ. Mammal., у. 37, М. 3, 456-457.

Chittleborough К.С. 1958. The breeding cycle of the female Humpback
Whale, Megaptera nodosa (Bonnaterre). Austral. J. Mar. u Fresh-
water Res., v. 7, N. 9, 1-18.

Chittleborough R.N. 1959. Determination of age in the Humpback
Whale, Megaptera nodosa (Bonnaterre). Austral. J. Mar. u
Freshwater Res., v. 10, N. 2, 125-143.

Christensen G. 1955. The stoks of Blue Whales in the Northern Atlantic.
Norsk. Hvalfangst—Tidende, N. 11, 640-642.

Christensen G. 1965. The Greenland Right Whale and whales wich sleep.
Norsk Hvalfangst—Tidende, N. 2, 55.

Clark E.S. 1958. Cuvier’s beaked whale (Ziphius cavirostris) strands on
Cape Cod Beach, USA. Norsk Hvalfangst—Tidende, N. 6, 291 - 293.

Clarke R. 1953. Sperm whaling from open boats in the Azores. Norsk
Hvalfangst—Tidende, N. 7, 373 -385.

Clarke R. 1955. A giant squid swallowed a sperm whale. Norsk
Hvalfangst—Tidende, N. 10, 589-593.

Clarke R. 1956. Sperm whale of the Azores. Discovery Reports, 28,
257.298.

Collett R. 1881. Оп Halichoerus grypus and из breeding on the Его
Islands off Trondhjems—fjords in Norway.—Proc. of the Sci. Meet-
ings of the Zool. Soc. London.

Collett R. 1911-1912. Norges Pattedyr. Kristiania.

Collins G. 1940. Habits of the Pacific Walrus (Odobenus rosmarus).
Journ. Mammal., у. 21, №. 2, 138-144.

Colman J.S. 1937. The present state of the Newfoundland seal fishery. J.
of Animal Ecol., v. VI, N. 1.

688

962

Colman J.S. 1938. The Newfoundland seal fishery and its influence on
the Greenlanders. Polar Rec., Cambridge, 16.

Colman J.S. 1949. The Newfoundland seal fishery and the second world
war. J. of Animal Ecol., v. 18, N. 1.

Cook H.W. and Baver B.E. 1969. Seal milk. I. Harp seal (Pagophilus
groenlandicus) milk: composition and pesticide residue content.
Canadian J. Zool., у. 47, М. 6, 1129 - 1133.

Coulson J.C. and Hickling G. 1964. The breeding biology of the grey seal,
Halichoerus grypus (Fab.) on the Farne Islands, Northumberland. J.
Animal Ecol., v. 33, 485 - 512.

Cowan Jan. 1944. The Dall—porpoise Phocoenoides dalli (True) of
Northern Pacific Ocean. Journ. Mammal., у. 25, М. 3, 295 - 306.
Craig A.M. 1964. Histology of reproduction and estrus cycle in the female
fur seal, Callorhinus ursinus. Journ. Fish. Res. Bd. Canada, v. 21,

М. 4, 773-811.

Curry-Lindahl К. 1965. The plight of the grey seal in the Baltic. “Oryx”,
v. VIII, N. 1, 38-44.

Daan K.H. and Douglas C.W. 1953. Beluga of the Churchill region of
Hudson Bay. Bull. Fish. Res. Bd. Canada, N. 98, 27p.

Dall W. and Dunston D. 1957. Euphausia superba Dana from a Hump-
back Whale Megaptera nodosa (Bonnaterre) caught off Southern
Queensland. Norsk Hvalfangst—Tidende, N. 1, 6-9.

Darling F. 1948. The Atlantic seal. The New Naturalist. London.

Dathe H. 1934. Ein Beitrag zur Wirbeltier—fauna Dalmatiens. Zool.
Garden, N. 7. Leipzig.

Davies J.L. 1957. The geography of the gray seal. Journ. Mammal., у. 38,
М. 3, 297-310.

Davies J.L., 1958. Pleistocene geography and the distribution of northern
pinnipeds. Ecology, v. 39, N 1: 97-113.

Davies J.L. 1960. The southern from of the Pilot Whale. Journ. Mammal.,
v. 41, N. 1, 29-34.

Dawbin, W.H. 1956. Whale marking in South Pacific waters. Norsk
Hvalfangst—Tidende, N. 9, 485 - 508.

Dawbin W.H. 1959. New Zealand and South Pacific Whale marking and
recoveries to the end of 1958. Norsk Hvalfangst—Tidende, N. 5,
213 - 238.

Degerbol M. and Freuchen P. 1935. Report of the Mammals collected
by the fifth thule—Expedition to arctic North America. Report of
the fifth Thule—Expedition 1921 - 1924, Zoology I, vol. II, N 4-5.

Dell R.K. 1960. The New Zealand occurrence of the Pygmy sperm Whale,
Коза breviceps. Rec. Dom. Mus., у. 3, М. 2, 221-234.

Deshaseaux C. 1961. Cetacea. Traite de Paleontologie, v. 1.

963

Dornesco С.Т. et Marcoci G.V. 1958. Etude comparative du crane des
Pinnipedes. Ann. des Sci. Lat. Zool., 11 ser.

Doutt J.K. 1942. A review of the genus Phoca. Ann. Carnegie Mus., 29,
61-75.

Dunbar М... 1949. The Pinnipedia of the arctic and sub-arctic. Bull. Fish.
Res. Bd. Canada, v. 85, 1-22.

Dunbar M.J. 1955. The status of the Atlantic walrus Odobaenus rosmarus
in Canada. Arctic circular, v. 8, N. 1, 11-14.

Duncan A. 1956. Notes on the food and parasites of grey seals, Hali-
choerus grypus, from the isle of Man. Proc. Zool. Soc. London,
у. 126, М. 4, 635 - 644.

Ehlers К. 1958. Seltener Fang einer (Cystophora cristata Erxl.) bei 61° М
und 5° W. Polarforschung, Bd. 3, H. 1-2.

Ehlers K., Sierts W. and Mohr E. 1958. Die Klappmutze, Cystophora
cristata Erxl., der Tiergrotten Bremerhaven. Der Zoologische
Garten, Bd. 24, H. 3(4).

Ellerman J.R. and Morrison-Scott T.C.S. 1951. Checklist of Palearctic
and Indian mammals 1758 to 1946. British Mus. (Nat. Hist.)

Evans W.E. and Prescott J.H. 1961. Study of whistles and clicks obser-
vation capabilities of the bottlenose porpoise. Marinland of Pacific.
Biol. Lab., Contr. N. 15.

Fabricius O. 1791. Om de. gronlandske saele. Skriv. naturhist. Selsk., 1.

Fay F.H. 1957. History and present status of the Pacific Walrus popula-
tion. Wild-life Management Institute. Washington D.C., 431 - 445.

Felts W.J.L. 1966. Some functional and structural characteristics of
cetacean flippers and flukes. In.: K. Norris, ed. Whales, Dolphins
and Porpoises. Univ. Calif. Press, Berkeley, Los Angeles.

Fetcher E.S. 1939. The water balance in marine Mammals. Quart. Rev.
of Zool., v. 14, N. 4.

Fiscus C.H. 1961. Growth in the Steller sea lion. Journ. Mammal., v. 42,
№2, р. 218-223.

Fiscus С.Н. and Baines С.А. 1966. Food and feeding behavior of Steller
and California sea lions. Journ. Mammal., у. 47, М. 2, 195-200.
Fiscus C.H. and Niggol K. 1965. Observation of cetaceans off California,
Oregon and Washington. United States Fish Wildlife Service. Spec.

Scient. Rep. Fisheries N. 498, 1-27.

Fisher H.D. 1952. The status of the Harbour Seal in British Columbia,
with particular reference to the Skeena River. Bull. Fish. Res. Bd.
Canada, N. 93, Ottawa.

Fisher H.D. 1952. Harp seal of the northwest Atlantic. Fish. Res. Bd.
Canada, Atlantic Biol. Stat. Circ. N. 20.

689

964

Fisher H.D. and Sergeant D.R. 1954. А record of the whall in the Bay
of Fundy. Canadian Field—Nat., 68, 138 - 139.

Fisher H.D. 1954. Delayed implantation in the Harbour Seal, Phoca
vitulina L. Nature, v. 173, 879.

Fisher H.D. 1954. Studies on reproduction in the Harp Seal Phoca
groenlandica Erxleben in the northwest Atlantic. Fish. Res. Bd.
Canada, MLS. Rep. (Biol.) М. 588.

Fisher H.D. 1955. Utilization of Atlantic Harp Seal population. Trans.
20th. N. Amer. Wild. Conf.

Fisher H.D. and Mackenzie B.A. 1955. Food habits of seal in the
Maritimes. Fish. Res. Bd. Canada. Progress Rep. of the Atlantic
coast. Stat., N. 61, 5-9.

Fitter R.S.R. 1961. Status of the Grey Seal. Nature, v. 189, N. 4763, 506.

Нааи Е. und Jacobson Г. 1899. Handbuch der Anatomie des
Zentralnervensystems der Saugetiere. Bd. I. Berlin.

Foelsch D.W. 1968. Beobachtete geburt eines Seehundes (Ph. vitulina).
Zeitschr. fur Saugetierkunde, Bd. 33, H. 1, 50-54.

Fraser F.C. and Purves P.E. 1960. Hearing in Cetaceans. Bull. Brit. Mus.
Nat. Hist., Zool., у. 7. 1-40.

Freuchen P. 1935. Mammals. Pt. II. Field notes and biological obser-
vations. Report of the Sth Thule Exped. 1921-1924. Copenhagen
Nordish Forlag 2, 5-41.

Freund L. 1933. Pinnipedia. Die Tierwelt der Nord und Ostsee, Lief. 24.

Friedel E. 1882. Thierleben im Meer und am Strande von Neuvorpom-
mern. Zoologischer Garten. 23.

Fujinko K. 1955. On the Body Weigh of the Sei Whales localed in the
adjacent waters of Japan (2). Scient. Rep. Whal. Res. Inst., N. 10,
133 - 139.

Fujino K. 1956. On the Body proportion of the Sperm Whale (Physeter
catodon). Scient. Rep. Whal. Res. Inst., N. 11, 47-85.

Gaskin D.E. 1966. New records of the Pigmy Sperm Whale, Kogia bre-
viceps Blanville, 1838, from New Zealand and probable record from
New Guinea. Norsk Hvalfangst—Tidende, N. 2, 35 -37.

Gest O.W., Buckley J.L. and Manville R.H. 1960. Alascan Records of
the Narwhal. Journ. Mammal., У. 41, М. 2, 250-253.

Gill T. 1866. Prodrome of а monograph of the pinnipeds. Proc. Essex.
Inst., 5.

Gilmore R.M. 1955. The return of the Gray whale. Scientific Amer.
у. 192, М 1, 62 6).

Gilmore В.М. 1956. Rare right whale visits California. Pacific Discovery,
м. ЭМ. 4.

965

Gilmore R.M. 1957. Whales around in Cortes Sea. Pacific Discovery,
№10 №11: 22-27.

Gilmore КВ.М. 1959. On the strandings of Sperm Whales. Pacific Natu-
ralist, v. 1, N. 9-10.

Gilmore В.М. 1960a. A census of the California Gray Whale. U.S.
Fish. and Wildlife Service Special Scient. Rep., Fisheries N. 342.
Washington.

Gilmore R.M. 1960b. Census and migration of the California Gray
Whale. Norsk Hvalfangst—Tidende, N. 9, 409 - 431.

Gilmore R.M. and Ewing G. 1954. Calving of the California Grays.
Pacific Discovery, N. 7.

Gottberg G. 1927. Salfangsten ar 1926. Fiskeritidskrift for Finland,
INS 6-7.

Gottberg С. 1927. Statistik Over Salfangsten aren 1909-1918. Finlands
Fiskerier Landbruksstyrelsens Meddelanden, N. 167. Helsingf.

Gottberg G. 1931. Salfangsten 1930. Fiskeritidskrift for Finland, N. 8-9.

Gray J.E. 1825. Phocidae et Trichechidae. Ann. Phil.

Gray J.E. 1850. Catalogue of the specimens of Mammalia in the collec-
tion of the British Museum. Part II. Seals. London.

Grey J.E. 1864. Notes on the seals (Phocidae) including the description
of a new seals Halicyon richardii, from the west coast of the North
America. Proc. Zool. Soc. London.

Gray J.E. 1866. Catalogue of Seals and Whales in the British Museum.

Grosse A. and Transehe N. 1929. Verzeichnis der Wirbeltiere des Ost-
baltischen Gebietes. Arb. Natur. Ver. zu Riga, H. 18.

Haglund B. 1961. Sal. Stockholm.

Hall, E.R. and Kelson, K.R. 1959. The Mammals of North America, 2
volumes, The Ronald Press, New York.

Handley C.O. 1966. A synopsis of the genus Kogia. In.: K. Norris, ed.
Whales, Dolphins and Porpoises. Univ. Calif. Press, Berkeley and
Los Angeles. 62 - 70.

Harrison R.J. 1960. Reproduction and reproductive organs in common
seals (Phoca vitulina) in the Wash, East Anglia. Mammalia, T. 24,
М. 3, 372-385.

Havinga В. 1933. Der Seehund (Phoca vitulina L.) in den Hollandischen
Gewassern. Tijdschrift Nedd. Dierk. Ver., Ser. 3, Aft. 2-3.

Heck K. 1966. Eine Monchsrobbe Monachus m. monachus (Hermann,
1779) von Rhodos. Saugetierkundliche Mitteilungen, H. 1, 47.
Heer E. 1964. Beitrage zur Saugetierkunde Sud-Bessarabiens, Zeit.

Saugetierkunde, Bd. 29, H. 3, 173-178.

Heezen B.C. 1957. Whales entagled in deep Sea cables. Norsk

Hvalfangst—Tidende, М. 12, 665 -681.

690

966

Heinroth К. 1958. Uber Seehundgeburten und eine Seehundaufzucht
im Zoologischen Garten Berlin. Zoolog. Garten. Bd. 22, H. 1/3,
196 - 204.

Hershkovitz P. 1966. Catalog of living whales. Smiths. Inst. Bull. v. 246,
№. 259.

Hewer Н.К. Behaviour of the grey seal (Halichoerus grypus Fab.) in the
breeding season. Mammalia, v. 24, N. 3, 400-421.

Hewer H.R. 1964. The determination of age, sexual maturity, longevity
and life-table in the Grey seal (Halichoerus grypus). Proc. Zool. Soc.
London, v. 142, pt. 4.

Hewer H.R. and Backhouse K.M. 1959. Field identification on bull and
cows of the Grey Seal (Halichoerus grypus Fab.). Proc. Zool. Soc.
London, v. 3, pt. 4.

Hewer H.R. and Backhouse K.M. 1960. A preliminary account of a
colony of grey seals Halichoerus grypus, in the Southern Inner
Hebrides. Proc. Zool. Soc. London, v. 134, N. 2, 157-195.

Hickling G., Rasmussen B. and Smith E.A. 1962. Innvandring fra Storbri-
tannia til Norge av havert (Halichoerus grypus). Fisken of Havet,
N. 2, 4-10.

Hjort J. and Ruud J. 1929. Whales and plancton in the North Atlantic.
In.: Whaling and fishing in the North Atlantic. Rapport et Process
Verbaux, 61, 1-123.

Hobson E.S. 1966. Visual orientation and feeding in seals and sea lions.
Nature, v. 210, N. 5033.

Holm O. 1921. De bottniska sjalarternes vanor. Fauna och Flora, H. 6
Uppsala.

Holmgren A. 1865. Skandinaviens Daggdjur. Stockholm.

Hook O. 1960. Some observations on the dates of pupping and the inci-
dence of partial rust and orange coloration in Grey Seal caws, Hali-
choerus grypus Fab. on Lunga, Treshnish isles Argyll. Proc. Zool.
Soc. London, v. 134.

Horn D.R. van and Baker В.Е. 1971. Seal milk. П Harp. seal (Pagophilus
groenlandicus) milk: effect of stage of lactation on the composition
of the milk. Canadian Journ. Zool., v. 49, N. 8.

Hornschuch und Schilling. 1850. Kurze Notizen uber die in den Ostsee
vorkommenden Arten der Gattung Halichoerus. Greifswald.

Hoshina T. and Sugiura Y. 1956. On a skin disease and Nematode par-
asite of a dolphin Tursiops truncatus (Nontagu, 1821). Scient. Rep.
Whal. Res. Inst., М. 11, 133-138.

Hosokawa H. 1950. On the cetacean larynx, with special remarks on the
laryngeal sack of the sei whale and the arytenoepiglottideal tube of
the sperm whale. Scient. Rep. Whal. Res. Inst., N. 3.

967

Houck W.J. 1961. Notes of the Pacific striped Porpoises. Journ.
Mammal., v. 42, N. 1, 107-109.

Ichihara T. 1958. Grey Whale observed in the Bering Sea. Scient. Rep.
Whal. Res. Inst., М. 13, 201-205.

Ichihara T. 1961. Blue Whales in the water around Kergelen Island.
Norsk. Hvalfangst—Tidende, N. 1, 1-20.

Ichihara T. 1966. The Pygmy Blue Whale, Balaenoptera musculis brevi-
cauda, a New Subspecies from the Antarctic. Whales, Dolphins and
Porpoises, N. 6, 79-113. Univ. of Calif. Press.

Ichihara T. and Doi T. 1964. Stock assesment of Pygmy Blue Whales in
the Antarctic. Norsk. Hvalfangst—Tidende, М. 6, 145 - 167.

Imler R.H. and Sarber H.R. 1947. Harbour seal and sea lions in Alaska.
U.S. Fish. Wildlife Serv. Spec. Sci. Rep. 28.

Inukai T. 1942. Hair seals (azarashi) in our northern waters. Shokubutsu
Dobutsu (Botany and Zoology)., у. 10, М. 10, 11.

Irving L. 1938. Control of respiration in diving animals. Amer. Journ. of
Physiol., v. 123, 107-120.

Iversen B. 1955. Whaling activity in Iceland. Norsk Hvalfangst—Tidende,
М. 10, 598 - 606.

Janes J. 1952. On the whale brain with special reference to the weight
of the brain of the fin whale (Balaenoptera physalus.) Norsk
Hvalfangst—Tidende, М. 9, 480 - 486.

Japha A. 1910. Weiter Beitrage zur Kenntnis des Walhaut. Zool. Jahrb.

Johnson A.L. 1957. Seal marking 1956-57. Nature in Wales, v. 3, N. 1.

Johnson M.L., Fiscus C.H., Ostenson B.T. and Barbour M.L. 1966.
Marine Mammals. Envirinment of the Cape Thompsen region,
Alasca. Chapter 33. U. S. Atomic Energy Commission.

Jonsgard P.M. and Ke P.J. 1968. Principal fatty acids of depot fat and
milk lipids from harp seal (Pagophilus groenlandicus) and hooded
seal (Gystophora cristata).—Journ. Fish. Res. Bd. Canada, v. 25,
М. 11, 2419-2426.

Jonsgard А. 1951а. Studies on the little piked whale or minke whale
(Balaenoptera acutorostrata Lac.) Norsk Hvalfungst—Tidende, N. 5,
209-232:

Jonsgard А. 19516. On the growth of ше Fin Whale in different waters.
Norsk Hvalfangst—Tidende, N. 2, 57-65.

Jonsgard A. 1955a. The Stocks of Blue Whales (Balaenoptera musculus)
in the Northern Atlantic Ocean and adjacent Arctic waters. Norsk
Hvalfangst—Tidende, М. 9, 505-519.

Jonsgard A. 1955b. Development of the modern Norwegian small whale
industry. Norsk Hvalfangst—Tidende, N. 12, 697-718.

691

968

Jonsgard А. 1966. Biology of the North Atlantic Fin Whale, Balaenoptera
physalus. Universitet storlaget, Oslo.

Jonsgard A. and Nordli O. 1952. Concerning a catch of white sided
Dolphins (Lagenorhynchus acutus) on the West coast of Norway,
Winter 1952. Norsk Hvalfangst—Tidende, N. 5, 229-232.

Jonsgard A. and Ноа! P. 1957. Strandings of Sowerby’s Whale
(Mesoplodon bidens) on the West Coast of Norway. Norsk
Hvalfangst—Tidende, М. 9, 507 -512.

Jonsgard А. ор Оупез Р. 1952. Om bottlenosen (Hyperooden rostra-
tus) og spekkhoggern (Orcinus orca). Saertrykk ау “Fauna”, М. 1,
p. 1-18.

Juncker H. 1940. Die Aufzucht der Seehunde in den Tiergrotten der
Stadt Wesermunde. D. Zoolog. Garten, N.F., 12.

Karcker F.H. 1940. Uber den Algenbewuchs aufsiidlichen Walen. Zeit.
fur Fischerei, Beiheft 1.

Kasuya T. and Ohsumi S. 1966. A secondary sexual character of the
sperm-whale. Scient. Rep. Whal. Res. Inst., N. 20.

Kasuya T. and Rice D.W. 1970. Notes on Baleen plates and on arrarge-
ment of parasitic Barnacles of Grey Whale. Scient. Rep. Whal. Res.
Inst., N. 22, 39-43.

Kawakami T. and Ichihara T. 1958. Japanese whale marking in the North
Pacific in 1956 and 1957. Norsk Hvalfangst—Tidende, М. 6, 285 - 291.

Kellog W.N. 1953. Ultrasonic hearing in the porpoise. J. Comp. Physiol.
Psychol., v. 46.

Kellogg R. 1922. Pinnipeds from Miocene and Pleistocene deposits of
California. Univ. Calif. Publ. Geol. Sci., v. 13, М. 4, 23-123.

Kellogg R. 1928a. What is known of the migrations of some Whalebone
Whales. Ann. Rep. Smiths. Inst., v. 6, 467 - 484.

Kellogg В. 1928b. The history of whales, their adaptation to life in the
water. Quart., Rev. Biol. Baltimore, у. 3, 29-76. and 174-208.
Kellogg R. 1931. Whaling statistics for Pacific Coasts of North America.

Journ. Mammal., v. 12, N. 1.

Kellogg R. 1958. Echoranging in the porpoise. Science, v. 128, N. 3330,
982 - 988.

Kenyon K.W. 1956. Food of fur seal taken on St. Paul Island, Alaska,
1954, Journ. of Wildlife Management, v. 20, N. 2.

Kenyon K.W. 1960. Territorial behavior and homing in the Alasca Fur
Seal. “Mammalia”, Т. 24, М. 3 431-444.

Kenyon K.W. 1961. Cuvier beaked whales stranded in the Aleutian
Islands. Journ. Mammal., у. 44, М. 1, 71-76.

Kenyon K.W. 1965. Food of Harbour seal of Amchitka Island, Alasca.
Journ. Mammal., у. 46, М. 1, 103-104. |

969

Kenyon K.W. and Rice D.W. 1961. Abundance and distribution of the
Steller Sea Lion. Journ. Mammal., у. 42, М. 2, 223-234.

Kenyon K.W. and Scheffer V.B. 1955. The seals, sealions, and sea otter
of the Pacific coast. Fish and Wildlife Serv. Depart. of the Interior.
Circul. 32. Washington.

Kenyon K.W. and Wilke F. 1953. Migration of the northern fur seal
Callorhinus ursinus. Journ. Mammal., у. 34, М. 1, 86-98.

Kimura S. and Memoto T. 1956. Note on a Minke Whale kept alive in
aquarium. Norsk Hvalfangst—Tidende, М. 12, 666-674.

King J. 1956. The monk seals (genus Monachus). Bull. Brit. Mus. Nat.
Hist. Zool., N. 3.

King J.E. 1961. Notes on the pinnipeds from Japan described by Tem-
minck in 1844. Zool. Medel., v. 37, N. 13.

King J.E. 1964. Seals of the world. British Mus. London.

King J.E. 1966. Relationships of the Hooded and Elephant seals (genera
Cystophora and Mirounga). Journ. Zool. Proc. Zool. Soc. London,
у. 148, pt. 4, 385-398.

Kjaer Н. 1903. Om de forskjellige Saelarters Forekomst ved vore nordlige
Kyster i vinter. Norsk Fiskeritid, Bd. 8, H.

Knipowitsch N. 1907. Uber die Biologie der Seehunde und Seehundjagd
im europaeischen Eismeer. Rapp. et Proc.—Verb. du Conseil Intern.
pour | Explotatio de la Mer., v. VIII. Copenhague.

Kojima T. 1951. On the brain of the sperm whale (Physeter catodon L.).
Scient. Rep. Whal. Res. Inst., N. 6.

Kowalski-K. (ed.) 1964. Klueze do oznaczania kregowsow polski. Czessc
У Ssaki-Mammalia. Warszawa—Krakow.

Kritzler H. 1952. Observation on the Pilot Whale in captivity. Journ.
Mammal., у. 33, 321-334.

Kruger L. 1959. The thalamus of the Dolphin (Tursiops truncatus) and
comparison with other mammals. Journ. of Compar. Neurology, v. 3,
N. 1, 133-194.

Kumerlowe Н. 1966. Zum Vorkommen der Monchsrobbe Monachus
т. monachus (Hermann 1779) im libanesischen Kustengebiet.
Saugetierkundl. Mitt. Bd. 14, 1172.

Kuroda N. 1938. A list of Japanese mammals. Tokyo.

Kuhn O. 1930. Die Robben der Adria. Zool. Gart. Bd. 2, Leipzig.

Kukenthal W. 1889. Vergleichend—anatomische und Entwicklungs-
geschichtliche Untersuchungen an Waltieren, 1, 2, Jena.

Kukenthal W. 1900. Die Wale der Arctis. Fauna Arctica I, Lief 2, Jena.

692 Kiikenthal W. and Ziehen T. 1893. Uber das Zentralnervensystem der

Wierbeltiere. Denkschr. Med.—naturwiss. Ges. Jena. Bd. 3. 80-198.

970

Laws R.M. 1956a. Determination of age of the largere Whales
(Mystacoceti). Polar Record, N. 8.

Laws R.M. 1956b. Growth and sexual maturity in aquatic mammals.
Nature, у. 178, 193 - 194.

Laws КВ.М. 1959. The foetal growth rate of whales with special refer-
ence to the finwhale Balaenoptera physalus Linn. Discovery Reports,
N. 29, 281 -308.

Laws В.М. and Purves Р.Е. 1956. The ear plug of the Mystacocell as on
indication of age with special reference to the North Atlantic Fin
Whale. Norsk Hvalfangst—Tidende, N. 8, 413-425.

Lay D.M. 1967. A study of the Mammals of Iran. Field Mus. of Nat.
Hist., v. 54, Chicago, USA.

Lagendre В. 1947. Notes biologiques sur les pinnipédes. A propos d’un
Halichoerus grypus (Fabricius) observé wivant a Concarnean. Bull.
Inst. Oceanogr., Monaco, v. 44, N. 907.

Leis М. 1960. Viiger—hulgest Riia lahes. Eesti loodus, М. 1.

Lenfant C. 1969. Physiological properties of blood of marine mammals.
In: Andersen H.T. ed. The Biology of Marine Mammals Academic
Press, New York, London.

Lillie D.G. 1915. Cetacea. British Antractic (“Terra Nova”) Expedition,
1910. Nat. Hist. Rep. Zool., у. 1, М. 3.

Lillie D.G. 1956. The Hood seal (Cystophora cristata). Proc. 5th Tech.

Meeting Int. Un. Prot. Nat.

Lilljeborg W. 1874. Sveriges och Norges Ryggradsjur. I. Daggdjren, Upp-
sala.:

Linstow O. 1905. Helminthen aus Ceylon und aus arctischen Breiten.
Zeit. Wissen. Zool., Jg. 82, 182 - 192.

Lockie J.D. 1962. Grey seals as competitors with man for salmon. The
exploitation of natural animal populations. New York.

Lockley К. 1954. British mammals the Atlantic grey seal. “Oryx”, у. 2,
N. 6.

Lonnberg Е. 1898. Om ishavsfararnes “Trold—Sael”. Ofvers. Kgl.
Vetensk. Akad. Forhlandl. 55.

Lonnberg Е. 1899. Om de Ostersjoen forekommande Sjalhundsarterna
och deras Kannetecken. Svensk Fiskeritidskrift.

Lonnberg E. 1911. Remarks on the dentition of Delphinapterus leucas.
Archiv fur Zool., v. 7, N. 2.

Lonnberg E. 1929. A hybrid between Grey Seal, Halichoerus grypus.
Nills. and Baltic Ringed seal Phoca hispida annelata. Fauna och
Flora. Uppsala.

Lund H.M. 1954. The Walrus and its visite to the Norvegian coast since
1900. Norsk Hvalfangst—Tidende, N. 11, 639-646.

971

Mackintosh М.А. 1947. The Natural History of Whalebone Whales. Ann.
Rep. Smiths. Inst. (for 1946), 235 - 264.

Mackintosh N.A. 1951. Observation on Whales from ships. Nat. Hist.
Okt 1-10!

Mackintosh N.A. 1955. Whale marks recently recovered. Norsk.
Hvalfangst—Tidende, N. 12, 718-719.

Mackintosch N.A. and Brown S.G. 1956. Estimates of the Southern pop-
ulations of large Baleen Whales. Norsk Hvalfangst—Tidende, N. 9,
469 - 480.

Mackintosh N.A. and Whieller J.F. 1929. Southern Blue and Fin Whales.
Discovery Reports, v. 1, 259-539.

McBride A.F. and Kritzler H. 1951. Observations on pregnancy, part-
urition and postnatal behaviour in the bottlenose dolphin. Journ.
Mammal., У. 32, 251-266.

McLaren J.A. 1958. The biology of the ringed seal (Phoca hispida Schre-
ber) in the Eastern Canadian Arctic. Fish. Res. Bd. Canada, Bull.
118.

McLaren J.A. 1958. The economics of seals in the eastern Canadian
arctic. Fish. Res. Bd. Canada, Circular N. 1.

McLaren J.A. 1960. The netting of sea mammals. A report of the Belcher
Islands experiment. Fish. Res. Bd. Canada, Circular N. 6.

McLaren J.A. 1960. On the origin of the Caspian and Baikal seals and
the paleoclimatological implication. Amer. Journ. Sci., v. 258.

McLaren J.A. 1966. Population dynamics and exploitation of seals in
the Eastern Canadian Arctic. The Exploitation of Natural Animal
populations. Oxford.

McLaren J.A. 1958. Some aspects of growth and reproduction of the
Bearded Seal, Erignathus barbatus (Erxleben). Journ. Fish. Res. Bd.
Canada, v. 15, N. 2.

McLaren J.A. 1966. Taxonomy of Harbour Seals of the western North
Pacific and evolution of certain other hair seals. Journ. Mammal.,
v. 47, N. 3.

Maher W.J. and Wilmoysky N.J. 1963. Annual catch of Bowhead Whales
by eskimos of Point Barrow, Alaska, 1928-1960. Journ. Mammal.,
у. 44, М. 1, 16-20.

Mahken T. and Gilmore В.М. 1960. Suckerfish on porpoise. Journ.
Mammal., v. 41, N. 1, 134.

Makino S. 1948. The chromosomes of Dall porpoise with remarks on
the phylogenetic relation of the Cetacea. Chromosoma, Berlin, v. 3,
220. 251,

693

972

Mansfield A.W. 1958. The biology of the Atlantic Walrus Odobenus
rosmarus (Linnaeus) in the eastern Canadian Arctic. Fish. Res. Bd.
Canada. Manuscript Report Series (biological), N. 653.

Mansfield A.W. 1959. The Walrus in the Canadian Arctic. Fish. Res. Bd.
Canada, Circular N. 2, Arctic Unit.

Mansfield A.W. 1963. Seals of Arctic and Easter Canada. Fish. Res. Bd.
Canada, Bull. N. 137.

Mansfield A.W. 1966. The grey seal in eastern Canadian waters. Canad.
Audubon Magazine, XI - XII. Toronto.

Mansfield A. 1967. Distribution of the Harbour Seal, Phoca vitulina Lin.
in Canadian Arctic waters. Journ. Mammal., v. 48, N. 2.

Mansfield A.W. 1970. Population dynamics and exploitation of some
Arctic seals. Antarctic Ecol., v. 1.

Manvill R.H., Joung S.P., 1965. Distribution of Alaskan Mammals. Wash-
ington. 1-74.

Margolis L. 1954. List of the parasites recorded from sea mammals
caught off the West Coast of North America. Journ. Fish. Res. Bd.
Canada, v. II, N. 3, 319-325.

Margolis L. and Pike G. 1955. Some helminth parasites of Canadian
Pacific Whales. Journ. Fish. Res. Bd. Canada, v. 12, N. 1.

Matheson C. 1950. Longevity in the Grey Seal. Nature, v. 166, N. 4210.

Mathisen O.A. 1959. Studies on Steléer sea lion (Eumetopias jubata)
in Alaska. Trans. Twenty-Fourth North Amer. Wildlife Conference.
Washington, 346 - 356.

Mathisen O.A., Baade R.T. and Lopp R.J. 1962. Breeding habits, growth
and stomach contents of the Steller sea lion in Alaska. Journ. Mam-
mal., у. 43, М. 4, 469-477.

Matsuura [. 1935. On the Sperm Whale found in the adjacent waters of
Japan. Bull. Japan. Soc. Scient. Fish., v. 4, N. 2, 101-112.

Matsuura 1. 1936. On the Lesser Rorqual found in the adjacent waters
of Japan (in Japanese). Bull. Japan. Soc. Scient. Fish., v. 4, N. 5,
325 - 330.

Matthews L.H. 1932. Lobster-krill. Discovery Reports, v. 5, 467 - 484.

Matthews L.H. 1938a. The Humpback Whale, Megaptera nodosa. Dis-
covery Reports, 17, 7-92.

Matthews L.H. 1938b. The Sperm Whale—Physeter catodon. Discovery
Reports, 17, 93 - 168.

Matthews L.H. 1938c. Notes on the Southern Right Whale, Eubalaena
australis. Discovery Reports, 17, 169 - 182.

Matthews L.H. 1938d. Sei Whale, Balaenoptera borealis. Discovery
Reports, 17, 185 -289.

694

973

Matthews Г.Н. 1950. The natural history of the Grey Seal, including
lactation. Proc. Zool. Soc. London, c. 120, 763.

Matthews L.H. 1952. British mammals. London.

Matthews L.H. 1966. Comments on the Delphinoidea. In.: Norris K. (ed).
Whales, Dolphins and Porpoises. Univ. of California Press, Berkeley
and Los Angeles.

Matthews L.H. 1958. Calving time of grey seals. Nature, v. 182, N. 4645,
1325 - 1326.

Mercer M.C. 1967. Wintering of Pilot Whales, Globicephala melaena, in
Newfoundland Inshore waters. Journ. Fish. Res. Bd. Canada, v. 24,
N. 11, 2481 - 2484.

Millais J. 1904. The Mammals of Great Britain and Ireland. I.

Millais J. 1904. Exhibition of and remarks upon pelages of the Grey seal
(Halichoerus grypus) in various stages of growth. Proc. Zool. Soc.
London.

Miller G.S. 1923. List of the American recent mammals. Inst. U. S., Nat.
Mus., Bull. 128.

Mitchell E. 1965. Evidens for mass strandings of the Folse Killer Whale
(Pseudorca crassidens) in the eastern north Pacific Ocean. Norsk
Hvalfangst—Tidende, N. 8, 172-177.

Mitchell E. 1966. Faunal succession of extinct North Pacific Marine
mammals. Norsk Hvalfangst—Tidende, N. 3, 47-60.

Mitchell E. and Houck W.J. 1967. Cuvier’s beaked whale (Ziphius
cavirostris) stranded in Northern California. Journ. Fish. Res. Bd.
Canada, у. 24, М. 12, 2503-2513.

Mizue K. 1951a. Grey Whales in the east sea area of Korea. Scient. Rep.
Whal. Res. Inst., N. 5, 71-79.

Mizue K. and 1951b. Food of Whales of the adjacent waters of Japan.
Scient. Rep. Whal. Res. Inst., N. 5, 81-90.

Mizue K. and Jimbo H. 1950. Statistic study of foetuses of whales. Scient.
Rep. Whal. Res. Inst., N. 3.

Mizue K. and Murato T. 1951. Biological investigation on the whales
caught. Ibid., N. 6.

Mizue K. and Joshida K. 1965. On the porpoises caught by the salmon
fishing gill—net in Bering Sea and the north Pacific Ocean. Bull.
Fac. Fish. Nagasaki Univ., N. 19, 1-36.

Mizue K., Joshida K. and Takemura A. 1966. On the ecology of the
Dall’s porpoises in Bering Sea and the north Pacific Ocean. Bull.
Fac. Fish. Nagasaki Univ., N. 21, 1-21.

Miyazaki I. 1954. Survey of Whaling operations from Land Stations in
waters adjacent to Japan in 1953. Norsk Hvalfangst—Tidende, N. 4,
181 - 192.

974

Miyazaki I. 1955. Survey of Whaling operations from Land Stations in
Japan in 1954. Norsk Hvalfangst—Tidende, М. 4, 189-200.
Miyazaki I. 1956. Survey of Whaling operations from Land Stations in

Japan in 1955. Norsk Hvalfangst—Tidende, М. 5, 252 - 264.

Miyazaki [. 1958. Survey of Whaling operations from shore based sta-
tions in Japan 1956 and 1957. Norsk Hvalfangst—Tidende, N. 12,
597 - 606.

Mohr E. 1941. Ein neuer westpazifischer Seehund. Zool. Anz., Bd. 133,
N. 3/4.

Mohr E. 1952. Beitrag zur kenntnis der Mahnenrobben. Zool. Garten

Bd. 19, H. 2/4, 98-112.

Mohr E. 1952. Die Robben der Europaischen Gewasser. Monographien
der Wildsaugetiere, 12. Frankfurt am Main.

Mohr E. 1955. Der Seehund. Die neue Brehm—Bucherei. Wittenberg
Lutherstadt.

Mohr E. 1965. Uber Phoca vitulina largha Pallas 1811 und weissgeborene
Seehunde. Zeit. f. Saugetierk., Bd. 30, Н. 5.

Mohr E. 1963. Beitrage zur Naturgeschichte der Klappmutze Cystophora
cristata Erxl. 1777. Zeit. f. Saugetiekunde, Bd. 28, H. 2.

Monticelli F.S. 1889. Elenco degli Elminti raccolti dal Capitano
Chierchia durante il viaggio di circumnavigazione della R. corvetta
“Vettor Pisani’. Boll. Soc. Nat., T. 3, N. 1, 67-71, Napoli.

Moore J.C. 1960. New records of the Gulf—stream beaked whale Meso-
plodon Gervais and some taxonomic considerations. Amer. Mus.
Novitates, 1933, 1-35.

Moore J.C. 1968. Relationships among the living genera of beaked whales
with classifications, diagnoses and keys. Fieldiana Zoology, v. 53,
М. 4, 209-298. Field Museum of Natural History.

Morton B.J. 1962. Seasonal occurrence and movement of seals in the
north-west Britain. Proc. Zool. Soc. London, v. 138, N. 3.

Morton B.J., Lockie J.D. and Hewer H.R. 1962. The breeding colony of
grey seals on North Rona, 1959. Proc. Zool. Soc. London, v. 138,
N. 2.

Mohl В. 1968. Hearing in seal. Behaviour and physiology of pinnipeds.
New York.

Mohl B. 1968. Auditory sensitivity of the common seal in air and water.
Jour. And. Res., 8.

Moller D., Naevdal G. og Valen A. 1966. Rapport om arbeidet med
blodanalyser for populasjonsundersokelser. Fisken og Havet, М. 2.

Murie O.J. 1959. Fauna of Aleutian islands and Alaska peninsula. Wash-
ington. 1 - 406.

695

975

Mursaloglu В. 1964. Occurrence of the Monk seal on the Turkish coasts.
Journ. Mammal., v. 45, N. 2, p. 316.

Muller-Wille L.L. 1969. Biometrical comparison of four populations of
Phoca hispida Schreb. in the Baltic and White Seas and Ladoga and
Saimaa. Commentationes Biological Soc. Scient. Fennica, v. 31, N. 3,
Helsinki.

Myers B.J. 1959. The stomach contents of harp seals Phoca groenlandica
(Erxl.) from the Magdalen Islands, Quebec. Canad. Journ. Zool.,
у 37, №. 3:

Naevdal С. 1965. Protein polymorphism used for identification of Harp
seal Populations. Arbok for Universitetet i Bergen. Mat.—naturv.
serie, N. 9, Bergen—Oslo.

Naevdal G. 1969. Blood protein polymorphism in Harp Seals off Eastern
Canada. Journ. Fish. Res. Bd. Canada, у. 26, 1397 - 1399.

Nagasaki F. 1961. Population study on the fur seal herd. Tokai Reg. Fish.
Res. Lab., Special Publ. N. 7, 1-60. Tokyo.

Nagasaki F. and Matsumoto K. 1957. Homing tendency of the bachelor
Seals to their Native Rookeries. Bull. Tokai Reg. Fish. Res. Lab.,
N. 18, 15-19.

Nakajima M., Sawaura K. and Oda T. 1963. On the food quantity of the
northern fur seal. Dobutsuen Suizokuken Zani. No. 12; 4-10. (in
Japanese).

Nansen F. 1924. Blant sel og bjorn. Kristiania.

Nehring А. 1886. Uber die Robben der Ostsee namentlich die Ringel-
roble. Sitz.—Bericht. Naturf. Fr. Berlin.

Nehring A. 1887. Veber das Gefangen der Kegelrobbe. Zool. Garten, 28.

Nemoto T. 1955. White scars on whales Lampery Marks. Scient. Rep.
Whal. Res. Inst., N. 10, 6978.

Nemoto T. 1956. Note on a Minke Whale kept alive in aquarium. Scient.
Rep. Whal. Res. Inst., М. 11, 181-184.

Nemoto T. 1957. Food of Baleen Whales in the Northern Pacific. Scient.
Rep. Whal. Res. Inst., N. 12, 33-89.

Nemoto T. 1959. Food of Baleen Whal. with reference to the whale
movements. Scient. Rep. Whal. Res. Inst., М. 14, 149-290.

Nemoto T. 1964. School of Baleen Whales in the feeding areas. Scient.
Rep. Whal. Res. Inst., N. 18, 89-110.

Nemoto T. and Kasuya T. 1965. Foods of Baleen Whales in the Gulf
of Alaska of the North Pacific. Scient. Rep. Whal. Res. Inst., N. 19,
45-61.

Nemoto T. and Nasu K. 1963. Stones and other aliens in the stomachs
of Sperm Whales in the Bering Sea. Scient. Rep. Whal. Res. Inst.,
N. 17.

976

Nishimoto S., Tozawa М. and Kawakami Т. 1952. Food of Sei Whales
(Balaenoptera borealis) caught in the Bonin Islands waters. Scient.
Rep. Whal. Res. Inst., N. 7, 79-85.

Nishiwaki M. 1950. On the Body weight of Whales. Scient. Rep. Whal.
Res. Inst., N. 4, 184-209.

Nishiwaki M. 1952. On the age determination of Mystacoceti Chiefly
Blue and Fin Whales. Scient. Rep. Whal. Res. Inst., М. 7, 88-119.

Nishiwaki M. 1955. On the sexual Maturity of the Antarctic Male Sperm
Whale (Physeter catodon L.). Sc. Rep. Whal. Res. Inst. N. 10;

‚ 143-151.

Nishiwaki М. 1956. On the sexual Maturity of the Sperm Whale (Physeter
catodon) found in the North Pacific. Scient. Rep. Whal. Res. Inst.,
№ 11. 39-47:

Nishiwaki М. 1957. Age characteristics of ear pluge of Whales. Scient.
Rep. Whal. Res. Inst., М. 12, 23-32.

Nishiwaki M. 1959. Humpback Whales in Ryukynan waters. Scient. Rep.
Whal. Res. Inst., N. 14, 49-87.

Nishiwaki М. 1964. Revision of the article ““Taxonomical consideration
on genera of Delphinidae” in N. 17. Scient. Rep. Whal. Res. Inst.,
М. 18, 171-172.

Nishiwaki М. 1965. Whales and Pinnipeds. Univ. of Tokyo Press. 440 р.
(in Japanese).

Nishiwaki M. 1966. Distribution and migration of the Larger Cetaceans
in the North Pacific as shown by Japanese Whaling Results. In. Nar-
ris K ed. Whales, Dolphins and Porpoises, N. 10, Univ. of Calif.
Press.

Nishiwaki M. 1966. Distribution and migration of marine Mammals in
the North Pacific Area. Symposium N. 4 of the Eleventh Pacific
Science Congress.

Nishiwaki M. 1966. A discussion of rarities among the smaller Cetaceans
caught in Japanese waters. In: Norris K. ed. Whales, Dolphins and
Porpoises. Univ. Calif. Press. Berkeley and Los Angeles, 192 - 204.

Nishiwaki M. 1967. A discussion of rarities among the smaller cetaceans
caught in Japanere Water. Ocean. Res. Inst., Univ. Tokyo. Coll.
Repr. 5, p. 125 - 137.

Nishiwaki M. 1967. Distribution and migration of marine mammals in
the North Pacific Area. Bull. Ocean Res. Inst. Univ. of Tokyo, N. 1,
December.

Nishiwaki M. 1974. Additional data an the distribution and migration of
maine mammals in the North Pacific area.

696

977

Nishiwaki М. and Handa С. 1958. Killer whales caught in the coastal
waters off Japan for recent 10 years. Scient. Rep. Whal. Res. Inst.,
№13, 585 96:

Nishiwaki М. and Hibiya Т. 1951. On the sexual maturity of the Sperm
whale (Physeter catodon) found in the adjacent waters of Japan (2).
Scient. Rep. Whal. Res. Inst., N. 7, 121-124.

Nishiwaki M., Hibiya.T. and Kimura S. 1954. On the sexual maturity of
the Sei Whale on the Bonin waters. Scient. Rep. Whal., Res. Inst.,
№9165 177:

Nishiwaki M., Hibiya T. and Ohsumi S. 1958. Age studies of Fin Whale
based on ear plugs. Scient. Rep. Whal. Res. Inst., М. 13, 155 - 169.

Nishiwaki M. and Kamiya T. 1958. A beaked whale Mesoplodon stranded
at Oiso Beach, Japan. Scient. Rep. Whal. Res. Inst., N. 13, 53-83.

Nishiwaki M. and Kamiya T. 1959. Mesoplodon stejnegeri from the coast
of Japan. Ibid., М. 14, 35-48.

Nishiwaki M. and Kasuya T. 1970. Argeenland right whale caught at
Osaka Bay. Scient. Rep. Whal. Res. Inst., М. 22, 45 -62.

Nishiwaki M. and Nagasaki F. 1960. Seals of the Japanese coastal waters.
“Mammalia”, у. 24, М. 3, 460-464.

Nishiwaki M., Ohsumi S. and Maeda J. 1963. Change of form in the
Sperm Whale accompanied with growth. Scient. Rep. Whal. Res.
Inst., N. 17, 1-14.

Nishiwaki M. and Yagi T. 1953. On the age and growth of teeth in
a Dolphins (Prodelphinus caeruleoalbus). Scient. Rep. Whal. Res.
Inst., М. 8, 133 - 146.

Nordman А. 1840. Observations sur la faune pontique. Voyage dans la
Russie meridionale et la Crimee par la Hongrie, la Valachie et la
Moldavie de M.A. Demidoff, v. 3, Paris.

Nordqvist O. 1899. Beitrag zur Kenntniss der isolierten Formen der
Ringelrobbe (Phoca foetida Fabr.). Acta Societatis pro Fauna et
Flora Fennica, Т. XV, №. 7.

Norris K.S. 1964. Some problems of echolocation in cetaceans. Marine
BioAcoustics. N. Y.

Ogawa T. and Arifuku S. 1948. On the acoustic system in the cetacean
brains. Scient. Rep. Whal. Res. Inst., N. 2. 1-20.

Ohsumi S. 1960. Relative growth of the Fin Whales Balaenoptera
physalus (Linn.). Scient. Rep. Whal. Res. Inst., М. 15, 17-84.

Ohsumi S. 1964. Examination on age determination of the Fin Whales.
Scient. Rep. Whal. Res. Inst., N. 18, 49-88.

Ohsumi S. 1965. Reproduction of the Sperm Whale in the North-West
Pacific. Scient. Rep. Whal. Res. Inst., N. 19, 1-35.

978

Ohsumi 5., Nishiwaki М. and Hibiya Т. 1958. Growth of Fin Whales in
the North Pacific. Scient. Rep. Whal. Res. Inst., М. 13, 97 - 133.
Ohta K., Watarai T., Oishi T., Ueshiba Y., Hirose S., Yoshizawa Т.
1955. Composition of Fin Whale milk. Scient. Rep. Whal. Res. Inst.,

№ 19

Ohta K., Watari Т., Ueshiba Y., Hirose S. and Yoshizawa Т., Akikusa Y..,
Sato M. and Okano H. 1955. Composition of fin whale milk. Scient.
Rep. Whal. Res. Inst., М. 9, 151-167.

Okutani T. and Nemoto T. 1964. Squids as the food of Sperm Whales
in the Bering Sea and Alaskan Gulf. Scient. Rep. Whal. Res. Inst.,
N. 18.

Olds J.M. 1950. Notes on the hood seal (Cystophora cristata). Journ.
Mammal., 31, 450 - 452.

Olsen O.W. 1958. Hookworms Uncinaria lucasi Stiles, 1901, in the Fur
Seals, Callorhinus ursinus (Linn.) on the Pribilof Islands. Trans.
Twenty-Third North Amer. Wildlife Conference, Wildlife Manage-
ment Inst. Washington, 152 - 175.

Omura H. 1950a. On the Body weight of Sperm and Sei Whales loceted
in the adjacent waters of Japan. Scient. Rep. Whal. Res. Inst., N. 4,
1-14.

Omura H. 1950b. Whales in the adjacent waters of Japan. Scient. Rep.
Whal. Res. Inst., N. 4, 27-115.

Omura H. 1953. Biological study on Humpback Whales in the Antarctic
whaling areas ГУ and У. Scient. Rep. Whal. Res. Inst. М. 8, 81 - 102.

Omura H. 1955. Whales in the Northern Part of the North Pacific. Scient.
Rep. Whal. Res. Inst., М. 7, 395 - 405.

Omura H. 1957a. Osteological study of the Little Piked Whale from
coast of Japan. Scient. Rep. Whal. Res. Inst., N. 12, 1-22.

Omura H. 1957b. Report of two right whales caught off Japan for scien-
tific purposes. Norsk. Hvalfangst—Tidende, М. 7. 374-390.

Omura H. 1958a. North Pacific Right Whale. Scient. Rep. Whal. Res.
05. №. 13. 1-52.

Omura Н. 19585. Note on embryo of Baird’s Beaked Whale. Scient. Rep.
Whal. Res. Inst., N. 13, 213-215.

Omura H. 1959. Bryde’s whale from coast of Japan. Scient. Rep. Whal.
Res. Inst., N. 14, 1-33.

Omura H. 1962. Further information on Bryde’s Whale from the coast
of Japan. Scient Rep. Whal. Res. Inst., N. 16, 7-18.

Omura H. and Fujino K. 1954. Sei Whale in the adjacent water of Japan.
2. Further studies of the external characters. Scient. Rep. Whal. Res.
Inst., N. 9, 89-103.

697

979

Omura H., Fujino K. and Kimura S. 1955. Beaked Whales Berardius
bairdi of Japan with notes on Ziphius cavirostris. Scient. Rep. Whal.
Res. Inst., М. 10, 89-133.

Omura H. and Kawakami T. 1956. Japanese whale marking in the North
Pacific. Norsk Hvalfangst—Tidende, М. 10, 555-563.

Omura H., Maeda K. and Miyazaki I. 1953. Whaling in the adjacent
waters of Japan. Norsk Hvalfangst—Tidende, N. 4, 199-212.

Omura H., Nemoto T. 1955. Sei Whale in the adjacent waters of Japan. 3.
Relation between movement and water temperature of sea.—Scient.
Rep. Whal. Res. Inst., М. 10, 79-87.

Omura H., Nishiwaki M. Ichihara T. and Kasuya T. 1962. Osteological
note of a Sperm Whale. Scient. Rep. Whal. Res. Inst. М. 16, 35 - 46.

Omura H., Ohsumi S., Nemoto T., Nasu K. and Kasuya T. 1969. Black
Right Whales in the North Pacific. Scient. Rep. Whal. Res. Inst.,
№21, 676.

Omura H. and Sakiura Н. 1965. Studies on the Little Piked Whale from
the coast of Japan. Scient. Rep. Whal. Res. Inst., N. II, 1-37.

Orr R.T. 1953. Beaked Whale (Mesoplodon) from California, with com-
ments of taxonomy. Journ. Mammal., v. 34, N. 2, 239-249.

Ottestad P. 1956. On the size of the stock of Antarctic Fin Whales
relative to the size of the catch. Norsk Hvalfangst—Tidende, N. 6,
298 - 308.

Palmer R.S. 1944. Walrus remains from Northern New England. Journ.
Mammal., У. 25, М. 2.

Pedersen А. 1931. Fortgesetzte Beitrage zur Kenntnis der Saugetier
und Vogelfauna der Ostkuste Gronlands, Meddelelser om Gronland,
B. 77. Kobenhaven.

Peters М. 1938. Der neue deutsche Walfang. Verl. “Hansa”.

Petersom R.S., Le Boeuf B.J. and Delong R.L. 1968. Fur seals from the
Bering Sea breeding in California. Nature, v. 219, N. 5157, 899-901.

Pike С.С: 1950a. Stomach contents of Humpback Whales from the coast
of British Columbia. Journ. Fish. Res. Bd. Canada, v. 10, N. 6.

Pike G.C. 1950b. What the whales eat? Canadian Fisherman, v. 37, N. 9,
Hes

Pike G.C. 1953. Two records of Berardius bairdi from the coast of British
Columbia. Journ. Mammal., у. 34, М. 1, 98- 104.

Pike G.C. 1958. Whaling on the coast British Columbia. Norsk
Hvalfangst—Tidende, N. 3, 117-127.

Pike G.C. 1958. Food of the Northern Sea Lion. Fish. Res. Bd. Canada.
Progress Reports of the Pacific Coast Stations. N. 12, 18-20.

Pike G.C. 1962. Migrations and feeding of the Grey Whale (Eschrichtius
gibbosus). Journ. Fish. Res. Bd. Canada, у. 19, М. 5, 815-838.

980

Pike C.G. and Giovando L. 1963. Whales and dolphins of the West coast
of Canada. Fish. Res. Bd. Canada, Circ. N. 168, 15p.

Pike G.C. and Maxwell B.E. 1958. The abundance and distribution of
the northern sea lion (Eumetopias jubata) on the coast of British
Columbia. Journ. Fish. Res. Bd. Canada, v. 15, 5-17.

Pilleri G. and Gihr M. 1969. On the anatomy and behaviour of Risso’s
dolphin. In: Pilleri G. ed. Investigations on Cetacea. Vol. 1, 74-93.
Berne.

Pocock R.J. 1911. The hair of the “puppy coat” of a Grey Seal.—Proc.
Zool. Soc. London, v. 11.

Pocock R.J. 1933. The presence of the external pinna in the grey seal
(Halichoerus grypus). Proc. Zool. Soc. London.

Ponomarenko V.P. 1968. Some data on the distribution and migration of
Polar Cod in the seas of the Soviet Arctic. Conseil Perman. Intern.
pour РЕхрюга+. de la Mer. Rapp. et Proc.—Verb., v. 158, 131 - 135.

Price E.W. 1932. The Trematodes parasites of marine mammals. Proc.
U. $. Nat. Mus., у. 81, М. 13, 1-68.

Priemel R. 1909. Die Geburt einer Kegelrobbe. Zool. Garten, 50.

Purves P.E. 1955. The wax plug in the external auditory meatus of the
Mysiceti. Discovery Reports, v. 27, 293 - 302.

Qritsland Т. 1959. Klappmyss. “Fauna”, М. 2.

@ritsland Т. 1960. Flyleting etter klappmyss pa fangstfeltet 1 Danmark-
stredet. “Fauna”, N. 4. |

ОгиЗапа Т. 1964. Klappmysshunnens for plantningsbiologi. Fisken og
Havet, N. 1. Bergen.

@stby Н. 1956. Catch of small Whales 1955. Norsk Hvalfangst—Tidende,
М. 5, 241-249.

@stby H. 1957. Catch of small Whales 1956. Norsk Hvalfangst—Tidende,
М. 6, 300-308.

@Ostby Н. 1958. Catch of small Whales 1957. Norsk Hvalfangst—Tidende,
М. 8, 377-385.

@stby Н. 1959. Catch of small Whales 1958. Norsk Hvalfangst—Tidende,
№: 10, 521 531.

@ynes Р. 1964. Sel ра norskekysten fra Finmark Ш Mgre. “Fisket Gang”,
50.

@ynes Р. 1966. Sel i Sor—Norge. “Fisket Gang”, М. 45.

Rae B.B. 1960. Seals and Scottish Fisheries. H.M. Stationary Office.
London and Edinburg.

Rae B.B. 1962. The effect of seal stocks in relation to scottish marine
fisheries. The Exploitation of natural animal populations. New York.

Rae B.B. and Shearer W.M. 1965. Seal damage to salmon fisheries. Dep.
Agricult. Fish. for Scotland. Mar. Res. N. 2.

698

981

Railliet А. 1899. Sur quelques parasites rencontres а l’autopsie d’un
Phoque (Phoca vitulina L.). С. К. Soc. Biol., 128 - 130.

Ramsey D.H. 1968. Diurnal fluctuations in censuses of migrating Cali-
fornia Grey Whales. Norsk Hvalfangst—Tidende, N. 5, 101-105.
Ransom B. 1920. Synopsis of the Trematode family Heterophyidae with
descriptions of a new genus five new species. Proc. U. S. Nat. Mus.,

v. 157, 527-573. Washington.

Rasmussen B. 1957. Exploitation and protection of the East Greenland
seal herds. Norsk Hvalfangst—Tidende, N. 2.

Rasmussen B. 1960. Om Klappmysse bestanden i det Nordlige atlanter-
hav. Havforsningsinstitutt, N. 1.

Rasmussen B. 1962. Klappmyssens alderfordeling. Danmark. Fiskets
Gang., М. 5.

Rasmussen B. and Oritsland T. 1964. Norwegian Tagging of Harp Seals
and Hooded Seals in north Atlantic waters. Fiskeridirektoratets
Skrift., v. 13, N. 7.

Ray G. 1961. White whales from the aquarium. Anim. Kingdom, 64,
162-170:

Rice D.W. 1961. Census of the California Grey Whale, 1959/60. Norsk
Hvalfangst—Tidende, М. 6, 219-225.

Rice D.W. 1963. Progress Report on Biological Studies of the larger
Cetacea in the water of California. Norsk Hvalfangst—Tidende, N. 7,
181 - 187.

Rice D.W. 1965. Offshore southward migration of Grey Whales off
southern California. Journ. Mammal., v. 46, N. 3, 504-505.

Rice D.W. 1968. Stomach contents and feeding behavior of Killer Whales
in the Eastern North Pacific. Norsk Hvalfangst—Tidende, N. 2,
35 - 38.

Rice D.W. and Wolman A.A. 1971. The life history and ecology of the
Grey Whale (Eschrichtius robustus). Special publ. N. 3. The Amer-
ican Society of Mammalogist. 142 p.

Ridgway S.H. 1966. Dall porpoises, Phocoenoides dalli (True) obser-
vation in captivity and at sea. Norsk Hvalfangst—Tidende, N. 5,
97-110:

Ridgway S.H. Ed. 1972. Mammals of the sea Springfield.

Risting S. 1928. Whales and Whale Foetuses. Rapp. Cons. Explor. Mer,
v. 50. Copenhagen.

Robinson C.G. 1897. Ice-riding pinnipeds. A description of the migration
and peculiarities of Phoca grenlandica and Cystophora cristata.
Roest, A. 1964. Ribbon seal from California. Journ. Mammal., v. 45,

М. 3, 416-420.
Romer А. 1966. Vertebrate Paleontology.

982

Ronald K., Johnson E., Foster М. and Vander Р.О. 1970. The Harp seal
Pagophilus groenlandicus (Erxleben, 1777). I. Methods of handling,
molt and disecases in captivity. Canad. Journ. of Zool., v. 48, N. 5.

Ropelewski A. 1952. Ssaki Baltycru. Krakow.

Roppel A.Y. and Dakey S.P. 1965. Evolution of fur seal management
on the Pribilof Islands. Journ. Wildlife Management, v. 29, N. 3,
448 - 463.

Rudolphi C.A. 1819. Entosoorum synopsis, cul accedunt mantissa duplex
et indices locupletissimi. 811 p. Berolini.

Ruud J.T. 1973a. Seihvalen, Balaenoptera borealis (Lesson) og Brydeh-
valen, Balaenoptera brydei (Olsen). Norsk Hvalfangst—Tidende,
М. 5, 145 - 150.

Ruud J.T. 19376. Grenlandvales (Balaena mysticetus L.). Norsk
Hvalfangst—Tidende, М. 7, 254-257.

Ruud J.T. 1937c. Finnhvalen, Balaenoptera physalus (Linne) Norsk
Hvalfangst—Tidende, М. 3, 98 - 112.

Ruud J.T. 19374. Knolhvalen, Megaptera nodosa (Bonnaterre). Norsk
Hvalfangst—Tidende, N. 4, 113-121.

Ruud J.T. 1937e. Vagenhvalen, Balaenoptera acutorostrata (Lacepede).
Norsk Hvalfangst—Tidende, М. 6, 193 - 194.

Ruud J.T. 1952. Catches of Bryde Whale off French equatorial Africa.
Norsk Hvalfangst—Tidende, М. 12, 662 - 663.

Ruud J.T. and @ynes P. 1959. Whale Marking carried out by Norwe-
gian catchers in the Antarctic, seasons 1956/57 and 1957/58. Norsk.
Hvalfangst—Tidende, N. 2, 57-62.

Saemundsson В. 1939. The zoology of Iceland. Mammalia. у. IV.

Scheffer V.B. 1945. Growth and behavior of young sea lion. Journ. Mam-
mal., у. 26, 390-392.

Scheffer V.B. 1949. Notes on three Beaked Whales from the Aleutian
Islands. Pacific Science, v. 3, N. 4.

Scheffer V.B. 1950. The striped Dolphin, Lagenorhynchus obliquidens
Gill, 1865, on the coast of North America. Amer. Midland Naturalist,
у. 44, М. 3.

Scheffer V.B. 1950a. Growth layers on the teeth of Pinnipeds as an indi-
cation of age. Science, v. 112, 309-311.

Scheffer V.B. 1950b. The food of the Alaska fur seal. Wildlife Leaflet
329. Washington, 1 - 16.

Scheffer V.B. 1958. Seals, Sea Lions and Walruses. A Review of the
Pinnipedia. Stanford Univ., California.

Scheffer V.B. 1962. Pelage and surface topography of the Northern fur
seal. Nat. Amer. Fauna, N. 64.

699

983

Scheffer У.В. 1963. Record of ringed from seals the Pribilof Islands.

Journ. Mammal., v. 43, N. 3.

Scheffer V.B. and Jonson A:M. 1963. Molt in the northern fur seal.
Fish and Wildlife Service, Spec. Scient. Rep. Fish. N. 450, 1-24.
Washington.

Scheffer V.B. and Slipp J.M. 1944. The Harbour seal in Washington
State. Amer. Mid]. Natural., у. 32.

Scheffer V.B. and Slipp J.M. 1948. The whales and dolphins of Wash-
ington State, with key of the cetaceans of the west coast of North
America. Amer. Midland Natural., у. 39, М. 2, 257-337.

Scheffer T. and Sperry C. 1931. Food habits of the Pacific Harbour seal,
Phoca richardii. Journ. Mammal., v. 12, N. 3.

Schevill W.E. 1956. Lagenorhynchus acutus off Cape Cod. Journ. Mam-
mal. v. 37, N. 1, 128-129.

Schiller E. 1954. Unusual walrus mortality on St. Lawrence Island,
Alaska. Journ. Mammal., У. 35, М. 2, 203-210.

Schapp B., Helwing 5. and Chizelea С. 1962. Contributions to the knowl-
edge of the Black Sea seal (Monachus monachus) Herm. Trav. du
Mus. D’Hist. Nat. “Gr. Antipa”, у. Ш.

Schneider R. 1963. Der Kehlkopf der Klappmutze. Anat. Anz., B. 112,
54 - 68.

Schubart O. 1929. Die Seehunde der Ostsee und ihr Fang. Zool. Garten,
Bd. 1, H. 7/9.

Schwarz E. 1942. The harbour seal of the western Pacific. Journ. Mam-
mal., v. 23.

Scott R.F., Kenyon K.W., Buckley J.L. and Olsen S.T. 1959. Status and
management of the polar bear and Pacific Walrus. Trans. Twenty-
Fourth North Amer. Wildlife Conference. Washington.

Selfangsten. Utgitt av Fiskeridirektoren. Arberetning ved Kommende
Norges Fisheries, N. 10.

Sergeant D.E. 1953. Whaling in Newfoundlend and Labrador water.
Norsk Hvalfangst—Tidende, N. 12, 687-695.

Sergeant D.E. 1959a. Age determination in Odontocete Whales from
dentinal growth layers. Norsk Hvalfangst—Tidende, М. 6, 273 - 288.

Sergeant D.E. 1959b. Dolphins in Newfoundlend waters. Norsk
Hvalfangst—Tidende, М. 11, 562 -568.

Sergeant D.E. 1962a. The biology and hunting of beluga or White Whales
in Canadian Arctic. Fish. Res. Bd. Canada, Circ. N. 8.

Sergeant D.E. 1962b. The biology of the Pilot or Pothead Whale Glo-
bicephala melaena (Traill) in Newfoundlend waters. Fish. Res. Bd.
Canada, Bull. N. 13, 85 p.

984

Sergeant Р.Е. 1963. Harp seals and the sealing industry. Canad.
Audubon, v. 25, N. 2.

Sergeant D.E. 1965. Migrations of Harp Seals, Pagophilus groenlandicus
(Erxleben) in the Northwest Atlantic. Journ. Fish. oo ‘Bd. Canada,
у. 22, М. 2, 433 - 464.

Sergeant D.E. 1965. Exploitation and conservation of ua and Hood
Seals. Polar Record, v. 12, N. 80, 541-551.

Sergeant D.E. 1966. Reproductive rates of Harp Seals, Pagophilus
groenlandicus (Erxleben). Journ. Fish. Res. Bd. Canada, v. 23, N. 5,
757 - 766.

Sergeant D.E. and Fisher H.D. 1957. The smaller cetacea of Eastern
Canadian waters. Journ. Fish. Res. Bd. Canada, v. 14, N. 1, 83-115.

Sergeant D.E. and Fisher H.D. 1960. Harp seal populations in the west-
ern north Atlantic from 1950 to 1960. Journ. Fish. Res. Bd. Canada,
Arctic Unit, Circ. N. 5.

Sergeant D.E. 1963. Minke whales, Balaenoptera acutorostrata Lace-
pede, of the western North Atlantic. Journ. Fish. Res. Bd. Canada,
у. 20, М. 6, 1489 - 1504.

Shearer W.M. 1962. Seals and salmon nets. The Exploitation of Natural
Animals Populations (A Symposium of Brit. Ecologic. Society). New
York.

Siebenaler I.B. and Caldwell D.K. 1956. Cooperation among adult Dol-
phins. Journ. Mammal., у. 37, М. 1, 126-128.

Siebold C. und Stannius 1846. Lehrbuch der vergleichenden Anatomie.
H. Stanins. Th. 2. Wirbelthiere verlag von Veit & Comp. Berlin;
Buch 4: Die Saugethiere. 339 - 482.

Siivonen Г. 1967. Pohjolan Nisakkaat, р. 1-181. Helsinki.

Siivonen L. 1968. Nordeuropas daggdjur. р. 4- 183. Stockholm.

Simpson G.C. 1945. The principles of classification and a classification
of mammals. Bull. Amer. Mus. Nat. Hist., v. 85.

Sivertsen E. 1941. On the biology of the Harp Seal, Phoca groenlandica
Erxl., investigations carried out in the White Sea 1925-1937.
Hvalradets Skrifter N. 26. Oslo.

Slijper E.J. 1936. Die Cetaceen. Capita Zool., v. VII, pt. 1-11.

Slijper E.J. 1955. Geluiden van walvissen en dolphin. Vakbl. biol. v. 35,
N. 12.

Slijper E.J. 1958. On the vascular system of Cetacea. XVth Intern.
Congress Zool., Sect. Ш.

Slijper E.J. 1958. Organ weights and symmetry problems in porpoises
and seals, Archives Nedelandises de Zool. T. 12, 97-113.

Slijper E.J. 1960. Aus dem Leben der Wale. Verh. Naturf. Ges. Basel.
Bd. 71, М. 1. 140-148.

700

985

Slijper E.J. 1962. Whales. London. Hutchinson.

Slijper E.J. 1966. Functional morphology of the reproductive system in
Cetacea. In.: Norris K. ed. Whales, Dolphins and Porpoises. Univ.
of Calif. Press. Berkeley and Los Angeles.

Slijper E.J. 1967. White Whales in Netherlands waters. Z. f. Saugetier-
kunde, Bd. 32, H. 2, 86-89.

Slijper E.J., Heinemann. 1969. Uberfamilie Narwalartige. B.: Grzimeks
Tierleben. 11, p. 486-489.

Slijper EJ. and Utrecht W.L. van 1959. Observing whales. Norsk
Hvalfangst—Tidende, N. 3, 101-117.

Slipp J. and Wilke F. 1953. The Beaked Whale Berardius on the Wash-
ington coast. Journ. Mammal., v. 34, М. 1, 105-113.

Smith E.A. 1962. Trackigg the Grey Seals. Nat. Hist., v. 71, N. 3.

Soper J.D. 1944. The mammals of Southern Baffin Island, Northwest
Territories, Canada. Journ. Mammal., v. 25, N. 3, 221-254.

Spalding D.J. 1964a. Age and growth of female sea lions in British
Columbia. Journ. Fish. Res. Bd. Canada, v. 21, N. 2.

Spalding D.J. 1964b. Comparative feeding habits of the fur seal sea lion
and harbour seal on the British Columbia coast. Journ. Fish. Res.
Bd. Canada, Bull. N. 146, 1-52.

Starrett A. and Starrett P. 1955. Observations of young blackfish Globi-
cephala. Journ. Mammal., v. 36, N. 3, 424-429.

Steenstrup J. 1860. Maelketansaettet hos Reunesaelen, Svartsiden og
Fjordsaellen (Phoca barbata, Ph. groenlandica og Ph. hispida) Vid.
Meded. fra den Natur. Forening i Kjobnhaven.

Steven G.A. 1934. A short investigation into the habits, abundance and
species of seal on the north Cornwall coast. Journ. Mar. Biol. Ass.
О. К., у. 19, 489-501.

Steven С.А. 1936. Seals (Halichoerus grypus) of Cornwall coast. Journ.
Mar. Biol. Ass. U. K., v. 20, 493-506.

Stiles C.W. and Hassall A. 1899. Internal parasites of the Seal. In: Jordan,
David, Starr et al. The Fur Seal Islands of the North Pacific Ocean.
Pt. 3. 99-177. Washington.

Stutz S. 1966. Foetal and postpartum white coat pelage in Phoca vitulina.
Journ. Fish. Res. Bd. Canada, v. 23, N. 4.

Stutz S. 1967. Pelage patters and population distributions in the Pacific
Harbour Seal (Phoca vitulina richardi). Journ. Fish. Res. Bd. Canada,
v. 24, N. 2, 451-455.

Sundstrom K.-G. 1962. Grasael i norra Ostersjon. Sver. Natur. Arbok,
Aroj. 53. Stockholm.

Tanaka S. 1956. Estimation of Whale population on the Basis of tagging
Experiments. Bull. Japan. Soc. Scient. Fish., v. 22, N. 6. 330-337.

986

Tavolga М.С. and Essapian Е.С. 1957. The behavior of the bottlenosed
dolphin (Tursiops truncatus). Zoologica, v. 42, N. 1, 11-31.

Taarra T. 1950. On the respiratory pigments of whale (Studies on whale
blood, II). Scient. Rep. Whal. Res. Inst., N. 3, 96-101.

Taylor F.H. C., Fujinaga M. and Wilke F. 1955. Distribution and food
habits of the fur seals of the North Pacific Ocean. Report of Coop-
erat. Investigat, by the Governments of Canada, Japan and United
States of America, February - July, 1952, 1-86. Washington.

The Norwegian Small Whale Hunting in the Seasons 1953 and 1954.
Norsk Hvalfangst—Tidende N. 3, 135 - 138.

The Norwegian Small Whale Fisheries 1966. 1967. Norsk Hvalfangst-
Tidende N. 1, 9-15.

Thompson W. 1939. The “Horn” of the narwhal. BHUPO.

Tobayama T., Uchida S. and Nishiwaki M. 1969. A white-bellied right
whale dolphin caught in the waters off Ibaragi, Japan. The Ocean
Res. Inst., Univ. of Tokyo, Collected Reprints, у. 8, 648 - 657.

Tobayama T., Uchida S. and Nishiwaki M. 1969. A white bellied right
whale dolphin caught in the waters off Ibaragi, Japan. Journ. Mam-
malogical Soc. Japan, v. 4, N. 4, 5, 6. 112-120 (in Japanese).

Townsend C.H. 1935. The distribution of Certain Whales as shown by
logbook records of American Whale Ships. Zoologica, v. 19, N. 1.

Troitzky А. 1953. Contribution a l’etude des Pinnipedes a propos
de deux phoques de la Mediterranée ramenés de croisiere par
5. А. S. Le Prince Rainier Ш de Monaco. Bull. Inst. Oceanogr.
Monaco, N. 1032.

Trouessart E.L. 1898. Catalogues mammalium. Berolini.

True F.W. 1899. Mammals of the Pribilof Islands. The Harbour seal,
Phoca largha Pallas. In: Jordan D.S. The Fur Seals and Fur Seal
Islands of the North Pacific Ocean. Washington.

Uda M. 1954. Study of the relation between the whaling ground and
hydrographic condition. Scient. Rep. Whal. Res. Inst. М. 9, 179 - 187.

Van Utrecht W.L. 1958. Temperaturregulierende Gefassysteme in der
Haut und anderen epidermalen Structuren bei Cetaceen. Zool. Anz.,
Bd. 161, H. 3/4.

Van Utrech-Cock C.H. 1966. Age determination and reproduction
of female Fin Whales, Balaenoptera physalus (Linnaeus, 1758)
with special regard to baleen plates and ovaries. Bijdragen tot de
Dierkunde, 35. Amsterdam.

Venables U.M. and Venables Z.S.V. 1955. Observation on a breeding
colony of the seal Phoca vitulina in Shetland. Proc. Zool. Soc. Lon-
don, v. 125.

701

987

Venables О.М. and Venables Z.S.V. 1957. Mating behaviour of the seal
Phoca vitulina in Shetland. Proc. Zool. Soc. London, v. 128, pt. 3.

Vibe C. 1950. The marine mammals and marine fauna in the Thule
District (North-west Greenland) with observations on ice conditions
in 1939-41. Medd. om Gronland, 150.

Vik R. 1959. Sel, laks, parasitter og fredning. Fauna, 12, N. 2.

Vladykov V.D. 1943. A modification of the pectoral fins in the beluga
from the St. Lawrence River. Natur. Canad., v. 70, N. 1-2.

Vladykov V.D. 1944. Etudes sur les mammiféres aquatiques Ш. Chasse
biologique et valeur economique du morsouin blanc ou beluga
(D. leucas) du bleuve et du Golfe St. Laurent. Dept. Pécheries Proc.
Quebec, Canada, 194 p.

Vladykov V.D. 1947. Etudes sur les mammiferes aquatiques. IV.
Nourriture du marsouin blanc ou beluga (D. leucas) du fleuve
Saint—Laurent. Contrbs. Inst. biol. de Univ. de Montreal, М 15,
Canada, 155 p.

Wagner H. 1936. Geburt und Jugendentwicklung beim Seehund (Phoca
vitulina L.) Zool. Garten, N. 8.

Walker L.W. 1949. Nursery of the Grey Whale. Nat. Hist., v. 58, N. 6.

Walker E.P. 1964. Mammals of the World. Baltimore.

Waters W.E. 1965. Grey seal haulouts at St. Kilda. Proc. Zool. Soc.
London, v. 145, pt. 1.

Weber М. 1886. Studien wber Saugetiere. Ein Beitrag zur Frage nach
dem Ursprung der Cetaceen. Jena.

Wijngaarden A. van 1962. The Meditterranean Monk Seal from report
to the International Union for Conservation of Nature: Oryx., v. 6,
№5.

Wijngaarden А. van 1969. The Monk Seal colony et la Guerra Rio d’Oro.
IUCN Bulletin, New Ser., v. 2, N. 10.

Wilke F. 1954. Seals of Northern Hokkaido. Journ. Mammal., v. 35, N. 2.

Wilke F. 1957. Food of sea otters and harbour seals at Amchitka Island.
Jovrn. Wildlife Management, v. 21, N. 2.

Wilke F. and Kenyon K.W. 1957. The food of Fur Seal in the Eastern
Bering Sea. Journ. Wildlife Management, v. 21, N. 2.

Wilke F. and Fiscus C.H. 1961. Grey Whale observation. Journ. Mam-
mal., у. 42, М. 1, 108-109.

Wilke Е. and Nicholson A.J. 1958. Food of porpoises in waters off Japan.
Journ. Mammal., у. 39, М. 3, 441 - 443.

Wilke F., Taniwaki T. and Kuroda N. 1953. Phocenoides and
Lagenorhynchus in Japan with notes on hunting. Journ. Mammal.,
у. 34, М. 4, 488 - 497.

988

Winge Н. 1902. Groenland Pattedyr. Conspectus faunae groenlandicae
Mammalia. Medd. on Groenland, 21, Kjobenhavn.

Winge H. 1924. Pattedyr—Slaegter. П. Kjobenhavn.

Winge H. 1941. The interrelationships of the Mammalian genera. Roden-
tia, Carnivora, Primates, у. II. Kjobenhavn.

Wolk K. 1963. Foka szara, Halichoerus grypus (Fabr.) v ujsciu Swiny.
Przeglad Zologiczny, VII, N. 4, 342-345.

Wolk K. 1969. Factors affecting seal population levels in the Southern
Baltic Sea. Saugetierkundliche Mitteilungen, H. 2.

Wolleback A. 1907. Ueber die Biologie der Seehunde und die Seehund-
jagd im Europaischen Eismeer. Rapp. et Proc.—Verb. v. 8. Copen-
haven.

Worthington L.V. and Schevill W.E. 1957. Underwater sounds from
Sperm Whales. Norsk Hvalfangst—Tidende, М. 10, 573-757.

Yamada M. 1953. Contribution to the anatomy of the organ of hearing
of whales. Scient. Rep. Whal. Res. Inst. 8, 1979.

Yamada M. 1954. An account of rare porpoises, Feresa Grey from Japan.
Scient. Rep. Whal. Res. Inst., N. 9, 59-88.

Yamada M. 1953. Some remarks on the Pygmy sperm whale, Kogia.
Scient. Rep. Whal. Res. Inst., N. 9, 37-59.

Zukowsky L. 1921. Beitrag zur kenntnis von Pusa hispida pygmaea. Arch.
Naturgesch. Berlin, Abt. A, H. 10.

702

INDEX OF LATIN NAMES ОЕ
ANIMALS’

Acrodelphidae 436, 441
Agrophiidae 435, 436
Alachtherium 27
Allodesmus 54
—kernensis 19
Anarnak 654
—groenlandicus 654
Archaeoceti 430
Arctocephalinae 54, 75
Arctocephalus 16, 18, 19, 21, 22, 54, 75,
79
—australis 19, 54
—californianus 81
—doriferus 54
—forsteri 54
—gazella 54
—monteriensis 56
—philippii 19, 54
—pusillus 18, 19, 54, 79
Argyrocetus 441
Artiodactyla 6, 8
Aulophyseter 593

Balaena ampullata 653, 654

—glacialis 425

—rostrata 654

Balaenoptera musculus 422

physalus 422, 422,** 424, 427

Basilosauridae 430

Beluga 562

Berardius 420, 627, 629

— агпоих 629

—bairdii 433, 437, 552, 629, 630, 630, 631,
632, 635, 656

—vegae 630
—vegana 630

Caenolestia 6

Caenolestoidea 6

Callocephalus 163

Callophoca 118

Callorhinus 14, 16, 18, 54, 74, 75, 79, 80

—alascanus 81

—curilensis 81

—obscura 166

—ursinus 14, 19, 21, 24, 52, 54, 55, 56, 68,
80, 81, 82, 84, 85, 86

- ~alascanus 87

- -californianus 87

--curilensis 86

- -cynocephalus 87

- -kracheninnikovi 86

--mimica 86

- -niger 86

- -ursinus 86

Callotaria 80

Canoidea 21

Carnivora 6, 8, 11, 13, 15, 21, 52

Caspiopusa Behningi 197

—Dierzawini 197

—kisielewitschi 197

Catodon 594

—australis 594

—macrocephalus 594

Cefus albicans 562

Cephalorhynchus 442

Ceratodon 587, 588

Cetacea 3, 7

“Reproduced from the Russian original. Page numbers of the Russian original appear
in the left-hand margin in the text—General Editor.

**Раре numbers in italics indicate illustration

990

Chiroptera 7

Clymene 444

—euphrosyne 444

Clymenia 444

Cystophinae 118

Cystophoca 392

Cystophora 14, 18, 118, 119, 392, 393
—borealis 392, 394

—cristata 19, 118, 119, 122, 122, 126, 393

395, 396, 397

Cystophorinae 118, 119, 129, 377, 390,

391, 392

Dasyuria 6
Delphinapterus 482, 562, 568, 587
—beluga 562, 563
—borealis 482
—dorofeevi 563
—freimani 563

—leucas 433, 438, 563, 564, 567, 568
- -leucas 568

- -dorofeevi 568

- -maris-albi 563, 568
Delphinavus 441
Delphinidae 435, 436, 441
Delphinoidea 436, 441
Delphinapteridae 561
Delphinus 442, 446, 452
—acutus 489

—aduncus 472

—albicans 562
—albirostris 487, 492
—algeriensis 455

—alope 451

—bairdii 455
—bredanensis 444
—butskopf 654
—capensis 455
—coeruleo-albus 446
—curvirostris 455
—deductor 522, 523

—delphis 418, 422, 432, 433, 440, 455, 456,

457, 458, 459
- -bairdii 460
- -delphis 460
- -ponticus 455, 460
—desmaresti 647
—dubius 450
—euphrosyne 444, 446
—frontalis 451

—frontatus 444
—gladiator 507
—globiceps 523
—grinda 523
—grisus 518, 519, 520, 521
—hyperoodon 654
—intermedius 535
—lateralis 446
—leucopleurus 487, 489
—malayanus 450
—leucas 562
—melas 523
—nesarnak 472
—orca 506, 507
—pectoralis 501
—peronii 482, 486
—phocaena 536, 558
—phocaenoides 557, 558
—rissoanus 518
—roseiventris 551
—rostratus 442, 444
—sowerbiensis 638
—sowerbensis 643
—styx 446
—truncatus 472
—tursio 472
Desmatophoca 54
Diaphorocetus 593
Didelphida 6
Diplodon 641
Diprotodontoidea 6
Disigatus 54
Dolichodon 641
Doridontidae 430

Electra 486

—electra 487

—obtusa 501

Eocetus 430

Erignathini 131

Erignathus 14, 16, 118, 119, 127, 129, 130,
131

—barbatus 19, 21, 24, 119, 120, 123, 127,
130, 131, 132, 134, 137, 141

- -albigena 142

- -barbatus 139, 141, 142, 160

- -lachtak 142

- -leporinus 142

- -lepechini 142

- -nauticus 139, 141, 142, 144, 156

- - рагзопзи 142

Eumetopias 18, 54, 56, 74, 75

—gillespii 78

—jubata 54

—jubatus 14, 52, 55, 57, 57, 59, 60, 66, 68,
107

Euphrosyne 444

Euphyseter 626

Eurhinodelphidae 436

Eurinodelphinidae 441

Feresa 442, 501, 534
—attenuata 439, 534, 535, 536
—occuleta 535

—uttenu 535

Gladiator 506

Globicephala 442, 522

—macrorhyncha 523

—melaena 438, 522, 523, 523, 524, 526,
527

- -deductor 525

- -globiceps 525

- -melaena 525

- -melas 525

- -scammoni 525

- -sibo 525

- -sieboldii 525

—scammoni 523

—sieboldii 523

Globicephalidae 441

Globicephalus 522

—chinensis 518

—scammoni 523

—sibo 523

Globiceps 522

Grampidae 441

Grampidelphis 518

Grampus 442, 489, 506, 518

—griseus 438, 507, 518

—orca 507

—stearnsii 518

Gryphoca 118, 341

Halichoerus 18, 118, 119, 127, 129, 340

—atlantica 341

—baltica 341

—griseus 340, 341

—grypus 19, 118, 120, 123, 123, 127, 341,
342, 344, 349, 351, 352, 394

991

—gryphus 354
- -atlantica 354

- -baltica 353

- -gris 354

- -grisseus 354

- - вгуриз 353, 354

- -halichoerus 354

- -macrorhynchus 353

- -pachyrhynchus 353

- -thienemanni 354

—macrorhynchus 341

—pachyrhynchus 341

Halicyon 163

—richardii 163, 232

Haliphilus 163

Heliophoca 375

—atlantica 375

Hemisyntrachelidae 436, 441

Hemisyntrachelus 472

Histriophoca 118, 122, 127, 163, 165, 166
167, 286, 329, 330, 331, 332

—fasciata 19

Histriophocina 286

Hoplocetinae 593

Hydrurga 16, 118, 374

—leptonys 115, 119

Hyperoodon 627, 629, 653

—ampullatus 437, 654, 655, 656

—butskops 653

—planifrons 654

Hyperoodon rostratus 422

Hyperoodontidae 436, 627

Indopacetus 654
Inia 422
—goeffrensis 413
Insectivora 6, 8

Kogia 593, 623, 624, 625
—breviceps 437, 627
—simus 437, 623, 627
Kogiidae 436

Kogiinae 593

Lagenodelphis 442

Lagenorhynchus 442, 487, 488, 501
—acutus 440, 489, 490, 491
—albirostris 439, 489, 493
—cruciger 489

—electra 439, 442, 487, 489, 501, 501

992

—obliquidens 439, 489, 495, 496, 498
—ognevi 495

—thicolea 489

Lagomorpha 6, 8

Lemuroidea 6

Leptonychotes 118, 374

—weddelli 19, 115, 119, 129
Leptophoca 19, 118

Leucopleurus 487

Lissodelphis 442, 482

—borealis 438, 482, 483, 484, 485, 486
—peronii 438, 482, 484, 487
Lobodon 16, 118, 374
—carcinophaga 19, 115, 119
Lobodoninae 118, 374

Lobodontini 374

Macrosselidea 6

Marsupialia 6

Mesoplodon 627, 638, 640, 641

—bidens 437, 639, 641, 643, 645, 646

—bowdoini 641

—carlhubbsi 641

—densirostris 639, 641

—europaeus 639

—ginkgodens 639, 641

—grayi 639

—hectori 641

—layardi 639, 641

—mirus 639, 654

—planifrons 654

—stejnegeri 437, 641, 642, 643, 644

Mesotaria 118

—ambigua 393

Miophoca 18, 19

—vetusta 341

Mirounga 14, 15, 18, 118, 119, 374, 392

—angustirostris 18, 119, 392

—leonina 9, 115, 116, 119, 397, 392

Monachinae 16, 118, 119, 129, 372, 373,
374, 377, 393

Monachini 374

Monachus 18, 118, 119, 131, 372, 374, 375,
376, 377

—mediterraneus 377

—monachus 19, 21, 119, 122, 126, 377,
378, 380, 381, 382

—schauinslandi 18, 21, 119, 377, 382

—tropicalis 18, 19, 21, 377, 382

Monodon 562, 587

—monoceros 438, 587, 588, 588, 589, 590
—narhval 588

Monodontidae 436, 441

Monotherium 19, 118

Mustelidae 118

Mutica 3

Mysticeti 6, 419, 430, 431

Narwalus 587

—vulgaris 587, 588

Neomeris 557

Neephoca 18, 19, 54, 75

—cinerea 54

—hookeri 54

Neophocaena 442, 557

—phocaenoides 438, 558, 558, 559, 560
Neotherium 54

Notocetus 627

Odobenidae 4, 12, 13, 20, 21, 22, 23, 24,
25

Odobenus 16, 27

—rosmarus 24, 25, 26, 27, 31

- -arcticus 33

- -cookii 33

- -divergens 33, 41, 44, 45

- -laptevi 28, 33

- -obesus 33

- -orientalis 33

- -rosmarus 33

Odontoceti 3, 6, 430, 431, 435

Ommatophoca 118, 372, 374

—rossi 119

Orca 506

—ater 507

—fusca 507

—gladiator 506

—intermedia 534

—rectipinna 507

Orcaelidae 441

Orcaella 442

Orcinus 442

—orca 317, 438, 507, 508, 509, 510, 511

—rectipinna 512

Otaria 19, 54, 56, 75

—byronia 19, 54

—californiana 74, 75

—gilliespii 74, 75

—japonica 75

—kracheninnikovi 81

—stelleri 58

Otariidae 4, 11, 12, 13, 14, 15, 16, 19—23,
24, 25, 27, 53, 54, 54, 118

Otarlinae 56, 75

Otarioidea 14, 21, 27, 52

Pagomy 163

Pagophila 163

Pagophilus 118, 163, 165, 167, 279, 283,
286, 287, 290, 302

—groenlandica 122

Pagophoca 118, 163

Paleophoca 118

Pappocetus 430

Pelagias 374

Pelagios 374

Pelagius 374

Pelagus 374

Pelagocyon 374

—monachus 374

Peramelia 6

Perissodactyla 8

Phalangeria 6

Peponocephala 442, 487, 489, 501

Роса’ 22, 118, 119 S127 22 23127,
129, 163, 166, 167, 232, 236, 244,
244, 245, 246, 278, 341

—albigena 131

—albiventer 377

—annelata 168

—baicalensis 220

—barbata 130, 131

—bicolor 377

—canina 232

—caspica 14, 19, 119, 121, 121, 124, 124,
129, 166, 167, 171, 197, 199, 200, 201

—chorisii 232

—communis 168

—crinita 377

—cristata 392, 394

—cucullata 394

—curilensis 246

—dorsata 278

—equestris 328

—fasciata 4, 119, 122, 122, 125, 163, 164,
165, 166, 167, 328, 329 — 332

—foetida 163, 167, 168

—groenlandica 19, 21, 118, 121, 125, 127,
129, 163, 164, 165, 166, 167, 278, 279,
283, 286, 287, 290

993

—ochotensis 232

- -groenlandica 291

- -oceanica 291, 302

—grypus 340, 341

—halitschensis 166

—hermanni 377

—hispida 14, 19, 21, 119, 121, 121, 124,
125, 129, 164, 166, 167, 168, 169, 170,
171, 173, 177, 202, 215, 236, 243

- -annelata 178

- -beaufortiana 179

- -birulai 168, 178

- -botnica 178

- -gischigensis 168, 178

- -hispida 178, 179

- -krascheninnikovi 168, 178, 181, 193

- -ladogensis 178, 195

- -nummularis 178

- -ochotensis 176, 178, 193

- -pomororum 168, 178

- -pygmaea 168, 178

- -rochmistrovi 168, 178

- -saimensis 178

- -Schreb 202, 232

- -soperi 179

—insularis 21, 232

—isidorei 394

—Jjubata 56, 58

—lachtak 131

—ladogensis 168

—largha 119, 126, 232, 237, 244

—leonina 58

—lepechini 131

—leporina 131

—leucogaster 377

—leucopla 394

—linnaei 232

—littorea 232

—mimica 81

—mitrata 394

—monachus 374, 377

—nautica 131

—nigra 81

—nummularis 232

—oceanica 278

—ochotensis 168, 232

- -kurilensis 232

—oronensis 220

—parsonsii 131

—petersi 232

994

—richardi pribilofensis 232

—rosmarus 27, 28

—saimensis 168

—semilunaris 278

—schreberi 168

—scopulicola 232

—sibirica 4, 119, 121, 121, 124, 124, 129,
164, 166, 167, 171, 202, 220,
220—222

—steinegeri 232, 237

—thienemannii 232

—variagata 232

—vitulina 19—22, 24, 119—121, 121, 124,
‘125, 126, 126, 163, 164, 166, 167,
171, 197, 220, 231, 232, 237, 238, 242,
244

- -botnica 167

- -canina 245

- -chorisii 246

- -concolor 246

- -insularis 246

- -kurilensis 119, 125, 237, 246, 247, 248,
250, 253

- -largha 125, 126, 232, 236, 244, 244, 245,
246, 250, 251, 252, 282

- -limnaei 245

- -littorea 245

- -macrodens 244, 246

- -nummularis 246

- -mellonae 249

- -ochotensis 244

- -pallasii 232, 244

- -petersi 244

- -richardi 125, 163, 246, 249

- -scopulicola 245

- -steinegeri 246

- -thienemanni 245

- -variegata 245

--vitulina 119, 123, 125, 126, 233, 235,
236, 244, 245

—vitulinoides 166

—ursina 79—81

Phocanella 118

—minor 166

—pumila 166

Phocarctos 18, 19

Phocidae 4, 12, 12, 13, 13, 14, 15, 18—23,
24, 25, 52, 117, 118—120, 127, 129,
300, 372, 390

Phocinae 16, 22, 114, 118, 119, 127—129,
136, 163, 341, 372, 377, 378, 390, 393

Phocoena 442, 470, 536, 537

—communis 538

—crassidens 501, 502

—dalli 549

—dioptrica 538

—phocoena 439, 538, 539 — 541

- -communis 542

- -phocoena 542

- -relicta 542

- -vomerina 542

—relicta 538

—sinus 538

—spinnipinnis 538

—vomerina 538

Phocoenidae 441

Phocoenoides 442, 548, 552

—dalli 439, 549, 549—551, 555

- -dalli 512, 549, 553, 634

- -truei 512, 549, 553

—truei 548, 549

Phocoidea 21, 27, 52

Physeter 539, 593, 594

—australasianus 594

—bidens 638, 643

—breviceps 623

—catodon 436, 594, 595, 596, 598 — 601,
614, 616

- -australasianus 603

- -australis 603

- -catodon 602

- -macrocephalus 602

—macrocephalus 594

—simus 626

—sulcatus 594

Physeteridae 435, 436, 593

Physeteroidea 436

Pinnipedia 3, 7, 17, 23

Pithanotaria 52

Platanista 434

—gangetica 413

Platanistidae 436

Platanistoidea 436

Platyphoca 118, 131

Pliopedia 54

Polyprotodontoidea 6

Pontolis 54

Pontophoca 118, 374

Pontoporia blainvillei 419

Pristiphoca 118, 375

Procidae 16

Prodelphinus 444

Prophoca 118

Prorosmarus 27

—alleni 27

Protocetidae 430

Protocetus 430

Pseudorca 442, 501

—crassidens 439, 502, 503, 504

—mediterranea 502

Pusa 118, 163, 166—168, 171, 178, 179,
193, 202, 232, 236, 286, 287, 291, 341

Rigoon 375

Rodentia 8

Rosmarus 27

Rostrifer nestorésmirnovi 630

Schizodelphis 441

Semantor macrutus 20
Semantoridae 21

Simae 6

Siren cynocephala 80

Sirenia 8

Sotalia 441, 442, 444

Sousa 444

Squalodon 627

Squalodonttdae 435, 436, 441, 627
Stenella 442, 444, 445, 450
—asthenops 446

—clymene 446

—caeruleoalba 440, 446, 447 —449, 512
- -euphrosyne 450
—coeruleoalbus 446, 450
—crotaphiscus 446

—dubia 440, 446, 450

—frontalis 440, 446, 450, 453
—graffmani 446

—longirostris 440, 446, 451, 453—455
—mialayana 446

—pernettyi 446

Stenidae 441, 444

Steno 445

—attenuatus 444, 450
—bredanensis 439, 442, 442, 443
Stenorhynchinae 118, 372
Stenorhynchus 372

995

Sudidae 554

Tarsioidea 6
Tasmacetus 627, 627
Trichechodon 27
Trichechus 27
—arcticus 28
—cookii 28
—divergens 28
—manatus 27
—obesus 28
—orientalis 28
Tridacna 35

True 19

Tupaioidea 6

Tursio 472, 482
Tursiops 442, 446, 470, 472
—abusalam 472
—aducans 472
—gephyneus 472
—gillii 472
—nuuanu 472
—parvimanus 472
—truncatus 422, 439, 472, 473 —476, 480
- - gillii 477

- -ponticus 472, 477
- -truncatus 477
Tylopoda 6

Ursidae 52
Ursus marinus 81

Vegae 630
Vermes 557

Xiphus 647

Zalambdodonta 6

Zalophus 13, 14, 16, 18, 54, 74, 76
—californianus 4, 19, 54, 55, 75, 76, 77
- -californianus 78

- -japonicus 78

- -wollebaecki 78

—cavirostris 473, 647, 647, 648 —650
Ziphiidae 419, 435, 436, 627, 628, 629
Ziphius 627, 647

—grebnitzkii 648

а
и
№

а,

м,

ny i
i om
Peale

I i 41 oe age es м м
о : Te:

ey у ие

т УИ

Ай
и
И
и
ий
|

Hart

1

—#- и в шв и

роевомьиы | МИНИН

MAMMALS OF THE ‘SOVIET UNION
Volume I
V.G. HEPTNER, A.A. NASIMOVICH, A.G. BANNIKOV

This volume of Mammals of the Soviet Union is devoted to a description
of artiodactyls and perissodactyls found in the Soviet Union. These
animals are of great scientific and economic interest; information on them,
especially in recent Soviet literature, is voluminous. In recent years world
literature on this subject has likewise been considerably enriched with new
data. The abundance of information on ungulates explains the size of the
present volume. All the characteristics of groups have been described
according to a common scheme; deviations occur in a few cases, however.
These characteristics are stated briefly and pertain to the group as a
whole; they are not exclusive to species of the Soviet fauna. All species
are described according to a common plan, altered only in the case of
some extinct forms. In devising the scheme for descriptions of species not
only the convenience of the reader was kept in mind, but the hope that
gaps in our knowledge would become self-evident and stimulate further
research.

MAMMALS OF THE SOVIET UNION
Volume II, Part 2

This volume, is part of a three-volume monograph, and is a continuation,
of Volume II, Part 1, which was devoted to sea cows and carnivores. It
contains species descriptions of terrestrial carnivores and detailed informa-
tion on their external morphology, skull, body measurements and other
data, affinities with other species, geographic distribution in the historic
past and today, geographic variation, practical significance, and biology.
Descriptions are presented for orders, families, and genera, and keys given
for their identification.

The book is richly illustrated with photographs, sketches, and colored
illustrations by the famous wild life painter, А.М. Komarov, and the
zoologist-artist, N.N. Kondakov.

The results of original scientific studies are published here for the .
first time, providing readers a vast wealth of material heretofore unknown.
This work is intended for teachers and students of faculties of biology and
geography in universities, as well as pedagogic, agricultural, and forest
institutes, similar organizations, workers engaged in game, fur, forest and
fish trades; those interested in the conservation of nature, and all persons
interested in zoology and nature study.