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Full text of "Mammals of the Soviet Union"

Mammals 

of the. 

Soviet Union 



V. G. Heptner 



N.P. Naumov 







1Ш^'.Ш1 



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, i.e., 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, 
i.e., 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 moфhological 
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. 




f<--^<^^ 



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 II, Part 3 

PINNIPEDS AND TOOTHED WHALES 

Pinnipedia and Odontoceti 



Late V.G. Heptner, K.K. Chapskii, V.A. Arsen'ev 
and V.E. 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.: SI 1.2:Ar7/2 

Contents: v. 1. Artiodactyla and Perissodactyla — 
V. 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. II. Bannikov, Andrei Grigor'evich. 
III. Hoffmann, Robert S. IV. SludskK A. A. V. 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. Юишоу 
(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 
B.I. Badamshin (Fort Shevchenko), F.Sh. Khuzin, M.Ya. Yakovenko 
(Murmansk), AG. Beloborodov, Yu.I. Nazarenko, and V.A. Potelov 
(Arkhangel'sk), A.A. Berzin, G.M. Kosygin, Yu.V. Kurochkin, and late 
E.A. Tikhomirov (Vladivostok), A.P. Shustov and G.A 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 LP. 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. I.I. 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), 



VI 

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 




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 entre 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. 



Vlll 

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 v Yaroslavskoi i 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-tobthed) 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 M (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 a "G" sound in Russian. However, 
the surname was originally German, and in the original German began 
with the letter "H" of the Latin alphabet. Since Cyrillic has no equivalent 
of "H" this is usually transliterated into "G" 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 vqlume, 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 



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, i.e., 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 



Xll 

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 pfesent 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 arid 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 



Xlll 

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 califomianus). 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 vii 

FOREWORD xi 

CLASSIFICATION OF CLASS MAMMALIA xxvii 

KEY FOR IDENTIFYING ORDERS OF MAMMALS xxix 

PART I. ORDER OF PINNIPEDS 
ORDER PINNIPEDIA ILLIGER, 1811 

Cohort Ferungulata Simpson, 1945 3 

Superorder Ferae Linnaeus, 1758 3 

Order Pinnipedia Illiger, 1811 3 

Family Odobenidae Allen, 1880 (Walruses) 22 

Genus Odobenus Brisson, 1762: Walruses 26 

Odobenus rosmarus (Linnaeus, 1758): Walrus . 26 

Diagnosis 27 

Description 27 

Geographic Distribution 29 

Geographic Variation 32 

Biology 34 

Economic Importance 54 

Superfamily Otarioidea Smirnov, 1908 58 

Family Otariidae Gill, 1866 (Eared Seals) 58 

Subfamily Otariinae Boetticher, 1934 (Sea Lions) 64 

Genus Eumetopias Gill, 1866: Steller's Sea Lion 64 

Eumetopias jubatus (Schreber, 1776): Steller's 68 

Sea Lion 

Diagnosis 68 

Description 68 

Taxonomy 69 



* Pages 713-718 in the Russian original. The EngUsh table of contents is not a Uteral 
translation. 



XVI 



Geographic Distribution 69 

Geographic Variation 72 

Biology 72 

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 



XVll 



Phoca (Pusa) hispida Schreber, 1775: 


218 


Ringed Seal 




Diagnosis 


219 


Description 


219 


Taxonomy 


223 


Geographic Distribution 


223 


Geographic Variation 


231 


Biology 


234 


Economic Importance 


256 


Phoca (Pusa) caspica Gmelin, 1788: 


260 


Caspian Seal 




Diagnosis 


260 


Description 


261 


Taxonomy 


266 


Geographic Distribution 


267 


Geographic Variation 


269 


Biology 


269 


Economic Importance 


286 


Phoca (Pusa) sibirica Gmelin, 1788: Baikal 


290 


Seal or Baikal Ringed Seal 




Diagnosis 


290 


Description 


290 


Taxonomy 


295 


Geographic Distribution 


295 


Geographic Variation 


295 


Biology 


295 


Economic Importance 


304 


Subgenus Phoca Linnaeus, 1758: True Seals 


307 


Phoca (Phoca) vituhna Linnaeus, 1758: 


307 


Common Seal, Larga 




Diagnosis 


308 


Description 


308 


Taxonomy 


313 


Geographic Distribution 


315 


Geographic Variation 


323 


Biology 


330 


Economic Importance 


364 


Subgenus Pagophilus Gray, 1844: Harp or 


369 


Greenland Seals 




Phoca (Pagophilus) groenlandica 


369 


Erxleben, 1777: Harp or Greenland Seal 




Diagnosis 


370 



XVIU 



Description 


371 


Taxonomy 


381 


Geographic Distribution 


382 


Geographic Variation 


387 


Biology 


389 


Economic Importance 


429 


Subgenus Histriophoca: Ribbon Seals 


436 


Phoca (Histriophoca) fasciata 


436 


Zimmermann, 1783: Ribbon (Banded) Seal 




Diagnosis 


437 


Description 


437 


Taxonomy 


440 


Geographic Distribution 


440 


Geographic Variation 


442 


Biology 


442 


Economic Importance 


451 


Genus Halichoerus Nilsson, 1820: 


452 


Gray Seals 




Halichoerus grypus (Fabricius, 1791): Gray Seal 


454 


Diagnosis 


455 


Description 


455 


Taxonomy 


466 


Geographic Distribution 


467 


Geographic Variation 


469 


Biology 


472 


Economic Importance 


493 


Subfamily Monachinae Trouessart, 1904 




(Monk Seals or 8-incisored Seals) 


495 


Genus Monachus Flemming, 1822: Monk Seals 


499 


Monachus monachus (Hermann, 1779): Monk 


502 


Seal 




Diagnosis 


502 


Description 


502 


Taxonomy 


507 


Geographic Distribution 


508 


Geographic Variation 


512 


Biology 


512 


Economic Importance 


519 


Subfamily Cystophorinae Gray, 1866 (Hooded 


519 



XIX 



Seals and Elephant Seals, or 6-incisored 




Seals) 




Genus Cystophora Nilsson, 1820: Hooded Seals 


521 


Crystophora cristata Erxleben, 1777: Hooded 


524 


Seal 




Diagnosis 


525 


Description 


525 


Taxonomy 


529 


Geographic Distribution 


529 


Geographic Variation 


532 


Biology 


532 


Economic Importance 


543 



PART II. ORDER OF WHALES 
ORDER CETACEA BRISSON, 1762 

Cohort Mutica 551 

Order Cetacea Brisson, 1762 551 

Suborder Odontoceti Flower, 1867 575 

Superfamily Delphinoidea Flower, 1864 587 

Family Delphinidae Gray, 1821 (Dolphins) 589 

Genus Steno Gray, 1846: Rough-toothed 590 

Dolphins 

Steno bredanensis Lesson, 1828: Rough- 591 
toothed Dolphin 

Genus Stenella Gray, 1866: Spotted Dolphins 592 

Stenella coeruleoalba Meyen, 1833: 594 
Blue-white Striped Dolphin 

Diagnosis 594 

Description 596 

Geographic Distribution 598 

Geographic Variation 600 

Biology 600 

Stenella dubia [=attenuata\ G. Cuvier, 1812: 601 
Malay [Pan-tropical Spotted] Dolphin 

Stenella frontalis G. Cuvier, 1829: Bridled 602 
[Atlantic Spotted] Dolphin 

Stenella longirostris Gray, 1828: Long-snout 604 
[Spinner] Dolphin 

Genus Delphinus Linnaeus, 1758: Common 604 
Dolphins 



XX 



Delphinus delphis Linnaeus, 1758: Common 


607 


Dolphin 




Diagnosis 


607 


Description 


607 


Geographic Distribution 


611 


Geographic Variation 


611 


Biology 


614 


Economic Importance 


627 


Genus Tursiops Gervais, 1855: Bottlenose 


631 


uuipnins» 

Tursiops tmncatus Montagu, 1821: 


632 


Bottlenose Dolphin 




Diagnosis 


633 


Description 


633 


Geographic Distribution 


634 


Geographic Variation 


638 


Biology 


639 


Economic Importance 


644 


Genus Lissodelphis Gloger, 1841: Right Whale 


645 


Dolphins 




Lissodelphis borealis (Peale, 1848): Northern 


646 


Right Whale Dolphin 




Diagnosis 


646 


Description 


647 


Geographic Distribution 


647 


Geographic Variation 


648 


Biology 


648 


Lissodelphis peroni (Lacepede, 1804): 




Southern Right Whale Dolphin (Peron's 


651 


Dolphin) 




Geographic Distribution 


652 


Biology 


653 


Genus Lagenorhynchus Gray, 1846: Shorthead 


653 


Dolphins 




Lagenorhynchus (Lagenorhynchus) acutus 


656 


Gray, 1828: Atlantic White-sided Dolphin 




Diagnosis 


656 


Description 


656 


Geographic Distribution 


657 


Geographic Variation 


659 


Biology 


659 



XXI 



Lagenorhynchus (Lagenorhynchus) albirostris 660 
Gray, 1846: White-beaked Dolphin 

Diagnosis "°" 

Description ""^ 

Geographic Distribution 661 

Geographic Variation 663 

Biology 663 

Lagenorhynchus (Lagenorhynchus) 664 
obliquidens Gill, 1865: Pacific White-sided 
Dolphin 

Diagnosis 665 

Description 665 

Geographic Distribution 666 

Geographic Variation 667 

Biology 667 

Economic Importance 671 

Lagenorhynchus (Peponocephala) electra 672 
Gray, 1846: Broadsnout Dolphin 

Genus Pseudorca Reinhardt, 1862: 673 
False Killer Whales 

Pseudorca crassidens Owen, 674 
1846: False Killer Whale 

Diagnosis 674 

Description 674 
Geographic Distribution 675 

Geographic Variation 678 

Biology 678 

Economic importance 679 

Genus Orcinus Fitzinger, 1860: Killer Whales 679 

Orcinus orca Linnaeus, 1758 : Killer Whale 680 

Diagnosis 681 

Description 681 

Geographic Distribution 683 

Geographic Variation 687 

Biology 687 

Economic Importance 694 

Genus Grampus Gray, 1828: Risso's Dolphins 695 

Grampus griseus G. Cuvier, 1812: Risso's 696 

Dolphin 
Diagnosis 696 

Description 696 

Geographic Distribution 697 



XXll 



Geographic Variation 


698 


Biology 


699 


Genus Globicephala Lesson, 1828: Pilot Whales 


701 


Globicephala melaena Traill, 1809: Pilot 


702 


Whale 




Diagnosis 


702 


Description 


702 


Geographic Distribution 


704 


Geographic Variation 


706 


Biology 


708 


Economic Importance 


717 


Genus Feresa Gray, 1870: Pygmy Killer Whales 


718 


Feresa attenuata Gray, 1875: Pygmy Killer 


719 


Whale 




Genus Phocoena G. Cuvier, 1817: Common 


721 


Porpoises 




Phocoena phocoena Linnaeus, 1758: 


722 


Common Porpoise 




Diagnosis 


722 


Description 


722 


Geographic Distribution 


725 


Geographic Variation 


726 


Biology 


729 


Economic Importance 


735 


Genus Phocoenoides Andrews, 1911: Dall 


737 


Porpoises 




Phocoenoides dalli True, 1885: Dall Propoise 


738 


Diagnosis 


738 


Description 


738 


Geographic Distribution 


740 


Geographic Variation 


741 


Biology 


744 


Economic Importance 


748 


Genus Neophocaena Palmer, 1899: Black 


750 


Finless Porpoises 




Neophocaena phocaenoides G. Cuvier, 1829: 


750 


Black Finless Porpoise 




Diagnosis 


750 


Description 


750 


Geographic Distribution 


751 


Geographic Variation 


752 


Biology 


752 



XXlll 



Family Monodontidae Gray, 1821 (Narwhals) 


755 


Genus Delphinaptenis Lacepede, 1804: Belugas 


755 


or White Whales 




Delphinaptenis leucas Pallas, 1776: Beluga or 


757 


White Whale 




Diagnosis 


757 


Description 


757 


Geographic Distribution 


761 


Geographic Variation 


763 


Biology 


765 


Economic Importance 


785 


Genus Monodon Linnaeus, 1758: Narwhals 


790 


(Narwhals or Unicorns) 




Monodon monoceros Linnaeus, 1758: 


791 


Norwhal or Unicorn 




Diagnosis 


791 


Description 


792 


Geographic Distribution 


792 


Geographic Variation 


794 


Biology 


794 


Economic Importance 


798 


Superfamily Physeteroidea Gill, 1872 


799 


Family Physeteridae Gray, 1821 (Sperm Whales) 


799 


Subfamily Physeterinae Flower, 1864 (Sperm 


800 


Whales) 




Genus Physeter Linnaeus, 1758: Sperm Whales 


800 


Physeter catodon Linnaeus, 1758: Sperm 


801 


Whale 




Diagnosis 


801 


Description 


802 


Geographic Distribution 


808 


Geographic Variation 


812 


Biology 


813 


Economic Importance 


834 


Subfamily Kogiinae Gill, 1871 (Dwarf Sperm 


840 


Whales) 




Genus Ko^a Gray, 1846: Dwarf Sperm 


840 


Whales 




Ko^a breviceps Blainville, 1838: Dwarf 


841 


Sperm Whale 




Diagnosis 


841 


Description 


843 



XXIV 



Geographic Distribution and Biology 


843 


Kogia simus Owen, 1866: Owen's Dwarf 


845 


Sperm Whale 




Diagnosis 


845 


Description 


845 


mily Ziphiidae Gray, 1865 (Beaked Whales) 


846 


Genus Berardius Duvernoy, 1851: Pacific 


849 


Beaked Whales 




Berardius bairdi Stejneger, 1883: Baird's 


850 


Beaked Whale 




Diagnosis 


850 


Description 


850 


Geographic Distribution 


854 


Geographic Variation 


854 


Biology 


855 


Economic Importance 


861 


Genus Mesoplodon Gervais, 1850: Beaked 


862 


Whales [Sword-tooth Dolphins] 




Mesoplodon (Mesoplodon) stejnegeri True, 


866 


1885: Stejneger's Beaked Whale 




Diagnosis 


866 


Description 


866 


Geographic Distribution and Biology 


868 


Mesoplodon (Mesoplodon) bidens Sowerby, 


869 


1804: Sowerby's Beaked Whale 




Diagnosis 


869 


Description 


869 


Geographic Distribution 


872 


Geographic Variation 


873 


Biology 


873 


Genus Ziphius G. Cuvier, 1823: Goose-beak 


875 


[Cuvier's Beaked] Whales 




Ziphius cavirostris G. Cuvier, 1823: Cuvier's 


876 


Beaked Whale 




Diagnosis 


876 


Description 


876 


Geographic Distribution 


878 


Geographic Variation 


882 


Biology 


882 


Economic Importance 


883 


Genus Hyperoodon Lacepede, 1804: Bottlenose 


884 


Whales 





XXV 

Hyperoodon ampullatus Forster, 1770: 885 

Northern. Boltlenose Whale 

Diagnosis 885 

Description 885 

Geographic Distribution 887 

Geographic Variation 888 

Biology 889 

Economic Importance 892 

LITERATURE CITED 895 

OTHER SOURCES 957 

INDEX OF LATIN NAMES OF ANIMALS 989 



CLASSIFICATION OF CLASS 
MAMMALIA 



In this publication the old and widely used system of the major units of 
the class, i.e., 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 



xxvm 



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, 
i.e., 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 

Subclass THERIA 

Infraclass METATHERIA 
Infraclass EUTHERIA 



Cohort UNGUICULATA 

Cohort GLIRES 
Cohort MUTICA 



Cohort 
FERUNGULATA 



Order MONOTREMATA 


Order MARSUPIALIA 


"•Order INSECTIVORA 


Order DERMOPTERA 


* Order CHIROPTERA 


Order PRIMATES 


Order EDENTATA 


Order PHOLIDOTA 


" * Order LAGOMORPHA 


* Order RODENTIA 



Superorder _ 

FERAE 
Superorder 
PROTUNGULATA 

Superorder 
PAENUNGULATA" 

Superorder 
MESAXONIA 
Superorder 
PARAXONIA 



'Order CETACEA 

'Order CARVINORA 
'Order PINNIPEDIA 

Order TUBULIDENTATA 

Order PROBOSCIDEA 
Order HYRACOIDEA 
'Order SIRENIA 

"Order PERISSODACTYLA 
'Order ARTIODACTYLA 



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



1( 2). Hind limbs absent. Body fishlike with large bilobate caudal fluke 
set horizontally CETACEA. 

2 ( 1). Hind limbs present. Body not fishlike; tail, if present, not in the 

form of a bilobate fluke. 

3 ( 4). Forelimbs in the form of leathery wings CHIROPTEIIA. 

4 ( 3). Forelimbs of different structure. 

5 ( 6). Fore- and hind limbs very short, paddle-shaped, and in the form 

of fins, i.e., all digits right up to very tips enclosed in a common 
skin PINNIPEDIA. 

6 ( 5). Fore- and hind limbs of different structure, not in the form of 

fins. 

7 (10). Hooves on legs. 

8 ( 9). Only one hoof on each limb PERISSODACTYLA.^ 

9(8). Two large hooves and two small ones set above them occur on 

each limb ARTIODACTYLA.^ 

10 ( 7). Hooves absent on legs (claws present). 

11 (14). 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. 

12 (13). Two incisors on upper jaw RODENTIA. 

13 (12). Four incisors on upper jaw; small blunt one occurs behind each 

of two large sharp ones LAGOMORPHA. 

14 (11). Diastema between incisors and molars absent or much smaller 

than length of molar row. Canines present. 

15 (16). 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 hmbs 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.) 



'' Skulls of the extinct Steller's cow, a 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 



COHORT OF CARNIVORES AND 
HOOFED MAMMALS 

COHORT Ferungulata Simpson, 1945 

Superorder of Carnivores 
Superorder FERAE Linnaeus, 1758 

ORDER OF PINNIPEDS 
Order PINNIPEDIA lUiger, 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 

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





и {5 

^41 


^''^^iSesS 






■*«:___/ 




8 


Г? 


^ 


^ , 
















12 Fig. 1. Body shape and dimensions of Pinnipedia (seals): A — lateral view; 

В — dorsal view (K.K. Chapskii). 1 — body length from anterior end of snout 
(nostrils) to tip of tail in a straight line (Lev); 2 — same, length along the dorsal 
surface (Lc); 3 — total body length along the dorsal surface (with hind flippers, 
Lp); 4 — axillary girth; 5 — 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. 






13 Fig. 2. Shape of fore flippers and disposition and growth of claws on them 

in the various families of Pinnipedia. A — eared seals (Otariidae); В — walruses 
(Odobenidae); С — 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 



shortened but quite independent and not fused. The humerus is even 
more shortened, but bears a highly developed deltoid crest; the for. 

12 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 

13 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: 

T 3 ^ 1 „ 4 „ 1 ,, 

The number of teeth is less in some seals and walruses. The more 
common permanent teeth formula among walruses is: 

113 

^ С ^ P ^ = 18. 

13 



I 




14 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 
14 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. 



8 

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 ' 
15 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. 



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, i.e., 
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 



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 
16 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 



^ Ice-loving, according to the terminology of N.A. Smiraov (1914, 1927, and 1936). 
^ Land-loving, ice-fearing species. 



11 

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, 
Hydrurga, 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 
18 of cold and moderately cold waters in which the surface temperature does 
not exceed 20° С 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° N lat. to 



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

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



12 



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13 

40° S 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 schauinslandL Pinnipeds are totally absent in 
the entire tropical portion of the Atlantic Ocean from 20° N 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 inhabit 
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 Z^mlya. 

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° С 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 
19 (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, Halichoems 
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 califomianus; 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 antarctic 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 сд5/?/сй)— 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 



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 sea.\,Allodesmus kemensis 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 
20 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 



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 et 
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 
21 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 



^ The origin of a group or two highly proximate famiUes 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 



^ 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. 



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 
22 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, i.e., 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,^ 
i.e., about 39% of all the species known in the world fauna. 



^ 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). 

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



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- 
23 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, 



20 

for soap-making, etc., for making medicinal ("fish") oil, 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.) 

Key to Families of Pinnipedia 
Identification Based on External Features 

1 (4). 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. 

2 (3). 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 
closed mouth Eared seals, Otariidae (p. 58) 

3 (2). 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 

Walruses, Odobenidae (p. 22) 

4 (1). 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. 
True seals, Phocidae (p. 142) 

Identification Based on Skull Features 

1 (4). 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- 
24 sive protuberance directed downward. All or almost all cheek 

teeth with single root (the last with two roots) and with simple 
undivided crown. 

2 (3). 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 



21 



25 



into incisors, canines, and cheek teeth (premolars and molars 
similar). Upper canines similar to lower ones in size and structure. 

Eared seals, Otariidae (p. 58) 

3 (2). 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 

from cheek 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 

True seals, Phocidae (p. 142). (K. Ch.) 



4(1). 




24 



Fig. 6. Transverse notch in middle upper incisor of eared seals, Otariidae, and 
the fur seal, Callorhinus ursinus (figure by K.K. Chapskii). 






24 Fig. 7. Structure of nasals in pinnipeds of various families: A — eared seals (family 

Otariidae); В — walruses (family Odobenidae); С — 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, 







В 

24 Fig. 8. Left zygomatic bone in pinnipeds: A — eared seal (fur seal, Callorhinus 

ursinus); В — walrus {Odobenus rosmanis); С — bearded seal (Erignathus 
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, 
26 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. 




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



24 




Fig. 11. Skull of an adult walrus, Odobenus rosmarus (figure by N.N. 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 



I 



0-0 



or 



0-0' 



1-1' 



M 



3-3' 



27 



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 



25 





26 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, I960*). 

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, I960*). 

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 Odobenus Brisson, 1762 

1762. Odobenus. Brisson. Regnum animale. Ed. 2, p. 30. Phoca rosmarus 
Linnaeus. 

1766. Trichechus. Linnaeus. Syst. Nat., ed. XII, I, p. 49. Nee Linnaeus 
1758 (pertains to manatee Trichechus manatus Linnaeus, 1758). 

1772. Rosmarus. Briinnich. Zoologiae fundamenta, p. 34. Phoca ros- 
marus Linnaeus, 1758. (V.H.) 
See description of the family. 

28 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.) 



27 

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

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

1831. Trichechus cookiL Fremery. Bijdrag. Nat. Vetensk, 6, p. 385. Ici 
Cape zone in Alaska (Chukchi Sea, 70° N lat. and 163° 18' W 
long.). 

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

1940. Odobenus rosmarus laptevL 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. 
29 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). 



28 




28 



Fig. 13. A group of walruses of different ages (figure by N.N. Kondakov). 




29 



Fig. 14. Front view of a walrus head (figure by N.N. 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 g, kidneys 3,544 g, and pancreas 2,148 g. (V.A) 

30 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° N lat.) and in the 1880s were caught in thousands every 
year. Walruses were reported on Cape Kronotskii (56° N 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 



30 




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). 



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. 
33 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 (x = 
369), in females 303-342 mm (X = 314); the maximum width of skull in 
males is 268-291 mm, in females 222-257 mm (x = 234.2). The length 
of male tusks along the curvature from the edge of the alveolus to the 



33 




34 

tip is 34-38 cm, in females 27-33 cm (Ognev, 1935). The tusk length 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 (O. r. divergens (Illiger, 1815)) (syn. cookii, ? orientalis). 

Largest form of the species. 

The maximum body length of males is 450 cm (x = 336), of females 
367 cm {x = 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 (x = 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). 
34 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 

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 moUusks and partly of crustaceans. 
35 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. 








.^fKasagSMiS-^rgg^fi; 



34 Fig. 17. A group of walruses on 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. Kiylov). 

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 
moUusks, 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. 



37 



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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). 

37 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). 

38 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. 




37 Fig. 19. A male walrus with a broken tusk. Chukchi Peninsula (photograph by 

V.I. Kiylov). 



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 m of 
freshly killed animals. They did not react even to gunshots, the clatter 
39 of operating winches on a ship, nor to any other noise. Yet, in general, 
their hearing is better than their sight. 




37 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 walrases. Chukchi Peninsula (photograph by 

P.G. Nikulin). 



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. 
40 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 









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



43 




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 



^ 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). 



44 




41 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. 
41 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 2^mlya 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 



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 
42 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, ft^om the eastern part of the East Siberian Sea to Point 
Barrow in Alaska, only in August. 

In mid-October, when intense forination 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.O. 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- 
ruses 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; Юeinenberg et ai, 
1964). 



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° N lat.), then traveled still northward. 
In mid-April, it was sighted on Helligwer Islands (67° N 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 
43 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). 



47 




43 Fig. 25. The carcass of a female walrus found on the coast of Mednyi Island. A 

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). 



48 




44 Fig. 26. Embiyo of the Pacific walrus, Odobenus rosmarus divergens. Bering Sea 

(photograph by V.I. Krylov). 



44 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 (N. 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 ЮevezaГ, 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 



50 




45 Fig. 27. Annual layers on a polished section of a 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 
moUusks 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. Kiylov) 

Age Male Female 



Newborn 

1-4 months 
Year-old 

2 years 

3 years 

4 years 

5 years 

6 years 

7 years 

8 years 

9 years 

10 years 

11 years 

12 years 

13 years 

14 years 

15 years 

16 years 

17 years 

18 years 

19 years 

20 years 

21 years 

22 years 

23 years 

24 years 

25 years 

26 years 

27 years 

28 years 

29 years 

30 years 

31 - 32 years 

33 years 

34 - 38 years 



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, 



No. of 


Mean 


No. of 


Mean 


animals 


length 


animals 


length 


4 


138 


8 


129 


3 


158 


5 


146 


5 


198 


11 


185 


4 


238 


5 


218 


6 


247 


3 


238 


7 


260 


7 


258 


11 


281 


11 


268 


13 


292 


6 


276 


24 


297 


7 


276 


25 


306 


6 


278 


30 


306 


11 


286 


49 


322 


13 


290 


32 


337 


8 


307 


45 


338 


18 


308 


24 


33& 


8 


298 


30 


352 


17 


307 


35 


354 


25 


299 


43 


351 


18 


308 


34 


358 


22 


304 


35 


358 


15 


312 


31 


362 


8 


315 


29 


370 


10 


311 


19 


371 


8 


312 


12 


370 


4 


328 


17 


375 


7 


326 


14 


371 


8 


321 


14 


369 


4 


321 


13 


372 


4 


321 


4 


367 






12 


370 






7 


371 






12 


363 






6 


370 






5 


372 






7 


374 







52 

they molt every year. The period and duration of molt 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 land 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 stmmosum Rudolphi has been found in seven 
species of pinnipeds and two species of cetaceans; Corynosoma semerme 



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 bearded seal, which also feeds on benthic invertebrates, 
competes with the walrus for food to some extent, as moUusks 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. 

48 The inhabitants of the Chukchi Peninsula and Alaska are permitted to 

hunt walruses, however, to meet their personal requirements. 



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- 
49 pie and sledge dogs, and material for building canoes and even houses 
("yarangas") 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, 



55 





■-1 :'■''.■-' '-~'\ijtjh<::* 



48 Fig. 28. A coastal rookery of walruses in the Gulf of Anadyr (photograph by 

A.V. Yablokov). 



piercing the body of the prioriy 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 (P.C. 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 




■'^.■ 



A; 



L?#^ 



, ■ %**f f <rr ^ 





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 



57 



50 



Table 3. Walrus hunting in the Chukchi Peninsula (Kiylov, 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 


1,273 


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 



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 
51 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.) 

SUPERTAMILY 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 
52 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. 



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: 



•I 



-h 



^i 



M 



2- 1 



= 36 - 34. 



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 




A В 

52 Fig. 31. Nasal portion of skull. A — S teller's sea lion, Eumetopias jubatus; 

В — northern fur seal, Callorhinus ursinus (figure by N.N. Kondakov). 1 — nasal 
bone; 2 — premaxillary bone. 



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. 
54 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 lion, Otaria, with one species, O. byronia [= O. 
flavescence]; the genus of Steller's sea lion, Eumetopias, with one species, 
E. jubatus; the genus of Californian sea lion, Zalophus, with one species, 
Z. califomianus (with three subspecies); the genus of Tasmanian sea 
lion, Neophoca, with two species, N. cinerea (Australia) and N. hookeri 
[= Phocarctos hookeri] (New Zealand); the genus of southern fur 
seals, Arctocephalus, with six species: A. pusillus (South Africa), forsteri 



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, С 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 {Otariidae) [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 {Lcvf^ 135-200 cm, of females 110-150 cm. Color 

^^ Not along the dorsal curvature 



63 

of adult males dark brown, young males and females silvery-gray, 

adult females darker with gray streaks 

Northern fur seal, Callorhinus ursinus (p. 98) 

2 (1). 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 
(II -IV). Hair coat on trunk either lacking underfur or latter 
scanty. 

3 (4). 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 

Steller's sea lion, Eumetopias jubatus (p. 68) 

4 (3). 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 

Californian sea lion, Zalophus califomianus (p. 92) 

Identification Based on Skull Features 

1 (2). 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. 
Steller's sea lion, Eumetopias jubatus (p. 68) 

2 (1). Distance between last upper premolar and first molar (IV 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. 



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 

up to 250 mm 

Californian sea lion Zalophus californianus (p. 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 
56 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 

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

Subfamily of Sea Lions 
Subfamily ОТАШШАЕ 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 lion (figure by N.N. 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. 




iSijiU^'*'' 



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



66 



The genus Eumetopias occupies within the subfamily of sea lions 
(Otariinae) a position at the commencement of a series of specializations. 
This is a genus neighboring Otaria (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, 
57 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). 




57 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, Eumetopias jubatus [dorsal view] 

(figure by N.N. Kondakov). 




57 Fig. 37. Skull of young Steller's sea lion, Eumetopias 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.) 



68 

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

1766. Phocajubata. 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. Diet, class. H.N. 13, p. 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 in old 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; 
A.S. 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. 

Sleller'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). 



69 

females 308-320 mm (x = 314.7); zygomatic width in males 211-261 mm 
(x = 239.8), in females 176-185 mm (x = 180.9); maximum width of 
the skull in males 198-238 mm (x = 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.) 

59 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 lony 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 



— ~ 


140 150 160 


17C 


180 170 






•I V.^ \ \ 




4^=,^===^--^^^^ 






'^ V—^ ^^ ^^-"^ * '^ 


^ 


^^^V ^У'^^-^^"^^^^\^ 




70 




\ 
i 


''f¥\' 


60 


60 




> 

\ 


W^\ 


50 




'Л^^ГЖ^ 


\ 


л7\ \ 






1%^ — y'y si 1 




/ /y \ \ 




50 






W\^ 


40 


40 


jf'i/^ ^ 1 V \ ) 




250 250 500 750 1000 km 

hrn-rb ^ 1 \ 1 






120 130 140 


1 1 1 

150 160 170 





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° N 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 (Attn, Kiska, Amchitka, etc.), on the islands of the 
Gulf of Alaska, on the Pribilov Islands and on nearby Vancouver Island, 



72 



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 











62 Fig. 40. Rookey of Steller's sea lion. Mednyi Island, July, 1969 (photograph by 

S.V. Marakov). 



73 




:. -:!ыП. -. ■ 



lit^'-- ;-=:=^. 





63 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 lony 
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. On 
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 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- 
64 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 



76 

developed a conditioned reflex to the working of fishing trawlers. On 
hearing the noise of a trawl being hauled, the aiiimals in the vicinity 
quickly gather around the boat, dive into the triiwl 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 ifeed 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° N 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. 
65 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). " 



^^ A 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 has 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- 
66 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 Hon, Eumetopias jubatus on 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 



79 




67 



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. 



80 




68 Fig. 46. Rookery of bachelor Steller's sea lion, Eumetopias 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 
67 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° N lat.). The 



81 




K#SfeS:S^^^ 



68 Fig. 47. Steller's sea lions in water. Mednyi Island (photograph by F.G. 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 heiice 
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; 



82 

69 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 lony 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 lony 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 
70 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 


— 


— 


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 years 


5 


213 


198-233 


5 years 


5 


223 


196-244 


6 years 


6 


227 


220-232 


7 years 


7 


230 


210-242 


8 years 


5 


235 


228-245 


9 years 


7 


232 


206-254 


10 years 


7 


235 


221-246 


11 years 


8 


232 


216-254 


12 years 


6 


240 


226-254 


13 years 


7 


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 



84 

The newborns are covered with dark brown soft hair. After molting 
of the juvenile hair, the ftir 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 ftir. 

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, hice, Proechinophthirus 
fluctus (Ferris) гпй 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 
71 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 arid 
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 



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 
73 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 




72 Fig. 49. Steller's sea lions in a гсюкегу of fur seals. Mednyi Island (photograph 

by S.V. Marakov). 



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 (see 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 receiit decades, 
essentially as a result of human intervention. Mention should be made 
first of one of the largest rookeries on lony 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 
74 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 cliffe (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 



88 





72 Fig. 50. Steller's sea lion pup. Kuril Islands, July, 1962 (photograph by G.M. 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.) 



89 




73 Fig. 51. Steller's sea lion diving from a cliff. Mednyi Island (photograph by 

F.G. Chelnokov). 

Genus of Northern [or California] Sea Lions^^ 

Genus Zalophus Gill, 1866 

1866. Zalophus. Gill. Proc. Essex. Inst., Salem, Communications, 5, p. 6. 
Otaria gilliespii MacBain = Otaria califomiana 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 V) on the hind 

flippers are somewhat longer and broader than the middle ones (II-IV). 

75 The upper surface of the hand is naked only terminally; the basal half of 



^^ In view of the unusual presence of these sea lions among the Soviet fauna (see below), a 
brief description of the genus as well as the species is given, which suffices for identification. 
The moфhology is mainly taken from V.B. Scheffer (1958), M. Nishiwaki and F. Nagasaki 
(1960), K.K. Chapskii (1963), J. King (1964), etc. (V.H.) 



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 2' ^ 1' ^ 4' ^ ^-..36-34. 

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 tne equa- 
tor and 49° N 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 Japanese 
islands. 

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

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



92 

CALIFORNIA SEA LION 
Zaiophus californianus (Lesson, 1828) 

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

California. 
1858. Otaria gilliespii MacBain. Proc. Edinb. R. Phys. Soc, 1, p. 422. 

California. 
1866. Otaria japonica. Peters. Monatsschr. K. Preuss. Akad. Wiss., 

Berlin, p. 669. Sea of Japan. (V.H.) 

77 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. 

78 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. Lev) 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.) 



93 










11 Fig. 53. California sea lion, Zalophus califomiamis (figure by N.N. 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° N lat.), i.e., about 
8,000 km. Nevertheless, the species identity of the three populations is 
indubitable. 

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



94 




77 Fig. 54. Skull of the California sea lion, Zalophus califomianus (figure by 

N.N. Kondakov). 



(reconstructed range); 2) Z. с wollebaecki (Sivertsen, 1953) — Galapagos 
Islands; and 3) Z с 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 
EUerman and Morrison-Scott, 1966*) to the sighting of this species on 



95 




78 Fig. 55. Range of the Japanese sea lion, Zalophus califomianus 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 
79 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 A. I. 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. 



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- 
mochefys 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 CaUorhinus Gray, 1859 

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

1866. Arctocephalus. Gill. Proc. Essex. Inst. 5, p. 11, Nee Geoffray Saint- 
Hilaire et Cuvier, 1826 {A. pusilus Schreber, 1776). 
80 1892. Callotaria. Palmer. Proc. Biol. Soc. Washington, 7, p. 126. Substi- 
tute for CaUorhinus 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 on 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° N lat.) and the coastal belt on the eastern coast of Japan 
(36 - 38° N 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° N 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) 



98 

NORTHERN FUR SEAL^^ 
CaUorhinus 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. V.H.). 
81 1811. Phoca nigra. Pallas. Zoogr. rosso-asiat., 1, p. 107. "Darnie Kuril'- 
skie o-va" [Remote Kuril Islands] ("Ex ulterioribus insulis curili- 
cis"). The description is evidently of a young (black) animal. 

1828. Otaria kracheninnikovl Lesson. Diet, 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 califomianus. Gray. Catal. of Seals and Whales, Brit. 
Mus., p. 51. Monterey. California. 

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

1898. CaUorhinus curilensis. lordan et Clarck. Ibid., p. 45. Tyulenii Island, 
Sakhalin. (V.H.)^* 



Only species of the genus. 



Diagnosis 



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 



^' 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. 

^^ 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.) 



99 




81 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, v/hile 
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 
82 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 




82 Fig. 57. Skull of adult male fur seal, Callorhinus ursinus (figure by N.N. Kondakov). 




82 Fig. 58. Skull of adult female fur seal, Callorhinus ursinus (figure by N.N. Kondakov). 

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



Age in Years 


She 


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 (r = 
232.3), in females 182-200 mm (x = 189); width at the zygoma in males 
135 - 142 mm (Jc = 138.7), in females HI - 116 mm (x = 114.3); maximum 
width of skull in males 118-131 mm (x = 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 (x = 54). 



101 



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

Age in Years Weight 

1 15-20 

2 about 20 

3 about 29 

4 35-36 

5 about 55 

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) 
3 


25.6 


4 


29.1 


5 


30.7 


6 


35.5 


7 


33.0 


8 


41.0 


9 


31 


(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 
83 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, 



102 





83 Fig. 59. Head of a bull and a female fur seal, Bering Island, 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 



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 
84 of fur seals are often seen in winter in Olyutorsk Gulf and close 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 




84 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 



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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 
86 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° N lat. and 133-136° E long.). The southern 
boundary of their extent passes along 36-37°N 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 



106 

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

1. Commander fur seal, C. 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, С 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. 

87 3. Pribilov Alaskan fur seal, C. ursinus cynocephalus (Wallbaum, 1792) 
(syn. califomianus, 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 



107 



"■^^NV 




86 Fig. 62. Nostril area of the fur seal, Callorhinus ursinus (figure by N.N. 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, 



m M7914 So^ ^^^> 



(•ai)}33iAairA'4d H 11 4У 




87 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 arty 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.) 

88 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 on 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 a§ 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 master serves as the 
predominant food (Table 6). 
90 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 followed 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. 



110 



89 Table 6. Food of the fur seal in the western part of the Pacific Ocean (Arsen'ev and 

Fedorov, 1964) 



Pacific waters of 
Japan 



Sea of Japan 



Sea of Okhotsk 



Bering Sea, region 
of Commander Islands 



Fishes 

Headlight fish, 
Diaphus sp. 



Fishes 

Pacific salmon, 
Oncorhynchiis sp. 



Fishes 



Fishes 



Japanese anchovy. Pacific salmon, 
Engraulis japo- Oncorhynchus sp. 



Gissu, Ptero- 
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 cali- 
fomiense 

Anchovy, 
Notoscopelus 
elongatus 

Pacific saury, 
Cololabis 



Humpback salmon, 
Oncorhynchus 
gorbuscha 

Alaska pollock, 
Theragra chalco- 
^amma 

Atka mackerel, 
Pleurogrammus 
rrwnopterygius 

Asian greenling, 
Pleurogrammus 
azonus 

Sand fishes, Tricho- 
dontidae 

Cephalopoda 



Squids 



Pacific salmon, 
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. 



Watasenia scintillans Asian greenling, 

Pleurogrammus 
Gonatus fabricii azonus 

Gonatus master 

Cephalopoda 
Ommatostrephes sloani- 
pacificus Squids 

Gonatus sp. 



Humpback salmon, 
Oncorhynchus 
gorbuscha 

Alaska рюИоск, 
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. 



Ill 



1 

Lantern fish, 
Electrona sp. 

Barracuda, 

Sphyraena sp. 
Pacific cod, 

Gadus macroce- 

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 master 
Chiroteuthis veranyi 
Ommatostrephes 
sloani-pacificus 



Gonatus fabricii 
Gonatus magister 

Gonatopsis 

borealis 
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 



H2 



91 Table 7. Food of the fur seal in the eastern part of the Paciflc 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 
Maffiisudes baryosoma 
Myctophidae 
Tarletonbiania crenularis 
Cololabis saira 
Merlucius productus 
Syngnathus califomiensis 
Trachipterus trachipterus 
Trachurus symmetricus 
Brama rayi 

Medialuna califomiensis 
Scomber japonicus 
Sebastodes sp. 
Sebastodes jordani 
Anaplopoma fimbria 
Atherinopsis califomiensis 
Citharidithus sp. 
Liopsetta adlis 
Polichthyus notatus 

Cephalopoda 

Tremoctopus sp. 
Loligo opalescens 
Onychoteuthis sp. 
Onychoteuthis banksii 
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 
Salrrw 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 banksii 
Gonatus sp. 
Gonatus fabricii 
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 
Arwtopterus pharap 
Microgadus proximus 
Theragra chalcogramma 
Gasterosteus aculeatus 
Sebastodes sp. 
Sebastodes ahitus 
Anoplopoma fimbria 
Pleurogrammus monopterygus 
Reinhardtius hippoglossoides 
Aptocyclus ventricosus 
Trichodon trichodon 
Ammodytes hexapterus 
Bathymaster sigfiatus 
Anarhichas orientalis 
Atheresthes stomais 
Hippoglossus stenolepis 

Cephalopoda 

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



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 m^ 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. By 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 t-o 
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 
91 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. 



115 




92 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 



116 














93 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 JFor 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 




W t 



f* 



Щ^М^:^Ф^ ;^.-'-Ai:*i 





95 Fig. 69. Harem temales returning to the beach after being out to sea on 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 

96 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). 



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 
98 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° N 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 



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 
99 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 
to 15° С but the animal is seen more often at temperatures of 6 to 11° С 

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 iive 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° N lat.); seven — eight-year-oW 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. 



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. 
100 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- 
ern 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° N 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 



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). 




101 Fig. 73. A group of "black" pups. Tyulen' Island, July, 1968 (photograph by 

V.A. Arsen'ev). 



126 

101 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 mohths. 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. 

102 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, 

101 





Table 8. 


Age 


-related gestating females (Arsen'ev and Fedorov, 


1964) 




years 






Gestating among them 


Age, 


No. of females 
in sample 


Number 


Percent 



Western part of Pacific Ocean 

4 847 413 53 

5 637 531 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 

7 545 434 80 

8 609 519 85 

9 555 501 90 
10 513 455 89 
10+ 2,641 2,171 82 



127 



102 Table 9. Change in length and weight of embryos in Japanese waters (Arsen'ev and 

Fedorov, 1964) 









Length, 


cm 




Month 




Males 




Females 






Number in 
sample 




Average 
length 


Number in 
sample 


Average 
length 


February 

March 

April 

May 

June 


5 

17 

151 

125 

18 




27.0 
33.6 
43.9 
51.5 
57.0 


12 

3 

146 

116 

24 


13.2 
31.2 
42.2 
50.8 
53.9 








Weight, kg 




Month 




Males 




Females 






Number in 
sample 




Average 
weight 


Number in 
sample 


Average 
weight 


February 

March 

April 

May 

June 


1 

17 

151 

125 

18 




0.4 
1.3 
2.1 
3.7 
6.2 


6 

3 

146 

116 

24 


0.1 
0.8 
1.9 
3.3 

4.8 



103 



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 
sample 


Average 
length 


Number in 
sample 


Average 
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 


7 


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 
sample 


Average 
weight 


Number in 
sample 


Average 
weight 


1 


1 


23.0 


— 


— 


2 


47 


24.0 


1 


25.0 


3 


404 


31.5 


11 


25.6 


4 


190 


40.9 


40 


29.1 


5 


21 


56.4 


26 


30.7 


6 


9 


71.0 


10 


35.5 


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 



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. 





103 Fig. 74. Annual marks (layers) on the canines of fur seals. A — male, В — female 

(figure by N.N. Kondakov). 



130 










104 Fig. 75. Large bull fur seal with a 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 
104 (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 



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 tiemditode, 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 



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 
stmmosum 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 oif 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 
106 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. 



133 

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





Subspecies 


of fur seals 




Helminth species 








Commander 


Kuril 


Alaskan 


Phocitrema fusiforme 


+ 






Admocephalus septentrionalis 


+ 






Cestobothrium glaciale 


+ 






Diphyllobothrium krotovi 


+ 


+ 


+ 


Diphyllobothrium lanceolatum 


+ 






Diphyllobothrium macrocephalus 


+ 






Diphyllobothrium tetrapterus 






+ 


Anisalds pacificus 


+ 






Contracaecum osculatum 


+ 






Phocascaris phocae 


+ 


+ 




Terranova azarasi 


+ 






Terranova decipiens 


+ 


+ 


+ 


Uncinaria lucasi 






+• 


Bolbosoma bobrovoi 




+ 




Bolbosoma nipponicum. 




+ 




Corynosoma semerme 






+ 


Corynosoma villosum 






+ 


Corynosoma strumoswn 


+ 




+ 


Corynosoma ventronudum 


+ 







•At the end of the 1960s U. 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, i.e., 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 л11 ages: 5%, 10%, and 15%. This 
question has not been finally resolved. 



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 
107 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 Uons 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. 




107 Fig. 76. Steller's sea lion, Eumetopias jubatus, in a rookery of the northern fur 

seal. Mednyi Island (photograph by S.V. Marakov). 



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). 
108 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, 1964a*). 

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 



138 

yielded the toiai number of mothers. By determining the number of 
parents of both sexes, the number of young males that could be killed 
no 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 
so 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 



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 US 
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 
112 1910, the leasing system was abolished and the government established 
its control over fur seal hunting. As a result of the measures adopted. 




111 



Fig. 78. Young female sleeping on the coast. Bering Island, July, 1967 (photo- 
graph by P.G. 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 scieiitists 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 Pribilov 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). 



фГ"' -' " ^'^^P?^ 




Ill Fig. 79. Year-old male fur seal sleeping on water. Bering Island, October, 1958 

(photograph by S.V. Marakov). 



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 
113 of byproducts — meat, blubber, and liver. The carcass is used for feeding 
ranch-maintained fur aniinals, 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 



^^ During 1963-1965 there was a marked decline in all the populations of fur seals, 
which shaфly 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 a 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 abo^'e. 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 



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 
114 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 dorsoventralfy. 

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 



144 

with the paroccipital and not directed downward. An 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). 
ТЪе 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: 

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. 
115 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 



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), i.e., 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 (Hydmrga 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. 
116 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 leonind). 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. 



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, 
118 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, lO-incisored (I j) 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 (I |) seals of the subtropical regions of 
the Northern hemisphere covering monk seals (Monachus) as also the 
antarctic seals Lobodon, Ommatophoca, Hydnirga, and Leptonychotes; 
and 3) Cystophorinae, 6-incisored (l \) 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, Hydnirga). 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, 
Halichoems, 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 Halichoems 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 (Halichoems 
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 schauinslandi), ribbon 
seal (Phoca fasciata), and two forms of the common seal (Ph. vitulina), 
i.e., pagophilic (Ph. v. largha) and pagophobic (Ph. v. 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 
p. 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,^^ 
i.e., 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 entiire territorial waters of the USSR except the Black Sea, 

^^ Couibining Phoca vitulina and Ph. 1агфа into a single species. 



151 



120 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. 
(1С Ch.) 

Key to Species of True Seals (Phocidae) 

Identification Based on External Features 

1 (16). Upper incisors six (three on each side of jaw). Claws well devel- 

oped on fore as well as hind flippers. 

2 ( 3). 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. . . . 
Bearded seal, Erignathus barbatus (pp. 166-211) 

3 ( 2). 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. 

4 ( 5). 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) 

5 ( 4). 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 




120 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 K.K. Chapskii). 



152 

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. 

6 (13). Color of hair coat usually dark and spotted, dorsally with 

fairly sharp light-colored streaks or speckled, or altogether 
monochromatic, dark. 

7 ( 8). 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- 
121 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)^^ 

8(7). 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. 

9 (10). Cheek teeth with vertically disposed high accessory cusps. 
Color monochromatic. Light-colored gaps or dark-colored spots 
absent Baikal seal, Phoca sibirica (pp. 290 - 306) 

10 ( 9). Cheek teeth with accessory cusps directed forward and backward 

(fanlike). Color of skin not monochromatic, with pattern. 

11 (12). 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 

Ringed seal, Phoca hispida (pp. 218 - 260) 

12 (11). 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 

Caspian seal, Phoca caspica (pp. 260 - 290) 

121 Fig. 82. Structure of the crowns of upper and lower cheek teeth of the common 

seal, Phoca vitulina (figure by K.K. Chapskii). 

*^See note at the end of this Key. 



153 



A В С 

121 Fig. 83. Structure of the crowns of lower (cheek) teeth in the small species of 
the genus Phoca. A — Baikal seal, Ph. sibirica; В — ringed seal, Ph. hispida; 

С — Caspian seal. Ph. caspica (figure by K.K. Chapskii). 

13 (6). 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. 

14 (15). 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). 
Greenland seal, Phoca groenlandica (pp. 369 - 436) 

122 15 (14). 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 

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). 



154 



16 ( 1). Upper incisors four (two on each side of jaw); color either dull 

and dark or bright and spotted. 

17 (18). 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 

pairs of teats 

Mediterranean monk seal, Monachus monachus (pp. 502-515) 

18 (17). 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.) 



123 



Identification Based on Skull Features 

1 (16). 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). 

2(3). 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 




122 Fig. 85. Rostral-facial part of the skull of seals. A — ribbon seal, Phoca (Histrio- 

phoca) fasciata (lateral contours with convexity); В — hooded seal, Cystophora 
cristata (concave lateral contours) (figure by K.K. Chapskii). 



155 



below resembles a trapezoid 

Bearded seal, Erignathus barbatus (pp. 166-211) 

3 ( 2). Alveoli of upper incisors distinctly compressed laterally. 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. 

4 (13). 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. 

5 ( 6). 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 part (Fig. 87) 

Gray or long-snouted seal, Halichoerus grypus (pp. 454-495) 

6 ( 5). 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; 





\jifv4t^ 




123 Fig. 86. Position of the infraorbital fpramen. A — seal of the genus Phoca; В and 

C— gray seal, Halichoerus grypus: 1 — anterior lower edge of orbits in all seals of 

the genus Phoca s. 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). 



156 



/ l\\ 




123 Fig. 87. Contour of nostril. A — larga, Phoca vitulina [largha]; В — gray seal, Hali- 

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. 
7(8). 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 

124 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)^* 

8(7). 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 
general tooth row. 
9 (10). 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 

(Fig. 89) Caspian seal, Phoca caspica (pp. 260 - 290) 

10 ( 9). 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 
rostral part of skull less than length of orbit. 



^^ See note at the end of this Key. 



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; В — 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 

Baikal seal, Phoca sibirica (pp. 290-306) 

12 (11). Lower cheek teeth (except' the first) with deflected (fanlike) 

anterior and posterior cusps, considerably shorter than middle 



158 

125 (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) 

13 ( 4). 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. 

14 (15). 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. 
Greenland seal, Phoca groenlandica (pp. 369-436) 

15 (14). 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) 

126 16 ( 1). 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). 

17 (18). 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) 

18 (17). 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). . . . 
Hooded seal, Cystophora cristata (p. 524). (K. 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 



159 



Pacific species, i.e., 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.) 




125 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 K.K. Chapskii). A — pagophobic forms of the com- 
mon seal: Atlantic, Phoca v. vitulina; Kuril (Island), Ph. v. Kurilensis; Richard's, 
Ph. V. richardi; В — pagophilic form of seal: Far-Eastern larga. Ph. v. largha. 




125 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 K.K. Chapskii). 



160 




125 Fig. 92. Nasal and premaxillary bones in different forms of the common seal, 

Phoca vituUna. A — ^Atlantic common seal, Ph. v. vitulina; В — larga. Ph. v. 1агфа 
(figure by K.K. Chapskii): 1 — nasal bone; 2 — premaxillary bone; 3 — facial part 
of nasal bone; 4 — maxillary part of nasal bone. 



127 



Subfamily of True, or 10-incisored, Seals 



Subfamily PHOCINAE Gill, 1866^^ 

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. 



^^ 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.). 



161 





126 Fig. 93. Structure of the palato-choanal region in different forms of the common 

seal, Phoca vitulina. A — ^Atlantic common seal, Phoca v. vitulina; В — 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: 



I 



2' 



cl 



pl, Mi = 34.20 
4 1 



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 



' Sometimes a second molar is seen. 



162 

the pups of the pagophobic members of the subgenus of ringed seals, 
Phoca s. str. (genus Phoca) and the bearded seal (genus Erignathus), 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 Hali- 
choerus, and the pagophobic form of the subgenus Phoca s. str., genus 
Phoca) whelp and molt on land. 
129 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 Mioc^e) 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). 



^^ The karyotype of the bearded seal (2 n = 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 




tc 



\u 



s 



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 Halichoenis 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.) 

130 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 



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: 

T 3 ^ 1 „ 4 , 1 ^, 
I 5, с J, P -, M -=34. 

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, 



j>xi^&ii2L£^ 



A 

130 Fig. 95. Structure of the crown of undamaged lower teeth in a young bearded 

seal, Erignathus barbatus. A — from the inner side; В — from the outer side (figure 
by K.K. Chapskii). 



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- 
131 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 л = 34 and in Phocinae, 2 n = 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 Erignathus 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 Erignathus 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 (i.e., 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. Miiller. 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. 



167 

1811. Phoca nautica. Pallas. Zoographia Rosso-Asiatica, I, p. 108. Sea 

of Okhotsk. 
1811. Phoca albigena. Pallas. Ibid., p. 109. Kamchatka. 
1817. Phoca lachtac. Desmarest. Now. Diet. Se. Nat., 25, p. 581. Paeifie 

Ocean. 
1828. Phoca parsonsii. Lesson. Diet. Class, d. Hist. Nat., 13, p. 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 

132 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- 

133 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"). 



168 




132 



Fig. 96. Bearded seal, Erignathus barbatus (photograph by G.M. 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 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 
134 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 



170 




133 Fig. 98. Color of the head of a five-year-old bearded seal. Barents Sea, May, 

1963 (photograph by V.A. Potelov). 



135 



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 



171 



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. 





134 Fig. 99. Skull of the bearded seal, Erignathus barbatus (figure by N.N. Kondakov). 



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 
136 height decreases from 72 to 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 (Lev) 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 (Lev) 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 (Le) (males 
240 cm, females 239 cm) while in a straight line (Lev) males are larger 
(225 cm) than females (217 cm) (Kosygin, 1966).^ 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 mm,^'^ 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); 



^^ 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. 

^ 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). 

'^^ Data of Chapskii (1938), Potelov (1968*), and G.M. Kosygin. 



174 

the Barents Sea males weigh on 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 
138 floes in very deep waters. Particularly F. Nansen (north of 82° N 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, i.e., 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 



175 



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176 

the meridian 40° E 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, Kanin- 
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 Ziemlya. 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 
139 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 (lokhel'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, i.e., 
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 (V. 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 



^ 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, 
A.N. Tyulin, K.K. Chapskii, I.K. Yakimovich, and others. 



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 in summer. 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 (Razumovskii, 
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 
140 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° E 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 



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). 

141 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; 

^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 




s 



t 



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; 
142 and others), no one has conclusively defined their morphological charac- 
teristics. "The distinctive features oiE. 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, E. 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 {Lev) 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, E. 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 



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 (Lev) 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 
143 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. 



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 



ййж?ЙЯ«№ ё".. f-- 




'^-'9^-- 



143 Fig. 102. A female bearded seal emerging from water. Barents Sea, April, 1967 

(photograph by V.A. Potelov). 



184 

Russk Bend and Kara 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 
144 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 northern part 



185 



145 



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° N lat. could be regarded as the tentative boundary between 
the northern and Sakhalin populations of the Okhotsk Pacific bearded 
seal. 




144 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 (Tikhom.irov, 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 

^' 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. 



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 
146 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 Ul'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 Ul'bansk 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 



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 
147 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 the 
limit. 



189 




147 Fig. 104. Pacific bearded seal on an ice floe. Bering Sea (photograph by G.M. 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 



190 

composition of the fish is quite diverse with the polar rod holding 
first place, followed by plaice {Pleuronectes platessa), common sand eel 
(Ammodytes hexaptems), herring (Clupea harengus), and more rarely 
148 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, as for example the Shoina. Crustaceans are represented almost 
exclusively by decapods, mainly crab {Ну as 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 (WoUeback, 1907), capelin and sea bass are 
consumed from time to time. 

From among the crustaceans, this seal ronsumes 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^lcry- 
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 — Echiums echiurus 
and polychaetes— ylren/co/a and Harmathoe, and from among the other 



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 
149 {Sclerocrangon and some others); sea slater^, and amphipods are sec- 
ondary. Mollusks are almost invariably represented by gastropods {Buc- 
cinum, 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. 



^ 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 coarctatd), 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, Chiysodomus) 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 coarctatd), 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 My a, 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 



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 
151 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 



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 
152 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.^^ 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 

^^ Others suggest that it was so named because it "hops" while negotiating on ice (or on 
land). 



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 (Ul'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. 
153 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" (S. 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 



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, 
154 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 



199 




154 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, 1966d*).^^ 

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 (WoUeback, 1907; Laktionov, 1946). The mating 



^^ 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). 



200 

behavior involves a state of excitation in the animals, which pursue each 
other, and perhaps "mating calls," recognized as a manifestation of sexual 
reflexes (Dubrovskii, 1937). 
155 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 

^^An even smaller figure of 10% has been estimated (Tarasevich, 1963). 



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, 
i.e., 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 
156 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 (V. 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 

^^ 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. 



202 



t 




156 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, i.e., 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. 



203 



In the Sea of Okhotsk pupping probably occurs everywhere but 
mainly at places where these seals concentrate (see p. 178) in the spring, 
i.e., 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 
157 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 






157 Fig. 107. Newborn Pacific bearded seal. Bering Sea (photograph by G.M. Kosygin). 



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). 
158 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, Lev) 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 (Lev), 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 (Lev); 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 



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 
AG. 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 (Lev 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 (Lev) or 
170-175 cm (Lc). 

In terms of percentages, the average adult growth dynamics can be 
159 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; 
S. 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 

^^ 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. 



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). 
160 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 



208 

liver of the bearded seal can be a host for over a thousand O. 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. lanceolatum is encountered most often. 
The nematodes infecting the stomach and intestines are Contracaecum 
osculatum, Terranova d^ecipiens, 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, С 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; 
161 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.^ 

Population dynamics. Because of the lack of population data at any 
initial level, its dynamics could not be determined accurately. At the end 



^^ 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 II 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, i.e., 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 



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 
162 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 m^); 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 



211 

animals were caught in this region while 1,239 animals were caught in the 
"northern ice floes," ^^ i.e., 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. 
163 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.) 

■'^ According to the annual report of the Norwegian Fishing Directorate on Seal Hunting 
for 1963 (1964). 



, 212 

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. Diet. 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 

164 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 

^^ 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.) 

^^ 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 oval.^^ 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 



•'^ Deviations of this characteristic are most common in the ribbon seal. 

■'^ In the case of the Atlantic common seal and its ecological (pagophobic) Pacific coun- 
teфarts (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. 



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 
166 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'N 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 




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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 
167 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 (EUerman 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 {Ph. vitulina)^^ 
Pusa Scopoli, 1777, with three species (Ph. hispida, Ph. caspica, ahd 
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, 
i.e., 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 {Ph. 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 role in marine-animal-based industries. The Greenland [harp], 
Caspian, and ringed seals occupy first place in this respect. (K.Ch.)'^^ 



"•^ As a result of recent studies (Chapskii, 1967, 1969), the monotypical state of this 
subgenus has become extremely doubtful, if not erroneous. 

'^^ For the key to species of the genus, see under the characteristics of the family (p. 
151). 



218 

Subgenus of Ringed Seals 
Subgenus Pusa Scopoli, 1777 

RINGED SEAL 
Phoca (Pusa) hispida Schreber, 1775 

1776. Phoca foetida. Fabricius in Miiller, Zoologiae Danicae Prodromus, 

p. VIII, Nomen nudum. Greenland. 
1778. Phoca vitulina botnica. Gmelin. Linn. Syst. Nat. Ed. XIII, 1: 63. 

Gulf of Bothnia, Baltic Sea. 
168 1775. Phoca hispida. Schreber. Die Saiigethiere, Table LXXXVI. 1776, 

3: 312. Northern Atlantic. 
1811. Phoca ochotensis. Pallas. Zoographia Rosso Asiatica, p. 117. 

Northern part of the Sea of Okhotsk, between Tauisk and 

Gizhiginsk bays. 
1820. Phoca annelata. Nilsson. Scand. Fauna, 1: 362. Renamed 

Ph. foetida Fabricius. Baltic. 
1828. Phoca schreberL Lesson. Die. Class, d'hist. Nat., p. 414. North 

Atlantic. 
1828. Phoca communis. F. Cuvier. Dents mamm. 
1839. {Phoca communis F. Cuvier) B. var. Octonotata. Kutorga. Bull. 

Soc. Imp. Nat. Moscow, p. 189. Neva. 
1839. {Phoca communis F. Cuvier) B. var. Undulata. Ibid., p. 191. Neva. 
1899. Phoca foetida var. saimensis. Nordquist. Acta Soc. Fauna Flora 

Fenn., 15, 7: 28. I^ke Saimaa, Finland. 
1899. Phoca foetida var. ladogensis. Nordquist. Ibid., p. 33, Ladoga. 
1902. Phoca {Pusa) hispida gichigensis. J. Allen. Bull. Amer. Mus. N. H. 

16: 478. Sea of Okhotsk, Gizhiga. 
1921. Phoca hispida pygmaea. Zukowsky. Arch. f. Naturgesch. 87 A, 10: 

183. Barents Sea at 77° 3' N lat. and 49° 40' E long, (pygmy ringed 

seal); (V.H.) 
1929. Phoca hispida pomororum. Smirnov. Dokl. Ak. Nauk. (C. R. Acad. 

Sc.) Leningrad, p. 95. Western coast of Novaya Zemlya. 
1929. Phoca hispida pomororum natio rochmistrovi. Smirnov. Dokl. Ak. 

Nauk. (C. R. Acad. Sc.) Leningrad, p. 95. Sumsk environs, Onezhsk 

Bay, White Sea. 
1929. Phoca hispida birulal Smirnov. Dok. Ak. Nauk. (C. R. Acad. Sc.) 

Leningrad, p. 96. Lyakhov Island, Novosibirsk Islands. 
1935. Phoca hispida krascheninnikovi. S. Naumov and Smirnov. 

N. A. Smirnov, V. Adlerberg, Vinogradov, Smirnov, Flerov. Arctic 

Animals. Leningrad, 1935. Bering Sea. (V.H.) 



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) 




169 



Fig. 109. Ringed seal, Phoca hispida (figure by N.N. Kondakov). 



220 

169 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 olive- 
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 no significant age-related 
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 





170 



Fig. 110. Skull of a ringed seal, Phoca hispida (figure by N.N. Kondakov). 



222 

bony septum in the choanae runs 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 
171 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 Sea"*^ were: condylobasal 
length 161.5-182 mm; mastoid width 96.5-108.5 mm; width at the 

''^ According to the data of the 2^ological Museum of Moscow University. 



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 
(seepp. 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 s. 1., the common seal or larga (Phoca vitulina L.) 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). 
172 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 in 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 



''' 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), N. Smimov (1903), Wolleback and Knipovich (1907*), 
Formozov (1929), Sdobnikov (1933*), and Surkov (1965*). 



226 

regions of the White Sea and forms rookeries there*"* 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 
174 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 northwest 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. 

^Some 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. Smimov (1903). V. Nikol'skii (1927), Bianki (1965), 
Golenchenko (1961, 1963*), Nazarenko (1%7, 1968), and others. The unpublished data of 
A.P. Golenchenko, Yu.I. Nazarenko, V.A. Potelov, and others were also used here. 

*^ 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), raumov (1935), Moskalenko (1945), 
A.P. Golenchenko, Yu.I. Nazarenko, V.A. Potelova, Tsapko (1958*), and others. 

''* Based on the published and unpublished observations and information of several 
researchers: Krivosheya (1884), Gorbunov (1929), Lepin (1935*), Klyuge (1936), 
M.I. Vladimirskaya, A.P. Golenchenko, A.N. Dubrovskii, A.I. Zubkov, V.A Potelov, and 
K.K. Chapskii. 

''^ 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. 

''^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), Aul, Ling, and Paaver (1957), Bergman (I960*), Leis (1960), and V.Z. Zheglov. 



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 "ludas" [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 
175 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. 



''^ The distribution of the Ladoga seal has been described from the data of Chapskii and 
co-authors (1932*), Golenchenko (1935), N. Smirnov and colleagues (1954*), Sorokin and 
co-authors (1957, 1958*), A.S. Sokolov (1958*), A.A. 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 en the 
observations of several individuals and on data from the literature, including Zhitkov (1913), 
Heptner (1930, 1936), S. Naumov (1931), Kolyushchev (1933), Probatov (1933), Ии- 
mov (1935), Urvantsev (1935), Kiфichnikov (1937), Mikhel' (1937), Chernigovskii (1935), 
Antipin (1939*), Rutilevskii (1939), Laktionov (1947*), Ushakov (1953), Mikhailov (1958), 
G.G. Galkin, L.I. Leonov, V.F. Nikitin, V.A Potelov, AN. Tyulin, K.K. 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 the 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 

5^ According to the data of lokhel'son (1898), Buturlin (1913), Arsen'ev (1935), Mikhel' 
(1937), Fedoseev (1966b), and others. 



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- 
176 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, Il'pi 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 

^^ On the Commander Islands themselves, "random finds of stray animals" (including 
gestating females) have been reported at various points on the coast (Marakov, 1968). 



230 




176 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*). 

177 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 



231 

Basin inclusively, the Labrador coast, northern edge of Newfoundland, 
the northernmost part of St. Lawrence Bay (in the south up to 50° N 
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, 
178 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. annelatd). 

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. ^^ 

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 

^^ The greater тофЬо1о|1са1 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 (Miiller-Wille, 1969). 



232 







05 



(i, 



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 rochmistrovi). 

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 (x = 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). 



234 

The condylobasal length of the skull is 152.2 - 176 mm {x = 163) (S. 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) 
179 Ph. (P.) h. saimensis Nordquist, 1899 — Lake Saimaa in Finland; (3) 
Ph. (P.) h. beauforttana 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, i.e., 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).^^^ 
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, 1966b, c). Similar figures, i.e., 600,000 to 865,000 
were arrived at in subsequent aerial surveys (Fedoseev, 1968). The 

^''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). 



235 

differences in the calculations are explained as due to error in the inethod 
of calculation (G.A 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 
180 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 

^^ 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; 



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 

181 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 breed- 
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 

182 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. 




■■■■' ■■'^■'.;,-}^' 



ЫШ^^'У^К-Ш!^к^ %?M 




*^y 



^s>..'. \ ■■■%S \ 




181 Fig. 114. Immature ringed seal, Phoca hispida ochotensis, Bering Sea, June, 1964 

(photograph by G.M. Kosygin). 



239 

Food. The ringed seal feeds on fish and crustaceans; other animal 
groups (moUusks, 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, i.e., 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 Gammanis, Themisto, and 
Anonyx), shrimps (genera Pasiphaea, Pandalus, and Spirontocaris), mysids 
{Mysis), and partly sea slaters (Mesidothea and Idothea) play a lesser 
183 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). 



^^ The depth of submergence can sometimes reach 60 ш 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 ai, 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 



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 nugax), a mysid {My sis oculatd), a euphausid — ^black-eye 
(Thysanoessa inermis), and shrimps {Eualus gaimardi). 
184 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 (AS. 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 



242 



'Щ .Ф 



Ai ■: 



i.*"'. "■s.t./'V,'^'.^5fr:';-',v^'' ■■•'!.;■■ ■ 




184 Fig. 115. Ringed seal on a sandy shoal. White Sea, Sevemye Koshki, September 

20, 1970 (photograph by A.G. Beloborodov). 

185 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 



243 

(N.A 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, whien 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 
186 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 

^^ Air holes can also be concealed under a snow cover as a result of drifting snow. 



244 

also encx)untered directly in deep snow at almost a level surface. Such a 
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 (AN. 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, i.e., 
after every full meal. 

The more active spring behavior among the Okhotsk ringed seals 
disposed on broken drifting ice floes is somewhat different. "Rookeries 



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 
187 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; i.e., 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 



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 oj 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 
188 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; 



^^ 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, 1966d*). 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 
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). 





189 Fig. 116. Ringed seal on ice floes. Barents Sea, April, 1967 (photograph by 

V.A. Potelov). 



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 
190 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 (N. Smirnov, 1903; Knipovich, 1907; Ognev, 1935; A.N. Dubrovskii; 
M.I. Vladimirskaya; Yu.I. Nazarenko; V.A. 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 (ICirpichnikov, 
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 fi-om the first appearance of the newborn almost until 

^^ According to Tikhomirov (1970), hovyever, barren seals in the Far East constituted 
only about 7%. 



250 



191 



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. 




'ЩШШ 







190 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 (Lc, 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 ("kavadei" 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 



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 
193 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, 

^ From the same sources (see p. 208). 



254 

1953; Delyamure, 1955; A. 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 tr^matodes, 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., Diplogonopoms 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 g. 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, С validum, C. hadveni, С 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, Diplogonopoms 
tetrapterus, Terranova sp., Skrjabinaria spirocauda, Parafilaroides arcticus, 
P. kroicheninnikovi, Corynosoma hadveni, and С 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. 



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 
194 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, i.e., 
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 v^th 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, i.e., 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 



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 (Lev), inhabiting the arctic seas, and also Lake Ladoga. They do 
not form dense herds, remain on the shore ice in venter, 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.) 

195 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 

^^ Apart from Lake Ladoga in which the population is not high. 



257 




195 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 
196 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 (AP. 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, i.e., a 
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. 



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 
197 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 
(AP. 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). 



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 (Рта) 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., p. 414. Caspian Sea 

(nom. nud.). 
1929. Caspiopusa dierzawini. Dybowski. Ibid., p. 414. Caspian Sea (nom. 

nud.). (V.H.) 

198 Diagnosis 

The body length including the tail along a straight line (Lev) 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, i.e., 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 Юevezal', 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 с г 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^ray, 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. 




199 Fig. 120. Caspian seal, Phoca caspica. A — adult male (with variegated colors); 

В — ^adult female (mother) with pup (figure by N.N. Kondakov). 



ет^'^-^^'" 




р > 



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о >^ 

V о 



^ Е S 



■5 iJ 



о "? 

•к "^ 
I' в 



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а. 


_м 


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263 



r-^^^n,^,^,,.,, 




200 



Fig. 121. Juvenile Caspian seal, Phoca caspica (figure by N.N. 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 
200 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). 




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 



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) 
201 (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 ( ases 
it may even be absent (Gadzhiev, 1957). As in many other species, the 



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) 
202 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 (x = 5.9). 

The condylobasal length among the females varies from 
171.8-182.0 mm (x = 175.1), width at the mastoid 85.0-95.5 mm (x = 
89.6), width at the zygoma 89.3-98.7 mm (x = 93.6), width above the 
canines 23.2-27.0 mm (x = 24.8), and interorbital width 4.0-6.7 mm (M 
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. 
hispidd) and Baikal seal {Ph. sibiricd). These are no doubt extremely close 
but the interpretation of the Caspian seal (like the Baikal) as a subspecies 



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.) 

203 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 

A 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 (B.I. 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 Iran. (K.Ch.) 



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 
204 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. 



^^ 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. 



270 

they enter Tyub-Karagansk Bay (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 
205 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 



271 




Fig. 124. Adult female Caspian seal, February, 1953 (photograph by Yu.V. Kurochkin). 



206 



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 



273 




207 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 



274 




209 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. 
208 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, 1948b*, 
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 



276 • 

a 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 
209 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 (Юeinenberg, 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. 



277 

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 mbdng 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 (B.I. 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 



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 
211 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). 



279 

The mating season commences roughly mid-February, i.e., 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, i.e., 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 
212 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 a 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, 1965a*). 

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 jon 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 gi\e 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. 



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 
214 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, 1965a*). 

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 a "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 {Haliaeetus leucoryphus), white-tailed [gray] 
eagle (Я. albicilld), and partly the golden eagle (Aquila chrysaetus), but 



283 



-sSfVisiiiji 




213 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 
215 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 Uterature (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; Ciureana 
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 cofymbi 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 



*' "It 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*). 

^ It should be borne in mind that none of the three species of helminths that are 
common to the Caspian seal and the northern species "is 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.) 



285 




216 Fig. 129. Caspian seal in open water pool. Gur'evsk Channel, February 1, 1966 

(photograph by G. 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- 
216 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 



286 

of the animal at that time. However, some population reduction did 
occuf 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 
217 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, i.e., 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 



287 







218 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 K.K. 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, i.e., 
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 88,0(Ю (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. 



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 

218 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 

219 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). 



290 

These seals have never had a 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.) 

220 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, no. 2, p. 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. 

221 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 



^^ Belyak — white pup that has not shed the embryonic hair coat; khubun or nerpyash- 
molted underyearling; chemysh — one-, two-, and three -year-old seals; yalovka — unfertilized 
female; /пагАм -gestating or whelping female; and argal or sekach — mature male. 



291 




220 



Fig. 131. Adult Baikal seal, Phoca sibirica (figure by N.N. 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). 



292 



Jv*y-'V*' 







221 Fig. 133. Head of adult male Baikal seal, Phoca sibirica (photograph by V.D. Pastukhov). 




221 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. ТЪе teeth in the upper jaw are 



293 





222 



Fig. 135. Skull of the Baikal seal, Phoca sibirica (figure by N.N. Kondakov). 







^<\ -s 








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 



294 




222 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. 

223 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," i.e., 
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 craniologic^l 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 honidus, 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 



296 




224 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 
224 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 Acanthogammams. 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 fislies consumed by the Baikal seal (Pastukhov, 1956b) 

Species of fish 



Baikal omul, Coregonus autumnalis mi^atorius Georgi 

Stone sculpin, Paracottus kneri Dyb. 

Sand sculpin, P. kessleri Dyb. 

Big-heated sculpin, Batrachocottus baicalensis Dyb. 

Spotfin Baikal sculpin, B. multiradiatus Berg. 

Yellowfin Baikal sculpin, Cottocomephonis grewig^ Dyb. 

Longfin Baikal sculpin, С inermis Jakowl. 

Red Baikal sculpin, Procottus jettelesi Dyb. 

Humped sculpin, Asprocottus megalops Gratzian 

Big red Baikal sculpin, Procottus jettelesi major Tal. 

Big Baikal oil-fish (big golomanka), Comephorus 

baicalensis Pall. 
Lesser Baikal oil-fish (lesser golomanka), С dybowskii 

Korotn. 
Common perch, Perca fluviatilis L. 
Burbot, Lota lota L. 
Ide, Leuciscus idus L. 

Siberian dace, Leuciscus leuciscus baicalensis Dyb. 
Roach, Rutilus rutilus Pall. 



No. of seals 


Frequency, 


in which this 


% 


species was 




detected 




8 


6.6 


3 


2.5 


22 


18.0 


3 


2.5 


4 


3.3 


70 


57.4 


70 


57.4 


9 


7.4 


2 


1.6 


1 


0.8 


60 


49.2 


92 


75.4 


5 


4.1 


2 


1.6 


1 


0.8 


1 


0.8 


6 


4.9 



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. 

225 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 

226 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 



299 




225 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. 



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 
227 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. 



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, 
228 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). 



302 



A 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 
229 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 Oxiset 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, С о. 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). 



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 
230 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 a growiiig 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, 



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 
range. 

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 
231 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.) 



307 

Subgenus of True Seals 
Subgenus Phoca Linnaeus, 1758 

COMMON SEAL, LARGA^ 
Phoca (Phoca) vitulina Linnaeus, 1758 

1758. Phoca угп//шя. Linnaeus. Syst. Nat. Ed. X, 1:38. Northern part of 
the Baltic Sea. 
232 1811. Phoca largha. Pallas. Zoogr. Rosso- Asiatica, 113. Eastern coast 
of Kamchatka. 
1811. Phoca canina. Pallas. 2^ogr. Rosso- Asiatica, 1:114. Atlantic (?). 
1820. Phoca variagata. Nilsson. Skand. Fauna, 1:359. Atlantic (?). 

1823. Phoca scopulicola. Thienemann. Reise im Norden Europas, 1:59. 
Iceland. 

1824. Phoca littorea. Thienemann. Ibid. Northern Russia (?). 

1828. Phoca linnaeL Lesson. Diction, classique d'Histoire Natur., 

13:415. European waters. 
1828. Phoca thienemannii Lesson. Ibid, 13:414. New name for Phoca 

scopulicola Thienem. 
1828. Phoca chorisiL Lesson. Ibid., 13:417. Kamchatka. 
1844. Phoca nummularis. Temminck. Fauna Japon, p. 3. Japan.^^ 
1864. Halicyon richardit Gray. Proc. Zoolog. Soc. Lond., p. 28. Van- 
couver Island. 
1902. Phoca ochotensis. J. Allen. Bull. Amer. Mus. Nat. Hist., 16:480. 

Gizhiga Estuary, Sea of Okhotsk. Nee Phoca ochotensis Pallas, 

1811. 
1902. Phoca ochotensis macrodens. J. Allen. Ibid., 16, p. 483. Avachinsk 

Bay, Kamchatka. 
1902. Phoca steinegen. J. Allen. Ibid, 16, p. 485. Commander Islands 

(Bering Island). 
1902. Phoca richardi pribilofensis. J. Allen. Ibid, 16, p. 495. St. Paul 

Island, Pribilov Islands. 
1936. Phoca vitulina largha natio pallasii S. Naumov and N. Smirnov. 

Tr. Vses. n-i. in-ta rybnogo khozyaistva i okeanografii (VNIRO), 

3:177. Sea of Okhotsk. 
1941. Phoca petersL Mohr. Zoolog. Anzeiger, 133, p. 49. Coasts of the 

Korean Peninsula. 



^ Also mottled seal (Far East), rock seal (sometimes in Murman). 

^^ 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. 



308 

1942. Phoca ochotensis kuritensis. 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). 
(KCh.) 

233 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 

^The larga described here is the arctic Far Eastern form (for more details, see 
"Geographic Variation"). 



309 




233 Fig. 143. Common seal, Phoca v. vitulina. Atlantic Ocean (figure by N.N. Kondakov). 




235 Fig. 144. Skull of the common seal, Phoca v. vitulina (figure by N.N. Kondakov). 



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, i.e., 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 
234 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. 



69 



Breeding not associated with ice floes. 







D_ 03 



i2 z 



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. 



312 

Two main types are noticed in the structure of the skull (Fig. 144) 

235 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 (Lev) was around 150 cm in most cases, the 
largest of them reaching 160-165 cm^^ (Havinga, 1933). The maximum 

236 length evidently not measured in a straight line, but along the dorsal 
surface (Lc) among the Norwegian seals is 180 cm (CoUett, 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". 

^^ 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. 

^ 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 



^ Data on the Atlantic seals are extremely scant and fragmentary and there is no accurate 
information on their actual ages. Moreover, the oldest arid the largest rarely attract hunters. 



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 
237 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.) 



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 
239 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 



316 





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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; G.A. 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 Penin- 
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 

^'^ 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 (Smimov, 1903). 

^^ The larga hardly spreads along the polar coast of Eastern Siberia into the west beyond 
L6ng Strait although there are references (Rutilevskii, 1962) that it perhaps reaches even 
the estuary of the Indigirka River. 

'^ For accurate definitions of "larga" and "Kuril or island seal," see under "Geographic 
Variation". 



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 
240 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) on 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). 



319 

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 (Тук 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). 
241 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 Vakhir 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)^^ (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 



'^ 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. 




[L, 



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° N 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.) 

244 Geographic Variation 

Early in this century concepts regarding the species and subspecies of 
seals of the subgenus Phoca s. 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. v. largha, vicariating with the European Ph. v. vitulina, 
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 



325 




245 Fig. 147. Typical coloration of an adult pagophilic larga, Ph. v. largha. Verkho- 

turov Island, Kuril range, August, 1971 (photograph by S.V. Marakov). 

245 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 
N. Smirnov, 1936). One of them, n. 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, n. largha, inhabiting the Bering Sea, 



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, limnaei, thienemannii). 

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). 
246 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 
on 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) v. curilensis Inukai, 1945^^ (syns. 
chorisii (?), stejnegeri, nummularis,^^ macrodens, and insulaiis; the name 
richardi was also used sometimes). 



^® Known on the Kamchatka and Commander Islands from the end of the nineteenth 
century under the local name "antrus" or "antur". 

^' The first name for this form should be stejnegeri J. Allen, 1902 (editor's note). (VH.) 
^ See the note on p. 307 (synonyms). 



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. 




247 Fig. 148. Dark-colored Far Eastern island seal ("antur"), Ph. v. kurilensis. Iturup 

Island, Kuril, July, 1966 (photograph by S.V. Marakov). 



328 




247 Fig. 149. Color details on the ventral side of the female island seal, Ph. v. kurilen- 

sis, Iturup Island, July, 1966 (photograph by S.V. Marakov). 




248 Fig. 150. Variation in dark coloration of the adult island seal, Ph. v. kurilensis, 

Iturup Island, July, 1966 (photograph by S.V. Marakov). 



329 




248 Fig. 151. Island seal ("antur"), Ph. v. 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 

249 islands; (2) Ph. v. concolor De Kay, 1842 — ^western Atlantic American 



330 

and Greenland waters; and (3) Ph. v. mellonae 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 20^^), 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). 

^^ This calculation was based on the data of Tikhomirov (1966b). 



331 

A 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 
250 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 Kamni, Srednii, Ptichii, Utichii, and also Malyi 
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 19^2 but later dou- 
bled (27,500-35,000) (P.G. Nikulin). In some very large rookeries on 
the Shantarsk Islands, especially on the Sivuch'i Kamni (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). 



332 




250 Fig. 152. Larga, Ph. v. largjha (left) and island seal ("antur"), Ph. v. kurilensis, 

Mednyi Island, June, 1969 (photograph by S.V. Marakov). 



251 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 that it is not rare along 
eastern Kamchatka and Koryak Land (from Cape Olyutorsk to Cape 



334 

Navarin). It is quite abundant also in the coastal waters of the western 
and southern regions of the Gulf of Anadyr; 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, 1935a*). 

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 
252 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). 

^^ At the beginning of this century, this seal was known in Murman as "kamenka". 



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. 
253 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, 1964a*; 
Velizhanin, 1967). Some animals (males) at places, for example on the 
Lx)vushki Islands, rest on small isolated rocks almost fully submerged in 
the strait and overgrown with algae (Belkin, 1964b*). 






253 Fig. 155. Typical habitat of the island seal ("antur"), Ph. v. kurilensis, on Mednyi 

Island. Commander Islands, Zapalata Bay, June 6, 1962. Sea otters also live here 
(photograph by S.V. Marakov). 



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, i.e., 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- 
254 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 



337 




254 Fig. 156. Largas on a drifting ice floe in Litka Strait, Karaginsk Island, Bering 

Sea, eariy 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 Tyma. 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 
255 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 Юyuchi 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 m 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 



339 

advantage of the approaching herring; in summer it feeds partly on 
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 leucomaenis] rudd etc. Some 
256 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 moUusks, 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 cephalqpod 
moUusks. The following fishes were detected in the stomach of largcis: 
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 



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. 
257 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 Sderocrangon and amphipods of the genus Gammams 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.) 
(Gol'tsev, 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 E. 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) 



342 

correspondingly increases. Among the benthic and demersal fishes con- 
258 sumed are the flounder, halibut, goby, stichaeid blennies, and some 
poachers; among decapods the snow crab and others. The quantum of 
cephalopod moUusks 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 Coitus) 
and greenlings (Hexagrammidae); from among the invertebrates, they 
consume predominantly cephalopod moUusks, 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 



343 

the coasts 10 km away from its estuary, on Cape 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, 
saffi-on 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, 
1966b). 

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 
259 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, i.e., roughly 15 - 19.5% of the pup's 
own body weight. The young seals voraciously consumed various types 



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 rookeries 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 
260 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 or reef, forming a large herd in 
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. 



346 

At the conclusion of molt the larga again becomes highly active, 
261 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 usually 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 



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. 
262 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 




261 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 '4vhelped 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 on 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 



349 

beach rookeries. They abandon them only at the time of reproduction, 
which takes place in the neighboring ice floes (Belkin, 1964). 
263 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 cx)mmencement of the period of molt (Tikhomirov, 
1964, 1966).^^ 

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 
264 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 (CoUett, 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 

^ 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. 



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). 
265 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, 1966d*). 

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; Fines, 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, i.e., 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, 



353 

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). 
266 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%.*'* 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, 

^ According to E.A. 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. 



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 exte?nds for 
150-200 km^ (Tikhomirov, 1965b*). 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 (Lev) 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. 
267 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 



^ The hair coat of the newborn in the region of Peter the Great Gulf is a smoky-gray 
(Kosygin and Tikhomirov, 1969). 

^ 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 fi^om 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) or 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). 



356 
















.^'.iumsMMMm^^^'i^:if^^§l^g 



268 Fig. 158. Intensely molting white large pup. Bering Sea (photograph by G.M. Kosygin). 



268 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 



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- 
269 cated (Havinga, 1933) that, after the first year, the Dutch seals grow 
to 95.7-113 cm (length in a straight line. Lev); 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, i.e.. 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 



^•Ф ■■■^■.^> . '-i^a-i.^;' 




270 Fig. 159. Under-yeariing larga. Benng island, May, 1969 (photograph by LP. 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. 



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 
270 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 over a 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, i.e., 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). 



360 

271 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 so on 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 



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 
Ol'ga 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 
272 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 tnincatum 



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, С semerme, C. validum, С hadveni, C. 
ventronudum, and С osmerl 

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 С 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 T. 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 rec- 
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 5. spirocauda, parasitizing the heart, large blood vessels, 
and lungs.^^ 
273 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 



^^ Information on the helminths of the species under study pubHshed 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 p. 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 



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 
274 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 



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 
275 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 
to 10,000-15,000 animals. Its proportion in the seals killed by hunting 



366 

using ships did not, however, exceed 9.4% in the best years (Tikhomirov, 
1966aj. 

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. 
276 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). 



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 
277 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 



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. (KCh.) 

278 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., p. 170. Greenland, Ice- 
land. 

1811. Phoca dorsata. Pallas. Zoogr, Rosso-Asiatica, I, p. 112. White Sea. 
(V.H.) 



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.) 




279 Fig. 160. A — male Ьаф seal, Pagophilus groenlandica, with the final color of 

the "wing pattern"; В — adult female haф seal in the "semiwinged" phase of 
color (incomplete pattern of the "wings" covered in sparse spots) (figure by 
N.N. Kondakov). 



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 
279 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 ЮevezaГ, 
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 



372 



'A^'- 







i0'' 







281 Fig. 161. Main changes in age-related skin patterns among Ьаф seals. Top to bot- 

tom: gray form (1-3 years old); transitional phase of female (average 3-5 years 
old); and final coloration, i.e., the winged form (5 years and above) (photograph 
by R.Sh. Khuzin). 



373 

base of the tail. On a flat skin they resemble a typical horseshoe or 
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 
280 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, i.e., 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, 
i.e., spotted gray (or grayish) and the final "wing pattern".^^ 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, 



^ Our special literature used to refer to such 20-year-old females as the "second phase, 
with gray wings" (N. Smimov, 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). 



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% 
281 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 fi-om 
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 

^^ 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). 



375 

the spotted gray to the final phase is extremely short among males and 
often the intermediate semiwinged phase is bypassed. Nevertheless, 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 'Ving 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. 
282 The second stage, forms with a "semiwinged pattern," is distin- 
guished by fully developed contours of the lower edge of the '4ving 
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 

^^ 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 larghd) 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 





283 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. 



377 

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 
283 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 



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.^^ A paired fold on the lateral fronto-sincipital surface from 
where the temporal muscles originates is more intensely manifest among 
284 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 foUowingi^"* (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 



'' This is not a characteristic of the race (N. Smimov, 1929) but is of sex-related sec- 
ondary importance (Plekhanov, 1932* ; Chapskii, 1952; and others). 
** Data pertain to the White Sea population. 



379 

the bony 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 
'4ving 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- 
285 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 

Up to one month 41.5 (n = 28) 

From one to three years 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 22.7 

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. Lev 
varies from 153-188 cm and Lc from 167-202 cm^^ (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 (x = 217.0); zygomatic width 
107.0-142.0 mm; mastoid width 113.0-133.3 mm (x = 122.4); width of 

^^ 229 cm in an extraordinary case. 



381 

the snout above the canines 30.0-43.5 mm (x = 36.2); and the smallest 
interorbital width 8.8-21.0 mm (x = 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 (x = 119.5); the ros- 

286 tral width at the canines 28.6-40.0 mm (x = 32.8); and the smallest 

interorbital width 9.1 - 16.0 mm {x = 12.7) (Khuzin, 1963, 1967). 

The reliability of the craniological differences between males and 
females is also confirmed by statistical variance analysis (t 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,(ХЮ km from the breeding site of the latter and the transgressions 
of Jan Mayen animals into our waters are very rare (see p. 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 s. 1., Ph. groenlandica. The seals oiPhoca 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 



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. 
287 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 1п1ефге1а11оп of the structural features the skull is somewhat debatable in 
the light of some new facts (Chapskii, in litt.). 



383 



288 



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), i.e., 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 



^ 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). 



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 
289 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 



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 
290 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 

'^ Observations in the Kara Sea, southeast of the Blagopoluchiya Strait (K.K. Chapskii). 



387 




290 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). 



291 



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, as 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 ^ven forty ye^rs 
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 (x = 183). The skull dimen- 
sions (according to the same source) are: condylobasal length in males 
(37) 200-234 mm (x = 217), in females (300) 200.5-223 mm {x = 
209.5); mastoid width in males (37) 113-127 mm (x = 121), in females 
(27) 111-124.5 mm (x = 117); rostral width (at level of canines) in 
males (38) 32-46 mm (x = 36.2), in females (27) 27.5-37.0 mm (x = 
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. 

* * * 

Outside the USSR, usually only one subspecies is recognized, i.e., 
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 (x = 176.5), of females (127) 156-201 cm 
(x = 175.5); condylobasal length in males (39) 200.0-219.5 mm (x = 
208.5), in females (41) 191.0-219.0 mm (x = 204.5); mastoid width in 
males (40) 109.0-124.0 mm (x = 117.5), in females (41) 109.0-123.5 mm 
(x = 116.0 mm); rostral width in males (40) 30.0-40.5 mm (x = 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 (t = 8.52) 
and in the length of the palate (r = 4.61) as well as in the condylobasal 
length and mastoid width of the skull, etc. 



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, 
292 1963) and does not differ from the latter biochemically in the protein 
polymorphism (Meller, Naevdal and Valen, 1966*).^^ 

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). 

^'^ 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 (N. Smirnov, 
1928)^^1 was initially put at 600,000-700,000 (P.A. Rudakov and 
N.V. Provorov). Somewhat later, double this figure or 1,300,000 was 
cited (N.V. Provorov). 

The first postwar experiiiient 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 
and Khuzin, 1959)^^^ 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 
293 (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, 

^^^ That is, without pups. 

^^2 Data for the 1930s -1940s have been taken from Khuzin (1972*). 

^^^ Assuming that all the 100,000 animals killed were pups. 



391 

293 Table L5. Variation in the area of concentration (rookeries) of harp seals in the White 

Sea, km^ (Yakovenko, 1967) 



Type of rookery 






Year of observation 




1953 1954 1955 1956 1957 1958 1959 1960 


Rookeries 
Rookeries 


in the lactation period (nurseries) 294 
in the molting period (molting) 85 


290 200 167 143 

75 64 58 58 


126 

48 


130 120 
46 48 


293 Table 16. 
areas 


Total average annual toll of the 
of the Barents Sea at five-year i 


harp seal in the White Sea and in adjoining 
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 
(i.e., 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 

294 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 venter, 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 2^mlya). 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, 
Gammams, Themisto, Gammarocanthys, and others), and also shrimps 
(Crangon sp., Sderocrangon boreas, Pandalus sp., etc.); among the 



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 
295 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; WoUeback, 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. raschii) 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 moUusk (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 



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' (i.e., 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 (N. 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. 
296 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 

^^ The "invasion" in Murman waters of this seal and the disappearance of cod there is 
a consequence of sharp atmospheric cooHng (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 



396 

harp seal is mainly dependent on 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 
297 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 (Юyuche,* 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 



397 




298 Fig. 165. Part of the Ьаф 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 m^, i.e., at a 
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 



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 sjx years, and adult females) covered a total 
area of 76 km^ in 1963 (Yakovenko, 1967), i.e., average of 2,200 animals 
per km^. In other words, there was one animal in an area of 455 m^ 
(average distance of 20 m from each other). In fact, however, in this 

298 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 

299 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 sopewhat 
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). 



^^^ Discontinuities formed between the frozen masses of ice floes, sometimes compressed 
(during compression) and sometimes diverging or enlarging (in low tide) are called "gapes" 
by the coastal hunters. 



400 

The last period, i.e., 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 
300 to be manifested predominantly in daylight hours which rapidly increase. 
Nevertheless, their activity does not cease at night. 

The summer behaviot 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 heids, 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 



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 
301 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, i.e., 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 




404 

a 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 seals are seen even in Septem- 
ber and later at alrnost 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, i.e., 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 



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, 
304 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 move 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 



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. 
305 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 

^^ There are some other views, although hardly substantiated, that the animals do not 
mate in water but only on ice floes (L. Popov, 1966). 



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 
306 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). 




ШШёйШй 



ШИШ: 






306 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 
307 has not been adequately proved. A thorough analysis of the generative 
organs leads more to a contrary conclusion (G. Nikol'skii, 1933; 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 

^''^ In calculating the population dynamics, the barren animals among the females were 
frequently not taken into consideration (Yakovenko, 1967). 



410 

species as a 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). 
308 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. It 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. ^^^ Evidently, taking into consideration such anomalous 
instances, the total duration of the whelping period is sometimes put 

^^ In 1966, one white female pup with a firm hair coat was detected as late as April 26 
(M.Ya. Yakovenko). 



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. 
309 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 molt. 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. Lev) and along the dorsal surface 



'ljV,tf к iffVr ,./^J/,J ^;- i > tf ^r , «S"^^^- "'■>' "".««t- 




309 Fig. 168. White pup of the harp seal. White Sea (photograph by A.V. Yablokov). 



^*^ Significantly, however, this author pointed out that not a single hunter sa\y more than 
two fetuses in one womb. 



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 

310 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 (Lev) or 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 

311 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 



414 







310 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 



415 



311 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% 



^^^ 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*). 

^^^ 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. 



416 






■^f "-"gf 





312 Fig. 171. Normally molting pup of the haq? 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 
313 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 



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). 
314 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 

313 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 


39.24 


29.35 


1.21 2.95 1.76 


7.73 


18.81 11.17 


0.51 


1.47 


0.82 



418 




314 Fig. 172. A typical underfed pup (dwarf) of the White Sea Ьаф 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. It 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, 



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 
315 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 in body length (Lc) of the White Sea harp seal (males) in relation to 

age (Yakovenko et 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 


9.7 


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 


11 


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 


2.5 


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 



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. 
317 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 

^^^See: V.G. Heptner et al. Mammals of the Soviet Union, vol. 2, pt. 1. 



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 v^'ith 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) 
318 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 (L. Popov, 1971*). 
This factor is of equally great relevance to the infant mortality of the 



424 



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. 




arkhangel'sk 



319 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). 



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 
319 herd was taken at roughly 20% in the calculations of population dynam- 
ics (N. Smirnov, 1928; P.A. Rudakov, K.K. Chapskii). 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 



426 

ducts of the liver. The cestodes Diphyllobothrium cordatum, D. schis- 
320 tochilus, Diplogonoporos tetraptems, 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 molted 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 

^^^ 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 

Sea herds 



Helminth 


From Jan Mayen 


From White Sea 




region 


region 


Orthosplanchnus arcticus 




+ 


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 
321 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. 



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. ^^"^ 

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 
322 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.) 



^^'' The numerical data were borrowed from R.Sh. Khuzin who processed the hunt- 
ing statistics pubhshed by Sivertsen (1941), Iversen (1927*), and the Norwegian Fishing 
Directorate. 



429 

Economic Importance 

The harp seal is of primary importance to hunting. Its potential resources 
on 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 
323 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 



430 



322 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 
for the 




Min. 


Max. 


Average 


Min. 


Max. 


Average 


period 


1875 - 1880 


19.3 


48.3 


30.7 


5.6 


9.1 


7.3 


38.1 


1881-1885 


8.9 


58.3 


26.6 


3.1 


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 


52.1 


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 


1911-1915 


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 


231.1 


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 


2.7 


131.8 


55.0 


— 


— 


— 


55.0 



♦Calculation based on material compiled by R.Sh. Khuzin, who used such primary sources 
as lort 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 



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. 
324 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: 

1946-1950 166,300 

1951 - 1955 151,700 

1955-1960 115,000 

1961 - 1965 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 



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, i.e., 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 
325 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 


17.7 


212.4 


1951 


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 


19.1 


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 


11.2 


105.0 


1962 


106.9 


8.3 


115.2 


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 



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 




'■^:*5.i=*i 



326 Fig. 174. Early spring rookery of adult males. White Sea, April 18, 1966 (photo- 

graph by Yu.I. Nazarenko). 



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, 

326 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 

327 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). 



326 Table 22. Norwegian hunting of the harp seal in the Jan Mayen Island region from the 

1880s (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 


7 


1,977 






.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, I960*). 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.^^^ 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 



115 



See: Proceedings of the Murman Scientific-Hunting Expedition, St. Petersburg, 1912. 



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, 
"right 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. 
328 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 vierfiissigen Thiere, 3, p. 277. Kuril 
Islands. 



^^^ "Kozha" is a local term used by the coastal people for the haф seal. 
^^^ 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, p. III. 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 




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); 

В — 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 



439 





330 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. 
331 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 (Ognev, 1935; S. Naumov and N. 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 iiiformation on the seasonal changes 
in weight. (V.A.) 



440 



1} 



''^''^'^^TrrTT.r.rn^W^ 




330 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.) 

332 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, i.e.. 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 



441 



но l^sn 160 




250 О 250 500 750 1000 кГП 



331 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). 



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, i.e., at the time of spring-summer rookeries on the 




332 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). 



443 

ice floes. From this viewpoint, it 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 
333 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 aire disposed on drifting ice floes far away from the coasts but 
usually overlying depths not exceeding 200 m. 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 



444 



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. 



334 



Table 24. Food items of ribbon seals (Shustov, L965a) 



Food item 


Number of cases 


Percentage of 








number of stomachs 








with food 


remains 




Sea of 


Bering 


Sea of 


Bering 




Okhotsk 


Sea 


Okhotsk 


Sea 


CRUSTACEANS 










Crangon dalli 


1 


— 


2.0 


— 


Nectocrangon lew 


— 


3 


— 


9.4 


Pandolopsis sp. 


— 


6 


— 


18.7 


Pandalus borealis 


— 


8 


— 


25.0 


Pandalus goniurus 


5 


1 


9.8 


3.1 


Eualus gaimardii 


— 


6 


— 


18.7 


Spirintocaris murdochi 


— 


1 


— 


3.1 


Lebbeus sp. 


— 


3 


— 


9.4 


Temisto sp. (2 species) 


— 


6 


— 


18.7 . 


Stilomysis grandis 


— 


2 


— 


6.2 


Amphipods 


— 


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, Aptocydus ventricosus 


1 


— 


2.0 


— 


Pacific cod, Gadus morrhua macrocephalus 


4 


— 


7.8 


— 



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). 

334 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 huriter 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 

335 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. Its 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 A.P. Shustov). 



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 May 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 
336 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 



448 




336 



Fig. 183. Young ribbon seal. Bering Sea (photograph by A.P. Shustov). 



Table 25. Variation in body length of seals with age (measured along the dorsal 
surface) (Tikhomirov, 1968) 



Age 


;, years 


Males 




Females 






No. of animals 


Average 


No. of animals 


Average 






measured 


length (cm) 


measured 


length (cm) 


1 




16 


132.0 


21 


129.0 


2 




20 


145.0 


22 


145.0 


3 




29 


155.0 


26 


152.0 


4 




33 


156.6 


28 


153.9 


5 




17 


165.0 


22 


160.2 


6 




20 


158.9 


15 


161.5 


7 




17 


160.1 


16 


168.0 


8 




19 


161.0 


5 


169.0 


9 




16 


168.1 


16 


163.9 


10 




21 


163.8 


10 


163.0 


11 




13 


165.0 


— 


— 


12 




14 


167.7 


19 


163.9 


13- 


14 


22 


165.2 


— 


— 


15- 


17 


26 


168.7 


12 


168.8 


18 and above 


15 


163.1 


13 


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- 
337 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 



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 stnimosum Rudolphi, known 
338 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 no 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 ol 
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). 



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.) 

339 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. 





339 Fig. 184. Adult male ribbon seal on an ice-floe. Bering Sea (photograph by 

A.P. Shustov). 



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. 
340 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. 



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- 
sory 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; 

4' ^T' ^i' m} = ^"^ °' ^T " ^^• 

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 
341 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," Hali- 
choems 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^i^ 
Halichoerm grypus (Fabricius, 1791) 

1791. Phoca grypus. Fabricius. Scrivter of Naturhist.-Selskabet, Kjoben- 
havn 1, p. 167, pi. 13. Greenland. 

118 '"pevyak" 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. 



455 

1820. Halichoems griseus. Nilsson. Scand. Fauna, Dagg. Djur., I, p. 377. 

Greenland. 
1851. Halichoems macrorhynchus. Hornschuh et Schilling. Arch. Natur- 

gesch., 17, p. 28: Baltic Sea. 
1851. Halichoems pachyrhynchus. Hornschuh et Schilling. Ibid. Baltic 

Sea. 
1886. Halichoems grypus var. atlantica. Nehring. Sitz.-Ber. Ges. Natur- 

forsch. Freunde, Berlin, p. 122. Western coast of Norway. 
1886. Halichoems grypus var. baltica. Nehring. Ibid. Baltic Sea. (V.H.) 

342 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 folds^^^ 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 

343 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 

^^^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. 



456 







342 Fig. 185. Gray seal ("tevyak"), Halichoerus grypus (figure by N.N. 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 



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, i.e., 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 
344 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 



458 





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