Part 3 This is the fourth book of Mammals of the Soviet Union, representing the third part of the second volume; it is devoted to descriptions of the orders of Soviet aquatic mammals — pinnipeds (Pinnipedia) and, in part, cetaceans (Cetacea), toothed whales (Odontoceti). In the sequence of descriptions from the “higher” to the “lower” orders adopted in this series, pinnipeds should have pre- ceded carnivores, 1.е., should have appeared in the second book. The group- ing of the orders at a higher level is given after G.G. Simpson (1945). The sequence of genera and species within the orders has been retained as before, 1.е., from the less specialized to the more specialized. The order of pinnipeds or seals (Pinnipedia) is described in this volume. The cohort of ungulates and carnivores represented in Soviet fauna by orders of artiodactyls, and perissodactyls, sirenians, carnivores, and pinnipeds thus comes to an end and the cohort of whales (Mutica) commences. The toothed whales are described in this volume. While it has not always been pos- sible to maintain a totally uniform description of the genera and species as in the volumes already published, in spite of every effort to do so, the sequence has been adhered to, with some excep- tions, in the case of Pinnipedia. But, it was impossible to use sane format in describing the toothed whales (Odontoceti). Only a brief morphological description has been given for many species, which is more or less adequate for identifying the species. General information on their distribution and fragmentary biological data are also given. In some cases the total absence of such information is indicated. fy ef hel и Mammals of the Soviet Union Volume II, Part 3 MAMMALS OF THE SOVIET UNION In Three Volumes Edited by Late V.G. Heptner and N.P. Naumov Vysshaya Shkola Publishers Moscow, 1976 Mammals of the Soviet Union VOLUME о Part 3 —_ PINNIPEDS AND TOOTHED WHALES Pinnipedia and Odontoceti Late V.G. Heptner, К.К. Chapskii, V.A. Arsen’ev _and У.Е. Sokolov Edited by Late V.G. Heptner Scientific Editor JAMES G. MEAD Smithsonian Institution Libraries and The National Science Foundation Washington, D.C. 1996 Smin-B86 SIL 005 Mlekopitayushchie Sovetskogo Soyuza In Three Volumes V.G. Heptner and N.P. Naumov, editors Vysshaya Shkola Publishers Moscow, 1976 Translator: P.M. Rao General Editor: Dr. V.S. Kothekar ©1996 Oxonian Press Pvt. Ltd., New Delhi Library of Congress Cataloging-in-Publication Data (Revised for v. 2, pt. 3) Geptner, V.G. (Vladimir Georgievich), 1901 - 1975. Mammals of the Soviet Union. Translation of: Mlekopitaiushchie Sovetskogo Soiuza. Vol. by V.G. Heptner and A.A. Sludskii; illustrators, A.N. Komarov and N.N. Kondakov. Vol. 2, pt. 2 includes index. Includes bibliographical references. Supt. of Docs. no.: $1 1.2:Ar7/2 Contents: v. 1. Artiodactyla and Perissodactyla— у. 2 pt. 2 Carnivora (hyaenas and cats)—pt. 3. Pinnipeds and toothed whales (Pinnipedia and Odontoceti) 1. Mammals—Soviet Union—Collected works. I. Nasimovich, A. A. П. Bannikov, Andrei Grigor’evich. Ш. Hoffmann, Robert $. IV. Sludskii, А. A. У. Title. QL728.S65G4713 1988 599.0947 85-600144 Translated and published for the Smithsonian Institution Libraries, pursuant to an agreement with the National Science Foundation, Washington, D.C., by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110 001 Printed in India at Baba Barkha Nath Printers, New Delhi UDC 596.5 We gratefully acknowledge the assistance rendered by the follow- ing persons, which made the publication of this monograph possi- ble: V.D. Pastukhov (Listvenichnoe, Baikal), late P.G. Nikulin and D.I. Chugunkov (Petropavlovsk-Kamchatskii), I.F. Golovlev, V.A. Zem- skii, late B.A. Zenkovich, M.V. Ivashin, L.A. Popov, and S.K. Klumov (Moscow), S.L. Delyamure (Simferopol’), A.S. Skryabin, V.V. Treshchev, and M.V. Yurakhno (and other colleagues at the Helminthological Lab- oratory, Crimean State University), V.D. Kokhanov (Kandalaksha), late В.1. Badamshin (Fort Shevchenko), F.Sh. Khuzin, М.Уа. Yakovenko (Murmansk), А.С. Beloborodov, Уи.Г. Nazarenko, and V.A. Potelov (Arkhangel’sk), A.A. Berzin, G.M. Kosygin, Yu.V. Kurochkin, and late Е.А. Tikhomirov (Vladivostok), А.Р. Shustov and С.А. Fedoseev (Maga- dan), and many others cited at the appropriate places in the text. Further, S.V. Marakov (Kirov), V.I. Krylov (Moscow), and some others made available to us their original photographs; due acknowledgments have been given under the photographs. We express our appreciation to all these persons, and to all those others without whose assistance this publication would not have been possible. The authors are extremely grateful to IP. Mitina and E.V. Zubchaninova of the Department of Vertebrate Zoology, Moscow University, who put in much effort toward the preparation of this manuscript for the press and its publication; and to V.I. Korotkova, Chief Librarian of the Zoological Museum, Moscow University, who readily supplied to the authors rare and almost inaccessible reference materials. The authors are particularly grateful to Prof. Г.1. Barabash- Nikiforov (Voronezh) and S.V. Marakov (Kirov) who undertook the formidable task of reading through the manuscript, and for their extremely valuable suggestions. The authors also express their gratitude to O.L. Rossolimo, Director, Zoological Museum, Moscow University. Much of the preparatory work for this publication was completed in this museum. Most of the reference materials utilized were published before the end of the 1960s, although a few books and articles from more recent years have been useful. References are cited in the-text by the author’s name without initials and with the year of publication. Initials are used only when the surnames are identical. References to authors of unpub- lished private communications are also made in parentheses but with initials and without the year. The bibliography at the end of the book covers only the references cited (although many more were reviewed), У1 excluding those listed under synonyms. As in the previous publications, the authors have included, quite naturally, their own unpublished mate- rial in the text. The preceding volumes of Mammals of the Soviet Union have been warmly received within and outside the USSR. The first three books in the series were translated into German. The authors hope that this vol- ume, too, will be as warmly received although they are aware of its several shortcomings. An explanation for this, although not a justification, is the complicated nature of the material, especially that of whales. Save for some individual species, the museums in the USSR contain practically no collections of this group. V.G. Heptner 7 | Fe А | | | < 2 — J} gf и ‘ y= FOREWORD TO THE ENGLISH EDITION The Smithsonian Institution Libraries, in cooperation with the National Science Foundation, has sponsored the translation into English of this and hundreds of other scientific and scholarly studies since 1960. The program, funded with Special Foreign Currency under the provisions of Public Law 480, represents an investment in the dissemination of knowledge to which the Smithsonian Institution is dedicated. One of the values of this translated volume is to give English readers an insight into another philosophical system that devoted nearly a cen- tury to the studies of marine mammals. This work is important because it has an abundance of data that were taken from commercial harvest, particularly from small cetaceans, that have been generally unavailable to the English-speaking community. A monograph of this magnitude, with such an extensive bibliography, serves as an excellent entré into the Russian literature. This volume is the last to be published under the general editor- ship of V.G. Heptner and N.P. Naumov; Volume II, part 3, Pinnipeds and Toothed-whales appeared in 1976, the year after Professor Hept- ner’s death. Only one more volume of the original series remains to be published in an English edition, Volume II, Part 1, Sea Cows and Carni- vores (Dogs, Bears, and Mustelids), but editing of the English language manuscript is about 60 percent complete, and it is the intention of the Smithsonian Libraries Translation Program to publish the volume as soon as possible. After a lapse of two decades, the series has been revived with the sponsorship of the Russian Academy of Sciences’ Institute of Evolu- tionary Animal Morphology and Ecology in Moscow, with a new series title reflecting recent political changes—Mammals of Russia and Border- ing Regions. The first volume in this new series is, as had been antici- pated, Usatye Kity [Baleen Whales], by V.E. Sokolov and V.A. Arsen’ev (1994, Nauka, Moscow, 208 pp.). A second volume on Zaitseobraznye [Lagomorphs] is in production. It is hoped (but by no means certain) that English editions of the revived series can be made available. Vill English readers interested in Dr. Heptner’s contributions as a mam- malogist should refer to the Foreword to the English Edition of Volume: I. Conventions used in rendering geographic names, first stated there, are reprinted here for the convenience of the reader. Geographic names are generally transliterated directly, but a few exceptions were permitted (e.g. Moscow instead of Moskva, translation rather than transliteration of certain modifiers of place names, such as Northern, rather than Sever- naya Dvina). Soviet administrative units are numerous, and the following equivalents were employed in translation: Krai, territory; oblast’, district; raion, region; guberniya (archaic), province. Also, in the original Russian text, rivers, mountain ranges, and cities are often not explicitly identified, the Soviet reader being presumed sufficiently familiar with the geography of the country to be able to understand from the context of the sentence what sort of place is referred to. Complicating the matter is the lack of articles as parts of speech in Russian. To assist the English reader, the following conventions have been adopted: if a river is referred to, an article precedes it; if a mountain range is referred to, it is translated as a plural; if a city is referred to, it is singular, and lacks the article. Examples are: the Ural (river); the Urals (mountains); Ural’sk (city). Geographic place names are also inflected in Russian, and these have been simplified by omitting transliteration of the inflected ending. For example, the Russian phrase у Yaroslavskoi 1 Kostromskoi oblastyakh is translated “in the Yaroslavl’ and Kostroma districts.” In cases where the nominative form of the place name has an -sk ending this is, however, transliterated (e.g., Omsk); when a Russian “soft sign” is employed in a place name, this is transliterated as an apostrophe (e.g., Khar’kov). As in any translation, particularly from one orthography into another, there arise problems of standardization of names. For geographic names the National Geographic Atlas of the World, 4th edition, 1975 was used for this volume (principally the map on pages 122 - 123). Some of the geographic terms like gulf, bay, inlet, strait [zaliv, bukhta, guba, proliv] are used in slightly different contexts in the original Russian text. Because of the large number of place names in this volume, it was not possible to verify all of them, and some inconsistencies are likely to occur. We would appreciate it if readers would bring any errors they may notice to our attention. The usage of the common terms “dolphin” and “porpoises” varies geographically. In North America these terms are used interchangeably in reference to members of the family Delphinidae (sensu Simpson 1945) | but only the term “porpoise” is used to refer to members of the fam- ily Phocoenidae (sensu Simpson 1945). In most other English speaking ix countries, including England, the term “dolphin” is restricted to refer- ence to the family Delphinidae. In Russian, the term “del’fin” (dolphin) is used to refer to all of the members of the family Delphinidae and the term “morskaya svinya” (Sea pig), which is equivalent to the meaning of the Latin roots of “porpoise” (porcus = pig + piscis = fish) is used to refer to the members of the family Phocoenidae. We have therefore chosen to use the more restrictive English usage in this volume. A few of the vernacular names, those where the Russian name was vastly different from the generally accepted English name, have been changed, but translations of Russian vernacular names were employed where there was not a generally accepted English term (e.g. the vernac- ular name of the genus Lagenorhynchus). There is no English vernacular name for members of the genus Mesoplodon; Mead prefers to trans- late the Russian term “remnezubov” (sword-toothed) as Mesoplodon, rather than a literal translation. In addition, recent systematic studies have shown that the appropriate scientific name for the Malay dolphin (Stenella dubia) is Stenella attenuata (= Pan-tropical spotted dolphin) and the English common name of the Bridled dolphin (Stenella frontalis) is Atlantic spotted dolphin. Some terms placed in brackets [ ] indicate that they are additions of the scientific editor and are not found in the Russian text. In statisti- cal references, x replaces М (mean), a much more accepted symbol for “mean” in English works. One further point of confusion not apparent when Volume I was translated also requires clarification; that is the English transliteration of the senior author’s surname. This begins with the fourth letter of the Cyrillic alphabet, which usually has а “С” sound in Russian. However, the surname was originally German, and in the original German began with the letter “Н” of the Latin alphabet. Since Cyrillic has no equivalent of “H” this is usually transliterated шо “С” in Russian. However, Hoff- mann, from conversations learned that Dr. Heptner preferred to use the original Germanic form of his surname rather than the transliterated ver- sion, which is rendered as Geptner. The rules of transliteration employed by the Library of Congress do not permit such flexibility, and the atten- tive reader may notice that Library of Congress cataloging employs the latter. The Series Scientific Editor expresses particular thanks to the edi- tor of this volume, Dr. James G. Mead, Curator of Marine Mammal Project, Department of Vertebrate Zoology, National Museum of Nat- ural History, Smithsonian Institution. Dr. Mead’s broad knowledge of all aspects of marine mammal biology was critical to the quality of this English edition, and I am grateful to him for the considerable time he has xX devoted to ensuring the accuracy of the volume. Thanks are also due to P.M. Rao and Dr. V.S. Kothekar, Translator and General Editor respec- tively of Amerind Publishing Company, New Delhi, India, for producing the original translation under the provisions of U.S. Public Law 480. Credit also must be given to the general editorial staff of Amerind Pub- lishing Co., who confirmed all of the technical names of the prey species and parasites, checked the bibliographic references and translated many obscure Russian scientific terms. Robert S. Hoffmann Series Scientific Editor Assistant Secretary for the Sciences Smithsonian Institution, Washington, DC James G. Mead Volume Scientific Editor Curator of Marine Mammals National Museum of Natural History FOREWORD This is the fourth book of Mammals of the Soviet Union, represent- ing the third part of the second volume; it is devoted to descriptions of the orders of our [Soviet] aquatic mammals—pinnipeds (Pinnipedia) and, in part, cetaceans (Cetacea), toothed whales (Odontoceti). In the sequence of descriptions from the “higher” to the “lower” orders adopted in this series, pinnipeds should have preceded carnivores, 1.е., should have appeared in the second book. The microsystem of the class is given below to elucidate the natural sequence and the relationships between orders. This is a simple and presently more widely used system, but pin- nipeds have been assigned the status of an order and not a suborder as commonly accepted. The grouping of the orders at a higher level is given after G.G. Simpson (1945). The sequence of genera and species within the orders has been retained as before, i.e., from the less specialized to the more specialized. The order of pinnipeds or seals (Pinnipedia) is described in this vol- ume. The cohort of ungulates and carnivores represented in our [Soviet] fauna by orders of artiodactyls and perissodactyls, sirenians, carnivores, and pinnipeds thus comes to an end and the cohort of whales (Mutica) commences. The toothed whales are described in this volume and the baleen whales will be taken up in the next volume. While it has not always been possible to maintain a totally uniform description of the genera and species as in the volumes already published, in spite of every effort to do so, the sequence has been adhered to, with some exceptions, in the case of Pinnipedia. But, it was impossible to use the same format in describing the toothed whales (Odontoceti). Firstly, the biology of nearly all the species, except the Black Sea species, is not adequately known even in respect of principal features, and information on their distribution is more scant. Only a brief morphological descrip- tion has been given for many species, which is more or less adequate for identifying the species. General information on their distribution and fragmentary biological data are also given. In some cases the total absence of such information is indicated. Secondly, the faunal compo- sition of the Soviet Pacific waters is not yet clear. While the presence ХИ of some species already reported calls for confirmation, several species known from the coastal waters of Japan have not been reported in our waters. Some of them are found on the American coasts at the same lati- tudes. Their presence in our waters even as “Stray” finds is highly proba- ble; such probable species have been indicated, included in the keys, and briefly described. In this regard the present volume has achieved its aim, not only to describe the known species, but also to draw attention to the unknown. Such species total 14 while the number of relatively well- known species in the Soviet Pacific waters is quite large. While using the keys and morphological characteristics of the Pacific dolphins, reported here for the first time from our waters, their identification should be checked with the available literature and museum specimens. Interest- ing finds could be anticipated in the Soviet Far East and each case of a “stray” report or capture should be studied carefully. The probable species and genera have been given without numbers and in small print. The ranges of distribution of every species have been compiled on the same principle as in the preceding books. However, in view of the specificity of the biology of marine species (long migrations), in many cases they are given in a more general form. The range extending beyond our waters is mainly based on the works of Van den Brink (1958), Schef- fer (1958), Hall and Kelson (1959), King (1964), Hershkovitz (1966), Siivonen (1967), and some others, mostly of Japanese authors. The lat- ter have been cited in the text. The entire synonymy has been selected on the principles adopted earlier (see Foreword to Vol. I and Vol. II, parts 1 and 2). In view of the scant indigenous literature on whales, their synonymy is given very briefly. A more complete list of the synonyms of the species of this order can be found in the monograph by Hershkovitz (1966). The total number of species constituting this class is roughly 3,500, of which about 300 are represented in our fauna (Heptner, 1956). Almost all the illustrations in this book are original with the exception of those of the toothed whales taken from Hall and Kelson (1959). The original drawings are by the well-known Russian artist N.N. Kondakov who, apart from being a talented artist, is an experienced zoologist. Other sources of the drawings have been acknowledged at the appropriate place. The contribution of each author to this volume is as follows. K.K. Chapskii wrote the general outline of the order Pinnipedia and the family of seals (Phocidae), gave the keys to the families of the order and species of the family of seals, and wrote the text on all the species of this family except for the Baikal seal, Phoca sibirica, and the ribbon seal, Phoca fasciata. V.A. Arsen’ev wrote the characteristics of the walruses (Odobenidae) and the eared seals (Otariidae), and the entire description хШ of walruses, sea lions, fur seals, and the ribbon and Baikal seals. He also wrote the sections on the distribution, biology, and economic importance of all the species of toothed whales (Odontoceti), the characteristics of the suborder of baleen whales (Mysticeti), and the outlines of all the species of this suborder which, as mentioned above, will be included in the next volume. V.E. Sokolov wrote the general outline of the order of cetaceans, suborder of toothed whales, and the description of all the genera of this suborder and the morphological characteristics of its species. V.G. Heptner wrote the introduction to the book, keys to the species of eared seals (Otariidae), and an outline of the genera and species of sea lions (Zalophus californianus). Moreover, he worked out the entire synonymy and participated in the compilation of notes on the distribution of many species. As in the preceding books, the individual authorship has been shown by initials within parentheses at the end of the pertinent section. The overall format, the system adopted, selection and sequence of the species, their scope, and the numbers and scope of the subspecies have been confirmed by V.G. Heptner. TABLE OF CONTENTS: FOREWORD TO THE ENGLISH EDITION FOREWORD CLASSIFICATION OF CLASS MAMMALIA KEY FOR IDENTIFYING ORDERS OF MAMMALS PART I. ORDER OF PINNIPEDS ORDER PINNIPEDIA ILLIGER, 1811 Cohort Ferungulata Simpson, 1945 Superorder Ferae Linnaeus, 1758 Order Pinnipedia Illiger, 1811 Family Odobenidae Allen, 1880 (Walruses) Genus Odobenus Brisson, 1762: Walruses Odobenus rosmarus (Linnaeus, 1758): Walrus Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Superfamily Otarioidea Smirnov, 1908 Family Otariidae Gill, 1866 (Eared Seals) Subfamily Otariinae Boetticher, 1934 (Sea Lions) Genus Eumetopias Gill, 1866: Steller’s Sea Lion Eumetopias jubatus (Schreber, 1776): Steller’s Sea Lion Diagnosis Description Taxonomy XXVII * Pages 713-718 in the Russian original. The English table of contents is not a literal translation. Geographic Distribution 69 Geographic Variation 72 Biology WZ Economic Importance 87 Genus Zalophus Gill, 1866: Northern Sea Lions 89 Zalophus californianus (Lesson, 1828): 92 California Sea Lion Diagnosis 92 Description 92 Geographic Distribution and Geographic Variation 93 Biology 96 Subfamily Arctocephalinae Boetticher, 1934: 96 (Fur Seals) Genus Callorhinus Gray, 1859: Northern Fur 96 Seals Callorhinus ursinus (Linnaeus, 1758): 98 Northern Fur Seal Diagnosis 98 Description 98 Taxonomy 101 Geographic Distribution 101 Geographic Variation 105 Biology 108 Economic Importance 141 Superfamily Phocoidea Smirnov, 1908 142 Family Phocidae Gray, 1825 (True Seals) 142 Subfamily Phocinae Gill, 1866 (True, or 160 10-incisored, Seals) Genus Erignathus Gill, 1866: Bearded Seals 164 Erignathus barbatus (Erxleben, 1777): 166 Bearded Seal Diagnosis 167 Description 167 Taxonomy 174 Geographic Distribution 174 Geographic Variation 181 Biology 182 Economic Importance 209 Genus Phoca Linnaeus, 1758: True Seals and 212 Ringed Seals Subgenus Pusa Scopoli, 1777: Ringed Seals 218 Phoca (Pusa) hispida Schreber, 1775: Ringed Seal Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Phoca (Pusa) caspica Gmelin, 1788: Caspian Seal Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Phoca (Pusa) sibirica Gmelin, 1788: Baikal Seal or Baikal Ringed Seal Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Subgenus Phoca Linnaeus, 1758: True Seals Phoca (Phoca) vitulina Linnaeus, 1758: Common Seal, Larga Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Subgenus Pagophilus Gray, 1844: Harp or Greenland Seals Phoca (Pagophilus) groenlandica Erxleben, 1777: Harp or Greenland Seal Diagnosis XVil 218 219 219 223 225. 291 234 256 260 260 261 266 267 269 269 286 290 290 290 295 295 295 295 304 307 307 308 308 313 315 323 330 364 369 369 370 ХУШ Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Subgenus Histriophoca: Ribbon Seals Phoca (Histriophoca) fasciata Zimmermann, 1783: Ribbon (Banded) Seal Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Genus Halichoerus Nilsson, 1820: Gray Seals Halichoerus grypus (Fabricius, 1791): Gray Seal Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Subfamily Monachinae Trouessart, 1904 (Monk Seals or 8-incisored Seals) Genus Monachus Flemming, 1822: Monk Seals Monachus monachus (Hermann, 1779): Monk Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance Subfamily Cystophorinae Gray, 1866 (Hooded Syl 381 382 387 389 429 436 436 437 437 440 440 442 442 451 452 454 455 455 466 467 469 472 493 495 499 502 502 502 507 508 512 512 519 519 Seals and Elephant Seals, or 6-incisored Seals) Genus Cystophora Nilsson, 1820: Hooded Seals Crystophora cristata Erxleben, 1777: Hooded Seal Diagnosis Description Taxonomy Geographic Distribution Geographic Variation Biology Economic Importance PART II. ORDER OF WHALES ORDER CETACEA BRISSON, 1762 Cohort Mutica Order Cetacea Brisson, 1762 Suborder Odontoceti Flower, 1867 Superfamily Delphinoidea Flower, 1864 Family Delphinidae Gray, 1821 (Dolphins) Genus Steno Gray, 1846: Rough-toothed Dolphins Steno bredanensis Lesson, 1828: Rough- toothed Dolphin Genus Stenella Gray, 1866: Spotted Dolphins Stenella coeruleoalba Meyen, 1833: Blue-white Striped Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology Stenella dubia [=айепиаа] С. Cuvier, 1812: Malay [Pan-tropical Spotted] Dolphin Stenella frontalis G. Cuvier, 1829: Bridled [Atlantic Spotted] Dolphin Stenella longirostris Gray, 1828: Long-snout [Spinner] Dolphin Genus Delphinus Linnaeus, 1758: Common Dolphins Delphinus delphis Linnaeus, 1758: Common Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Tursiops Gervais, 1855: Bottlenose Dolphins Tursiops truncatus Montagu, 1821: Bottlenose Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Lissodelphis Gloger, 1841: Right Whale Dolphins Lissodelphis borealis (Peale, 1848): Northern Right Whale Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology Lissodelphis peroni (Lacépéde, 1804): Southern Right Whale Dolphin (Peron’s Dolphin) Geographic Distribution Biology Genus Lagenorhynchus Gray, 1846: Shorthead Dolphins Lagenorhynchus (Lagenorhynchus) acutus Gray, 1828: Atlantic White-sided Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology 607 607 607 611 611 614 627 631 632 633 633 634 638 639 644 645 646 646 647 647 648 648 651 652 653 653 656 656 656 657 659 659 Lagenorhynchus (Lagenorhynchus) albirostris Gray, 1846: White-beaked Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology Lagenorhynchus (Lagenorhynchus) obliquidens Gill, 1865: Pacific White-sided Dolphin Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Lagenorhynchus (Peponocephala) electra Gray, 1846: Broadsnout Dolphin Genus Pseudorca Reinhardt, 1862: False Killer Whales Pseudorca crassidens Owen, 1846: False Killer Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Ecanomic importance Genus Orcinus Fitzinger, 1860: Killer Whales Orcinus orca Linnaeus, 1758 : Killer Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Grampus Gray, 1828: Risso’s Dolphins Grampus griseus G. Cuvier, 1812: Risso’s Dolphin Diagnosis Description Geographic Distribution ххИ Geographic Variation Biology Genus Globicephala Lesson, 1828: Pilot Whales Globicephala melaena Traill, 1809: Pilot Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Feresa Gray, 1870: Pygmy Killer Whales Feresa attenuata Gray, 1875: Pygmy Killer Whale Genus Phocoena G. Cuvier, 1817: Common Porpoises Phocoena phocoena Linnaeus, 1758: Common Porpoise Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Phocoenoides Andrews, 1911: Dall Porpoises Phocoenoides dalli True, 1885: Dall Propoise Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Neophocaena Palmer, 1899: Black Finless Porpoises Neophocaena phocaenoides G. Cuvier, 1829: Black Finless Porpoise Diagnosis Description Geographic Distribution Geographic Variation Biology 698 699 701 702 702 702 704 706 708 VAY 718 719 721 722 72.2. 722, 725 726 729 735 737 738 738 738 740 741 744 748 750 750 750 750 751 sv 752 Family Monodontidae Gray, 1821 (Narwhals) Genus Delphinapterus Lacépéde, 1804: Belugas or White Whales Delphinapterus leucas Pallas, 1776: Beluga or White Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Monodon Linnaeus, 1758: Narwhals (Narwhals or Unicorns) Monodon monoceros Linnaeus, 1758: Norwhal or Unicorn Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Superfamily Physeteroidea Gill, 1872 Family Physeteridae Gray, 1821 (Sperm Whales) Subfamily Physeterinae Flower, 1864 (Sperm Whales) Genus Physeter Linnaeus, 1758: Sperm Whales Physeter catodon Linnaeus, 1758: Sperm Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Subfamily Kogiinae Gill, 1871 (Dwarf Sperm Whales) Genus Kogia Gray, 1846: Dwarf Sperm Whales Kogia breviceps Blainville, 1838: Dwarf Sperm Whale Diagnosis Description ххШ 755 755 IBY 150 757) 761 763 765 785 790 ТЭД Е 792 792. 794 794 798 799 799 800 800 801 801 802 808 812 813 834 840 840 841 841 843 XXIV Geographic Distribution and Biology Kogia simus Owen, 1866: Owen’s Dwarf Sperm Whale Diagnosis Description Family Ziphiidae Gray, 1865 (Beaked Whales) Genus Berardius Duvernoy, 1851: Pacific Beaked Whales Berardius bairdi Stejneger, 1883: Baird’s Beaked Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Mesoplodon Gervais, 1850: Beaked Whales [Sword-tooth Dolphins] Mesoplodon (Mesoplodon) stejnegeri True, 1885: Stejneger’s Beaked Whale Diagnosis Description Geographic Distribution and Biology Mesoplodon (Mesoplodon) bidens Sowerby, 1804: Sowerby’s Beaked Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Genus Ziphius G. Cuvier, 1823: Goose-beak [Cuvier’s Beaked] Whales Ziphius cavirostris G. Cuvier, 1823: Cuvier’s Beaked Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance Genus Hyperoodon Lacépéde, 1804: Bottlenose Whales 843 845 845 845 846 849 850 850 850 854 854 855 861 862 866 866 866 868 869 869 869 872 873 873 875 876 876 876 878 882 882 883 884 Hyperoodon ampullatus Forster, 1770: Northern Bottlenose Whale Diagnosis Description Geographic Distribution Geographic Variation Biology Economic Importance LITERATURE CITED OTHER SOURCES INDEX OF LATIN NAMES OF ANIMALS 957 989 hee ie и ie (Na nahi >. Lai | & soy У by т | i ve Gi > a к icv a ae : т a4 CLASSIFICATION OF CLASS MAMMALIA In this publication the old and widely used system of the major units of the class, 1.е., orders, has been adopted. There is only one feature in it which cannot be considered universal: pinnipeds are considered an inde- pendent order and not a suborder of carnivores (Carnivora). However, this approach, too, has many supporters. In the system of present-day mammals, these two groups are separated naturally and fully, no less than other orders. However, the close genetic relationship between pinnipeds and land carnivores is striking and beyond doubt. The system of orders adopted here is well founded, being based on the morphology of present-day forms by M. Weber (1928) and of fossil forms by G.G. Simpson (1945). The grouping of orders into taxa of a much higher rank and the sequence of orders is after Simpson. A tendency toward an extreme division of the orders has recently developed. Thus it has been proposed that the order of marsupials (Marsupialia) be divided into three (Polyprotodontoidea, Caenolestoidea, and Diprotodontoidea) and even five (Didelphia, Dasyuria, Peramelia, Caenolestia, and Phalangeria); insectivores into four (Insectivora proper, Zalambdodonta, Macroscelidea, and Tupaioidea); cetaceans into two (Odontoceti and Mysticeti); primates into three (Lemuroidea, Simiae, and Tarsioidea); and artiodactyls into two (Tylopoda and Artiodactyla proper). The number of orders has thus increased from 18-19 to 30-31. None of these suggestions is yet well founded. Further, mammalogists dealing with extensive paleontological material do not generally favor extreme division. An order is primarily regarded as an integrating and not a differentiating concept. Otherwise, there is the risk of a tendency, as in ornithological macrosystematics, for the concept of an order to become essentially indistinguishable from the concept of a family, and sometimes even simply substituted for it. As a result, the scientific aspect of the system of vertebrates has undoubtedly suffered, and is suffering even now, from the excessive ХХУШ division of the system which sometimes uses the term ‘fishlike vertebrates and fish”. From the established viewpoint, Duplicidentata are regarded as an independent order (Lagomorpha). Thus, we have here 19 instead of 18 orders. Of these, 10 or 52.6% are represented in our fauna. One, 1.е., the sirenians (sea cow), is extinct. The system of class adopted here is given below. The orders repre- sented in the fauna of the USSR are marked with an asterisk. (V.H.) CLASS MAMMALIA Subclass PROTOTHERIA Order MONOTREMATA Subclass THERIA Infraclass METATHERIA Order MARSUPIALIA Infraclass EUTHERIA “Order INSECTITIVORA Order DERMOPTERA _| “Order CHIROPTERA Order PRIMATES Order EDENTATA Order PHOLIDOTA ee LAGOMORPHA Cohort UNGUICULATA Cohort GLIRES *Order RODENTIA Cohort MUTICA “Order СЕТАСЕА Superorder _| “Order CARVINORA FERAE “Order PINNIPEDIA Superorder Canon PROTUNGULATA Order TUBULIDENTATA FERUNGULATA Order PROBOSCIDEA BS ies —| Order HYRACOIDEA PAENUNGULATA | , Order SIRENIA Superorder i | MESAXONIA Order PERISSODACTYLA Superorder PARAXONIA Order ARTIODACTYLA Key for Identifying Orders of Mammals [т the Soviet Union] 2). РВ. 3(4). 4(3). 5 ( 6). 6 ( 5). 7 (10). 8 ( 9). 9( 8). 10 ( 7). 11 (14). 12 (13). 13 (12). 14 (11). 15 (16). Hind limbs absent. Body fishlike with large bilobate caudal fluke SCEMONIZOM alll yore soe heii. tare ae caer ee Mee ES CETACEA. Hind limbs present. Body not fishlike; tail, if present, not in the form of a bilobate fluke. Forelimbs in the form of leathery wings...... CHIROPTERA. Forelimbs of different structure. Fore- and hind limbs very short, paddle-shaped, and in the form of fins, 1.е., all digits right up to very tips enclosed in a common SHAM О roc ate ay species and subspecies). As a rule, the more thermophilic species use the land (eared seals, Otariidae; subtropical seals with 8 incisors, Monachinae; and also some Phocinae with 10 incisors—true seals, and the West European form of gray seal). The rest of the 10-incisored seals, the hooded seal, and antarctic 8-incisored seals usually reproduce and molt on ice. Some pagophilic species (ringed and Baikal seals, Caspian seal, Greenland seal, and others) make round openings in the ice cover for breathing and ventilation, and for crawling onto the ice. Some bur- row holes in the ice for themselves and for concealing their pups under the snow cover. There is no winter hibernation. Polygamy is common to the aegialoid [littoral; pagophobic] pin- nipeds. There is no strict monogamy since the males and females come together only for a short while, invariably preceded by fairly severe fights among the suitors. Males play no part in the care of the progeny. Fertility is low: all pinnipeds usually deliver a single pup; twins are extremely rare. The newborns are large, fully formed, covered with a thick coat, with normally developed limbs and open eyes. Postnatal growth 2 Ice-loving, according to the terminology of М.А. Smirnov (1914, 1927, and 1936). 3 Land-loving, ice-fearing species. 18 т is quite rapic in some (hooded seal, Greenland seal, and most other pagophilic seals) and less so among others (Baikal seal, ringed seal). In some others, growth is very slow (walrus, fur seal, sea lion). The feeding habits of pinnipeds are not clearly differentiated, though there are some mainly benthic feeders among them (walrus, Odobenus; bearded seal, Erignathus); some live on very large plankton, mostly fish and cephalopods (hooded seal; elephant seal; fur seals, Callorhinus and Arctocephalus;, sea lion, Zalophus), and some live on planktonic crustaceans (crabeater, Lobodon). Most survive on mixed food consisting mostly of small and minute fishes, large plankton, some demersal crustaceans, as well as cephalopods and other mollusks. The leopard seal, HAydrurga, feeds also on large fish and warm-blooded animals, including birds (penguins). Among the sense organs, the most developed are those of hearing and vision. The bony tympanic bullae are highly bulged in true seals (Pho- cidae) but flattened in all others. The pinnae are either altogether absent (true seals and walruses) or highly reduced (eared seals, Otariidae). Pinnipeds are distributed in all the oceans except the Indian Ocean; they are found in the southernmost fringe of the latter (generally not above 30° S lat.) from the adjoining antarctic waters (Fig. 5). An overwhelming majority of the species are confined to the regions of cold and moderately cold waters in which the surface temperature does not exceed 20°C at any time of the year (Davies, 1958b*). The exceptions are primarily thermophilic seals of the genus Monachus, with isolated distribution in three regions of the subtropical belt (one species in the Mediterranean Sea, in the open ocean around Gibraltar and Western Africa, and in the Black Sea; another in the Caribbean Sea‘; and a third in the Hawaiian Islands). Many others can withstand relatively warm water conditions: the southern population of the northern elephant seals (Mirounga in California), the California, and Galapagos sea lions (Zalo- phus), the South African Cape fur seal (Arctocephalus pusillus), the South Australian and New Zealand eared seals of the genera Neophoca, Pho- carctos, and Arctocephalus, some populations of the northern elephant seal (Mirounga angustirostris), and others. Pinnipeds are absent not only in the Indian Ocean but also in the Malayan archipelago, southwestern Pacific Ocean south of 30° М lat. to * Here and throughout the text an asterisk (*)-after a reference in the text indicates that either the author is not listed in the “Literature Cited” or the author is listed but no entry given for the publication date cited - General Editor. 4The Caribbean monk seal (Monachus tropicalis) is extinct; the few survivors of its populations were finally exterminated in the first half of this century. ‘(irysdeyD УМУ) Bag yoRTG 94) рие элэцазииэц илэЧ 1 ш Aepunog WIsyVION OOS = ОБ ae Om. Bat Oe EOS 110$ ЭЧ -elpodiuutg лор ЕЕ LF Е. | | sey Ее р = oa SNL 19 13 40° $ lat., and in its entire central part south and north of the equator up to the 30th parallel. The only exception is the small area around the Hawaiian Islands occupied by the relict small population of the Hawai- ian monk seal, Monachus schauinslandi. Pinnipeds are totally absent in the entire tropical portion of the Atlantic Ocean from 20° М lat. to 30°S lat. and some other pelagic parts of the world oceans, including a large expanse of temperate latitudes of the Atlantic to north of the equator (Figs. 5, 80). At present, pinnipeds are most numerous in the boreal and arctic regions of the Northern hemisphere and in the temperate zone of the Southern, especially in the zones of confluence of cold and warm waters (zone “front,” convergence). Some 14 species inhabit the seas of the Arctic Ocean, northern and subarctic parts of the Atlantic, and the northern parts of the Pacific Ocean (mostly true seals with 10 incisors, i.e., Phocidae, walrus, and some eared seals); just as many species inhabit the waters of the South- ern hemisphere (mostly eared and 8-incisored true seals). The remaining species are the above-mentioned subtropical monk seal and the inhabi- tants of landlocked waters such as the Caspian Sea and Lake Baikal. The species most adapted to living in icy conditions (especially the ringed seal and to a lesser extent the bearded seal) penetrate into very high latitudes, into the Atlantic sector of the Arctic, and reach north of the 80th parallel and even the polar regions in eastern Severnaya Zemlya. The zoogeographic range of pinnipeds is divided (Skleter,* 1897; and others) into five regions: 1) Arctoatlantic, including the Arctic Ocean and North Atlantic (endemic monotypic genera: hooded seals, Cystophora; gray seals, Halichoerus, to which may be added the Greenland seal); 2) Arctopacific, covering the extensive area of the North Pacific (endemic genera: northern fur seals, Callorhinus; sea lions, Eumetopias); 3) Mesat- lantic (subtropical belt intersecting the Atlantic Ocean on both sides of the Tropic of Cancer and including the Mediterranean and Black seas) —area of distribution of the Caribbean and Mediterranean monk seals; 4) Mesopacific—similar to the small zone around the Hawaiian Islands where the third member of the genus of monk seals, Monachus schauins- landi, lives; and 5) Notopelagic—the zone encircling the Antarctic and extending north roughly along the 20°C isotherm of February surface temperature of the sea on the coasts of Chile and Peru and also covering the coastal waters of South Africa, South Australia, and New Zealand (endemic genera: monotypic genera of the Antarctic, 8-incisored true seals—Phocidae; also the southern eared seals—Otariidae, especially the genera Otaria, Neophoca, Phocarctos, and many other species of southern fur seals of the genus Arctocephalus). 14 The general range of the order (Pinnipedia) did not undergo serious changes in the historic past, though in the eighteenth, nineteenth, and - twentieth centuries significant “gaps” have been noticed. Some popula- tions of the various species of southern fur seals (Arctocephalus) were totally exterminated and some of them, A. philippi (South America), almost completely. The Caribbean monk seal, M. tropicalis, has also become extinct in the present century while the Mediterranean species, M. monachus, is facing extinction. The populations of many species have shrunk sharply while those of many others continue to shrink in the North and Arctoatlantic (common seal, Phoca vitulina; walrus and, at places, the gray seal, Halichoerus grypus; to a lesser extent populations of ringed seal, Ph. hispida, and some others) as also in the northern part of the Pacific Ocean (sea lion, Zalophus californianus,; walrus); the population of the bearded seal, Erignathus barbatus, and of the ribbon seal, Histriophoca fasciata, has decreased sharply as a result of unrestrained hunting. Conservation measures and a rational system of utilization have helped restore the populations. The population of the northern fur seal (Callorhinus ursinus) and that of the following southern fur seals have already reached high levels: Arctocephalus pusillus (in South Africa), A. australis (at the coasts of Uruguay and Argentina), and the elephant seal (Mirounga leonina) (especially on the island of South Georgia). Popu- lations of the other southern fur seals and elephant seals as also the northern fur seal (at the Pacific coast of North America), which faced extinction, have begun to rise steadily. The total world population of all the species of pinnipeds is esti- mated at 16-20 million. The maximum numbers exceeding 1 million are attributed to four species: 1) the ашагсис crabeater seal, Lobodon car- cinophaga, probably numbering 4-5 million; 2) the ringed seal, Phoca hispida, somewhat less numerous than the preceding species (1-2 mil- lion); 3) the Greenland seal, Phoca groenlandica, has a total population of 1.5-2 million; and 4) the northern fur seal, Callorhinus ursinus, has a total population of 2-2.5 million. The census of the southern sea lion, Otaria byronia, is estimated at 800,000, that of the southern ele- phant seal at 600,000, and for the bearded seal (Erignathus barbatus), hooded seal (Cystophora cristata), Weddell’s seal (Leptonychotes wed- delli), South African fur seal (Arctocephalus pusillus), and Caspian seal (Phoca caspica)—400,000 - 500,000 each. The population of most other species is even less (Chapskii, 1966; and others). Phylogenetically, the Pinnipedia are very close to the Carnivora from whose more primitive ancestors they undoubtedly evolved comparatively recently (in the geological sense of time), but evidently not later than in 20 15 the Oligocene. In fact, members of this order belonging to the Oligocene have not yet been discovered. The oldest known fossils are from the Lower and Middle Miocene. These were fully formed members of the same families to which the present-day Pinnipedia belong. Thus, a mem- ber of the family of eared seals could be recognized from the lower jaw with teeth of the Lower Miocene seal, Allodesmus kernensis Kellogg. Sim- ilarly, the Middle Miocene genera Leptophoca, True, Miophoca, Zapfe, or the Upper Miocene genus Monotherium van Beneden belong to the fam- ily of true seals (Phocidae). At that time (if not earlier), the present-day subfamilies of Phocidae already existed. The Miocene seals did not differ considerably from the present-day species and morphologically did not stand very close to any of the land or semiaquatic carnivores which could serve as the ancestral form for the evolution of pinnipeds. All this offers a basis for presuming that the initiation of the primary phyletic branches leading to the families of sea lions and fur seals (Otariidae), the walrus (Odobenidae), and true seals (Phocidae) should belong to a much earlier period than the Miocene. Many cardinal differences in body structure between the eared (Otariidae) and earless (Phocidae) seals greatly complicate establishing a single root which could be regarded as the base for the evolution of this group of mammals. Among these characteristics are: features of the tarsal region (especially the structure of the ankle bone) and the resultant differences in the ability of the animals to move on land, difference in the general structure of flippers, numerous elements of craniological dissimilarity (structure of tympanic region, mastoid, its association with the paroccipital process, and presence or otherwise of the alisphenoid canal). Quite some time ago (Maivart, 1885*) attention was drawn to the fact that one group of pinnipeds (eared seals and the walrus, Otari-- Oidea) was significantly close to bears in many craniological features, while another group (true seals, Phocidae) reveals a similarity, though less distinctly, with martens. From this arose the concept of diphyletic origin (Kellogg, 1922; Howell, 1929,* 1930*; McLaren, 1960; Chapskii, 1963; King, 1964; and others). In spite of some features complicating the adoption of this hypothe- sis, especially that bears represent a much younger branch of carnivores whose evolution is put in a much later period than Pinnipedia, the pos- sible diphyletic origin of the latter is not excluded. The evolution of true seals (Phocidae) from martens is quite convincingly demonstrated by the find of a seallike otter—seal semantor, Semantor macrurus, in the Pliocene formations of Western Siberia (Orlov, 1931*). This animal possessed distinct transitional features of structure between the sea otter 21 16 and true seals. It was closer to the seal than the sea otter with respect to the humerus (Kirpichnikov, 1955), though some other skeletal elements have revealed, on the contrary, a very close genetic similarity with the sea otter (K.K. Chapskii). The historical evolution of true seals (Phocidae) evidently proceeded in a similar manner, but in a much earlier geologi- cal age. The semantor was a somewhat incomplete branch formed in the same direction, independent of the emerging seals, but under different conditions. This could serve as an argument in favor of the validity of different ways of formation of the different groups of Pinnipedia and, hence the possibility of interpreting the latter not as a phylogenetically valid order (or suborder) but as a composite group of animals arising from different roots.° The extremely scant paleontological material on true seals in general, and especially the recent finds within the USSR, make it difficult to estimate the exact number of not only species, but even of genera of fossil Phocidae. At present, 42 genera of Pinnipedia are known. Of these, 25 are fossil (Simpson, 1945; McLaren, 1960; Mitchell, 1961,* 1968*) and 17 Recent. This proportion does not, however, indicate that Pinnipedia are at the end of their evolutionary development. The Recent genera form a fairly large figure of 40%, which is much higher than the corresponding figure (28%) for the “contemporaneity” of land carnivores (Heptner er al., 1967*). The other indices from which the potentially developing state of the order could be judged are seen from the areas of distribution, population, and biological stability of the different species. The range of the common seal (Phoca vitulina) is very extensive and is interrupted only by ecological barriers; the range of the bearded seal (Erignathus barbatus), the ringed seal (Ph. hispida), and the genus of southern fur seals (Arctocephalus) is fairly large though not continuous; the range of the northern fur seal (Callorhinus ursinus) and the Greenland seal (P. groenlandica) is also fairly large. The relative diversity of forms in spite of the narrow specialization of the order as a whole is per se significant. The high population of many species (see above), including even such relict species as the Caspian seal existing under conditions of intense anthropogenic pressure, is an impor- tant feature. In spite of their low fertility, pinnipeds can quite quickly 5 The origin of a group or two highly proximate families of the same order (suborder) could hardly be regarded as polyphyly. This, at best, is paraphyly or a form of monophyly. (V.H.) 17 restore their populations and thus largely withstand destructive hunt- ing. They cannot, however, survive incessant killing. Thus, one species (Caribbean monk seal, Monachus tropicalis) became totally extinct, while the population of another (Mediterranean monk seal, M. monachus) is vanishing right before our eyes; the population of yet another (Hawai- ian monk seal, M. schauinslandi) is very small and only exists because of protective measures. Some other populations, especially of antarctic species, are in a pitiable state as a result of irrational killing. The popu- lation of all species of pinnipeds is wholly dependent on their judicious utilization and conservation. The classification of the Pinnipedia is quite simple in its general fea- tures and more or less generally accepted. The differences arise only in the number and scope of systematic categories and the position. of the entire group which is interpreted sometimes as an independent order or suborder of carnivores (Carnivora), and sometimes even as a com- posite group deserving a rank of several families (or subfamilies) in the superfamily Canoidea of carnivores (Carnivora). Moreover, none of the researchers inclined to such an interpretation (Frian, 1956*; Lien and Waiens, 1956*) have been able to specifically accomplish such an arrangement. The Pinnipedia, regarded as an independent order, are divided into two superfamilies: Otarioidea and Phocoidea. The former consists of two families: eared seals, Otariidae (Steller’s sea lion, sea lions, and fur seals) and walruses, Odobenidae. The second superfam- ily comprises only a single family—true seals, Phocidae.® Quite often (N.A. Smirnov, 1929, 1935; and others), only two families are recog- nized: Otariidae s. lato, which includes not only eared seals, but also walruses, and Phocidae, true seals. Otarioidea are characterized by the following features: the bullae osseae are flattened and have complex angular outlines; the mastoid forms a single extremely massive process in adults; the process descends much below the tympanum part and fuses with the pr. paroccipitalis; the alisphenoid canal is present. The astragalus resembles more that of Carnivora than of members of Phocoidea. The hind limbs can bend forward and are adapted to locomotion on land. Phocoidea are characterized by the following features: the tympanic bullae are fairly strongly bulged and rounded; the mastoid process is not joined with the pr. paroccipitalis (when present), not bent downward, and generally not well developed. The alisphenoid canal is absent. The astragalus has a large process at the top (no similarity whatsoever with 6 The fourth family, Semantoridae (fossils of seallike otter—semantors; Yu.A. Orlov, 1931), sometimes placed in the Pinnipedia (Simpson, 1945), should not be included here. 22 18 that in land carnivores) and its articular surface is not saddle-shaped. The hind limbs do not bend forward and do not participate at all in locomotion on land. There is no basis for assuming that new species not known so far will be detected in the order.’ At the same time, the morphological features of the subspecies of many true seals (Phocidae) should be reviewed. The scope of some genera of this family should also be partly reviewed. How- ever, there is hardly any need to revise the classical division of phocids into subfamilies, as recommended by King (1966). In the family of eared seals, Otariidae, there is evidently need for a more rational argument for the independence of all the species of the genus Arctocephalus. Thus the system of eared seals of the Southern hemisphere at the level of genera and species cannot yet be regarded as conclusively estab- lished. There is no single opinion even about the scope of the genus of some northern seals of the subfamily Phocinae in view of the attempt of some theriologists to give a broader interpretation to the concept of genus. The antarctic 8-incisored seals can be regarded as conclu- sively established monotypic genera. The morpho-ecological rationale of changing the measurements of taxonomic differences prevailing among seals of the subgenera of the widely interpreted genus Phoca (true seals) is highly substantiated. At present, 32 -34 present-day species are included in the order Pin- nipedia, i.e., only some 1% of the total number of species of mammals of the world. On average, a genus has 2.1 species. There are a maxi- mum of 18-19 species in the family of true seals, Phocidae; eared seals, Otariidae, have 12-13 while there is only one species in the family of walruses, Odobenidae. The present-day pinniped fauna of the world is represented almost evenly in the North Atlantic basin (10 species includ- ing the Caspian seal), in the North Pacific Ocean (11-12 species) and in the seas of the Southern hemisphere (13 species). Two superfamilies, all the 3 families, and 8 genera, 1.е., 42% of the 19 present-day genera, are known in the seas and landlocked water bodies of the USSR. However, the fauna of the USSR comprises only 13 species,® 1.е., about 39% of all the species known in the world fauna. 7 The unexpected recent “discovery” in the Pacific Ocean of a “new” species, described under the name of the island seal Phoca insularis (Belkin, 1964), was actually one of the forms of the pagophobic common seal, Phoca vitulina (see p. 314, and also Chapskii, 1967, 1969). 8 If the common seal (Phoca vitulina) and the subspecies are regarded as a single species; see pp. 323 - 330. 23 19 The range of the order encompasses all the oceans of the USSR and some of the largest inland water bodies (Caspian Sea, Lake Baikal, Lake Ladoga). The practical importance of pinnipeds is very high. Almost all the species, but primarily those which periodically form massive herds, rep- resent game animals of great economic value and are hunted mainly for their fur and partly for their skin; the fat, meat, and some other body parts are also used. Historically, the hunting of pinnipeds has played a major role. Seal- ing vessels combed the seas of both hemispheres in search of herds of these animals, especially fur seals, and killed them mercilessly. This car- nage continued for decades and ultimately led to the near total depletion of stocks of the commercially valuable species. In the Southern hemi- sphere, all the rookeries of the southern fur seal were almost wholly rav- aged and the elephant seal population suffered. The latter faced extinc- tion even at very low latitudes. The rookeries of the northern fur seal on the Commander and Pribilov islands were subject to intense destruction. Effective conservation measures alone prevented the recurrence of the fate that befell many populations of the southern fur seal. The stocks of walrus were severely depleted, especially in the Atlantic sector of the Arctic, and also in other regions of its habitat. In spite of its sordid history, sealing at sea has not lost its importance to date. In some regions of the globe it continues to play an important economic role as a supplier of raw material for the fur, hide, and other industries. In some countries (such as Greenland), sealing is extremely important and represents the only source of livelihood for northern native populations. Hunting of pinnipeds is of great significance in the life of the native coastal populations of Alaska and the Chukchi Penin- sula, for the coastal villagers of the Soviet Arkhangel’sk region, and for hunters of some other regions of our country. It provides Eskimos and some coastal Chukchians meat for themselves and for their dogs, skin for making footwear, harnesses, and other requirements, as well as fat, etc. Moreover, it provides work for the people engaged at the collective farms and in the state-owned sealing industry. In the USSR and many foreign countries, seal skins are in demand as highly valuable, durable, and fashionable furs. The most important targets of sealing are the northern and southern fur seals, the Green- land seal, the ringed seal, the Caspian seal, the hooded seal, etc. Fur is generally obtained from fur seals and mainly young Greenland and Caspian seals, hooded seals, etc., whjle the skin is obtained from large animals. Blubber is mainly used commercially in the tanning industry, 24 20 for soap-making, etc., for making medicinal (“fish”) ой, and as an ingre- dient in other products. Meat is used locally. Some pinnipeds (e.g., larga seals) inflict much damage in fisheries by thriving on prime fish, espe- cially salmon. The most important commercial species of seals are now exploited almost everywhere on a rational basis, eliminating the danger of their depletion. (K. Ch.) 1 (4). 2 (3). 3 (2). 4 (1). 1 (4). 2 (3). Key to Families of Pinnipedia Identification Based on External Features Hind flippers bent at calcaneal joint with foot forward, serving as body support on hard ground. Fore flippers longer than hind ones, naked distally; claws on hands absent or poorly developed and placed well behind margin. Snout narrowed anteriorly and quite elongated; pinnae small. Hands long, roughly triangular, wing-shaped. Claws absent (or very small). Well-developed claws only on three middle digits of hind flippers. Upper canines (in adults) not protruding from СОЗ MOUS oe ogee yee yoke ops Eared seals, Otariidae (p. 58) Snout very broad, anteriorly somewhat truncated and short. Traces of pinnae not seen. Hands not highly elongated and not triangular; small reduced claws seen on digits of hands and all digits of feet. Upper canines notably protruding from closed mouth. ......... FUER A RAE APE BES RII oR OA Walruses, Odobenidae (p. 22) Hind flippers not bent forward at calcaneal joint, always held backward, and not serving as body support on solid ground. Fore flippers not long, slightly shorter than hind ones, and fully cov- ered with fur. Hand with well-developed claws at end of digits. AUT LONI Coa eee eer ne Sa Barada True seals, Phocidae (p. 142) Identification Based on Skull Features Tympanic bullae relatively small, flattened from top downward, with uneven rugose surface, and complex outline. Alisphenoid canal (Fig. 4) present. Mastoid part of temporals with very mas- sive protuberance directed downward. All or almost all cheek teeth with single root (the last with two roots) and with simple undivided crown. Rostral portion [of skull] slightly narrower than cranial part in tem- poral region. Ends of middle pair of upper incisors with transverse angular notch (Fig. 6). Nasals shifted backward and projection of frontals lodged in fork between their apices (Fig. 7). Supraorbital processes well developed. Lower jaw teeth sharply differentiated 25 24 24 3 (2). 4 (1). 21 into incisors, canines, and cheek teeth (premolars and molars similar). Upper canines similar to lower ones in size and structure. payne ча рН. she ha Maes Eared seals, Otariidae (p. 58) Rostral portion [of skull] hardly less wide than cranial portion in temporal region. Ends of nasals posteriorly, at junction with frontals, sharply incised transversely, not forming acute apices (Fig. 7). Supraorbital processes not developed. Lower jaw teeth similar to one another; upper canines large and may attain large size, bearing no similarity to the lower ones, which do not differ fromycheek teeth: ...‹........... Walruses, Odobenidae (p. 22) Tympanic bullae relatively large, bulging, with almost smooth semicircular surface and comparatively simple outline. Alisphe- noid canal absent. Mastoid part of temporals without mas- sive downwardly directed protuberance. Most cheek teeth with two roots and with divided crown. ......................0 000 Fig. 6. Transverse notch in middle upper incisor of eared seals, Otariidae, and the fur seal, Callorhinus ursinus (figure by К.К. Chapskii). Fig. 7. Structure of nasals in pinnipeds of various families: A—eared seals (family Otariidae); B—walruses (family Odobenidae); C—true seals (family Phocidae): 1—frontal, 2—maxillary, 3—premaxillary, 4—nasal bones. 22 25 Family of Walruses Family ODOBENIDAE Allen, 1880 This is one of the largest groups of animals of the order Pinnipedia. The trunk is massive; the head is rounded and, compared with the massive body, appears small. Pinnae are absent. The whiskers are long, dense, 24 Fig. 8. Left zygomatic bone in pinnipeds: A—eared seal (fur seal, Callorhinus ursinus); B—walrus (Odobenus rosmarus), C—bearded seal (EFrignathus barbatus); D—common seal (Phoca vitulina) (figures by K.K. Chapskii): 1—anterior lower corner (or process). 25 Fig. 9. Skeleton of a seal (figure by N.S. Kondakov). 23 very thick, and directed downward. The neck is short but movable and merges imperceptibly into a fairly clumsy trunk. The skin is thick, rough, and forms numerous folds and wrinkles. The hair coat is sparse and bristly; large portions of the body are naked in adults. The fore flippers are slightly larger than the hind ones, terminating in a frill of skin devoid of hair; the frill extends beyond the margin of the digital phalanges. The first digit (inner) is the longest and the rest shorter in the order of first to fifth. The hind flippers have a naked foot; the fifth digit is the longest; the first is almost equal to it. The claws on the flippers are weakly developed and disposed away from the margin of the flippers. The hind flippers can bend under the trunk and assist in movement on land (Fig. 10). There is one pair of teats. The testes are disposed under a layer of skin and fat (subintegumental); a scrotum is absent. The skull is massive and broad with an extremely massive elevated frontal portion, by which it differs from the skulls of all other Pinni- pedia (Fig. 11). The skull bones are highly massive. The width of the skull above the canines is almost equal to the width above the external auditory meatus. A supraorbital process is absent. The hind margin of the nasals forms an almost straight line. The orbits are relatively small, their transverse width equal to 1/3 or 1/2 the width of the palate in the line of molars. The bony palate is flexed inward like a boat, forming at the back a very gentle arc turned forward with a bulge. The exter- nal auditory meatuses are very small and their lower walls thickened. Z Ye y oe Fig. 10. Fore and hind limbs of the walrus (figure by N.N. Kondakov). 27 24 Fig. 11. Skull of ап adult walrus, Odobenus rosmarus (figure by М.М. Kondakov). The tympanic bullae are relatively small and flattened. The lower lat- eral angle of the temporal bone bears a massive projection. This bony mass exceeds the height of the tympanic bulla by.a few times. The ante- rior section of the lower jaw is very massive and its two halves are firmly fused. The dental formula of permanent teeth in most walruses is: 2—2 1-1 1-1 3—3 ол. oS The front incisors are small or altogether reduced, while the last pair of incisors is indistinguishable from the molars in size and shape. The upper canines are massive, up to 80 cm long, and directed vertically downward. The molars are massive, with a single root, their cusps sloping backward. In older animals the molars become worn and flat, and even have a slightly concave surface. The scapula is relatively long, without a perceptible arcuate notch on the hind margin. Its crest is comparatively elongated, reaching the hind margin. The os penis is very long, 508-512 mm, slightly S-shaped, its posterior end broadened and terminating in a head; the anterior end is obliquely truncated. Paired air sacs are formed by a projection of the upper section of the esophagus and the broad openings joined with it. There are no closing valves. Each sac can hold up to 50 liters of water [air]. The 26 25 И Ум Wi LENS Wb Fig. 12. Skull of a yearling walrus, Odobenus rosmarus (figure by N.N. Kondakov). air-filled sacs inflate and, spreading under the skin of the neck, hold it up; their ends lie between the scapulae (Sleptsov, 1940; Nikulin, 1941; Fay, 1960*). _ Walruses mainly inhabit the coastal shallow waters of arctic seas and feed on benthic invertebrates. A large part of their life is associated with ice floes. Walruses are distributed only in the Northern hemisphere, in the circumpolar region, with small interruptions. They inhabit the Bering, Chukchi, East Siberian, Laptev, Kara, and Barents seas and the waters of the Canadian archipelago in the northwestern part of the North Atlantic Ocean. Walruses constitute one of the three families of the order Pinni- pedia, but the taxonomic position of the family has not yet been clearly established. Some authors regard it as a subfamily of the family Otariidae (Smirnov, 1935; Romer, 1939; and others), while many separate it into an independent family. The latter view is more prevalent. The relative proximity of the family of walruses to eared seals (Otariidae) is, however, beyond doubt. This fact is particularly emphasized by combining them into one superfamily, Otarioidea Smirnov, and contrasting them in such a combined form with the superfamily Phocoidea Smirnov, comprising only the family of true seals. 26 In origin, the walrus family is closely related to the family of eared seals (Otariidae) and could be regarded as its derivative. Thus the ear- liest and most primitive known form of the walrus family, Prorosmarus alleni, from the Upper Miocene of the Atlantic coast of North America bears some features of the skull structure and dentition characteristic of Otariidae. In particular, the lower canines are preserved in this form (Tenius and Gofer, 1960*). Of the four genera of the family, only one is Recent. Apart from Prorosmarus, two other genera, i.e., Trichechodon and Alachtherium, are known from the Middle Pliocene and Pleistocene of Europe. The only present-day genus, Odobenus, is known from the Pleistocene of North America and Europe. Extinct as well as extant genera are known only from the arctic seas of the European and American continents, i.e., the present-day range of the family. The North Atlantic Ocean could perhaps be regarded as the center of origin of this family. The economic importance of this family is presently low, since the walrus population has been greatly depleted in recent decades. The family is represented by one genus, Odobenus Brisson, 1762, with a single species, O. rosmarus Linnaeus, 1758, widely distributed in the waters of the USSR. (V.A.) Genus of Walruses Genus Qdobenus Brisson, 1762 1762. Odobenus. Brisson. Regnum animale. Ed. 2, p. 30. Phoca rosmarus ) Linnaeus. 1766. Trichechus. Linnaeus. Syst. Nat., ed. XII, I, p. 49. Nec Linnaeus 1758 (pertains to manatee Trichechus manatus Linnaeus, 1758). 1772. Rosmarus. Brinnich. Zoologiae fundamenta, р. 34. Phoca ros- marus Linnaeus, 1758. (V.H.) See description of the family. WALRUS Odobenus rosmarus (Linnaeus, 1758) 1758. Phoca rosmarus. Linnaeus. Syst. nat. Ed. X, I, p. 38, North Atlantic Ocean. 1811. Trichechus arcticus. Pallas. Zoogr. rosso-asiatica, I, p. 269, Novaya Zemlya (“Frequens in Oceano arctico ... Copiosissimi in Insula Navaja Zemla.” (V.H.) 29 27 1815. Trichechus divergens. Illiger. АБВ. Acad. Wiss. Berlin, 1804—II, р: 68. 35 miles south of Ici Cape, Alaska (162° W long. and 70° М lat.), Chukchi Sea. (V.H.) 1815. Trichechus obesus. Illiger. Ibid., p. 64, Nom. nud. 1831. Trichechus сооки. Fremery. Bijdrag. Nat. Vetensk, 6, р. 385. Ici Cape zone in Alaska (Chukchi Sea, 70° М lat. and 163° 18’ W long.). | 1922. Trichechus orientalis. Dybowski. Arch. Tow. Nauk. Lwow.., I, р. 351, Nom. nud. 1940. Odobenus rosmarus laptevi. Chapskij. “Problemy Arktiki” (Prob- lems of the Arctic), No. 6, p. 94. Laptev Sea. (V.H.) Diagnosis Only species of the genus. Description In general form, the walrus differs considerably from all other species of Pinnipedia (Fig. 13). The body is large and massive, the skin thick, covered with wrinkles and folds, and the limbs broad (Fig. 13). Movement on land is slow and cumbersome and the animal utilizes all four limbs, but in water it is quite agile and moves fairly fast. It crawls onto ice floes with difficulty, using its tusks and fore flippers. The head is relatively small with a massive snout that is blunt in front. The eyes are small and shifted far back (Fig. 14). Long (up to 10-12 cm), numerous, very hard and thick whiskers occur on the front part of the snout; they are directed downward and arranged in 13-14 rows. Each side of the snout bears 300-350 whiskers. The whiskers in the middle part of the snout are usually very worn and measure hardly 0.5 to 1 cm. Both tusks in the upper jaw are very long and directed vertically downward. Instances are known of walruses with more than one pair of tusks. A walrus caught around 1915 had two well-developed tusks on each side, and all four were of almost normal length. Those on the right side grew parallel to each other while those on the left were twisted (Caldwell, 1964). Four skulls have been described with three tusks and five with five tusks each. A skull was found in which there was a single normally developed tusk on the left and five separate formations on the right side consisting of fragments of small tusks of normal shape, two dentine stubs 12 and 14 cm long rising from the surface of the gums, and two dentine concretions 12 and 20 mm in diameter concealed in the jawbone (Bel’kovich and Yablokov, 1960). 29 Fig. 14. Front view of a walrus head (figure by М.М. Kondakov). 29 The color of old walruses is a dirty olive on the back and rusty-brown on the belly. This coloration is caused by the color of the hair, as also the skin pigmentation, which is brownish. The tips of the flippers, devoid of hair, are similar in coloration. Sexual and age-related dimorphism are manifest in the body dimen- sions, shape and size of tusks, and hair coat. Adult males are about 0.5 m longer than females; the male skull is more massive and the tusks longer and thicker than in the female. The tips of the tusks diverge sideways in the male but in the female are somewhat proximate with a slightly spiral curvature. The female, usually darker, attains maturity one to two years earlier than the male. Pups of both sexes are identical in size, but the female lags behind in growth during the period of sexual maturity and ceases to grow altogether soon thereafter; cessation of growth sets in later in the male. A clear example of sexual dimorphism is the presence of large wartlike formations on the breast and shoulders of males, which are absent in mature females. The hair coat of young walruses is fairly dense and dark brown. It becomes bristly, sparse, and brownish-yellow with age, with large bald patches. Evidently the hair color undergoes no seasonal variations. (For skull description, see under characteristics of the family; body and skull sizes are given under “Geographic Variation”.) The weight of an adult male can reach almost 1,500 kg and that of an adult female 800-900 kg. The average weight of the visceral organs (Seven specimens) is: heart 6,167 g, lungs 14,062 g, liver 29,640 g, spleen 4,146 g, stomach 5,312 g, intestine 33,640 р, kidneys 3,544 р, and pancreas 2,148 р. (V.A.) Geographic Distribution Arctic seas of the Atlantic and Pacific Oceans. Geographic Range in the USSR (Reconstructed) In the early twentieth century, walruses were probably regular inhab- itants of the northern White Sea (Morzhovets Island). In the Bering Sea during the nineteenth century, walruses bred regularly on Karaginsk Island (59° М lat.) and in the 1880s were caught in thousands every year. Walruses were reported on Cape Kronotskii (56° М lat.), Cape Shipunskii (53° N lat.), and on the Commander Islands (Grebnitskii, 1902; Suvorov, 1914; Arsen’ev, 1928*; Ognev, 1935; N. Smirnov, 1935; Nikulin, 1941) (Fig. 15). Direct references are available to the breeding of walruses in the northern part of the Sea of Okhotsk. The reports of a Yakutian army commander in 1651 refer to the hunting possibilities of walruses in the Зоиихо MOU SI зпем э41 YOY ит $10131 Dy) $9007 рэцозец pue э8ие1 рэ12пл11510991 ay) Jo Азерипод ay) зэюцэр aut] рноб -(aa,uasTy 7.) 31550] Э41 Ul $тириибол пиздоро ‘зпцем эчз JO чоппана ‘ст i) fe Te ; 7 et 06 OL =F у, [ el В. Sey RS ay №. а | ut \ Ps | О =} у re) д д К {3 LB) eS “a 75% a eo & é is Bet Fy >, к $ ==) WZ а к к К re 31 Sea of Okhotsk. In the record for 1652, he reported sighting many walruses on the beach of Cape Morzhov for two or more “versts” [1 verst = 1.067 km] (Akinfov, 1848). These data pertain to the northern part of the Sea of Okhotsk not far from the present Magadan. (V.A.) The veracity of this report was confirmed by a reference to the possibility of finding “fish teeth,” i.e., walrus tusks, at these places. This is also supported by the find of tusks in the coastal rock mounds along the northern coast of the Sea of Okhotsk, and the sighting of live walruses. A young walrus was found around 1890 on Yamsk Island in the northeastern part of the Sea of Okhotsk and a pair of tusks was found around 1900 on the coastal dumps of Shelikhov Strait. Walrus tusks were also found in Nogaev Bay (B.A. Zenkovich). The present-day distribution of walruses in the USSR forms but a small part of the range which prevailed in the past. Walruses inhabit the waters of Franz Josef Land, Novaya Zemlya, Barents and Kara seas, Severnaya Zemlya islands (more often on the eastern coast of the archipelago), Vil’kitsk Strait, and in the shallow waters of the Ob’-Yenisey. They inhabit the Laptev Sea (mostly its western part close to the eastern Taimyr coast), Lena delta, landlocked waters of the Novosibirsk archipelago (mostly its northwestern fringe), and are encountered in the western part of the East Siberian Sea, mainly in the region of Novosibirsk Islands and De Long Islands. Farther east, they are found in the Chukchi Sea from De Long Strait in the west to Wrangel Island in the north, Bering Strait and Anadyr Strait to Cape Navarin (Chapskii, 1936, 1939, 1940, 1941, 1963; Tsalkin, 1937; Belopol’skii, 1939; L.N. Popov, 1939; Rutilevskii, 1939; Nikulin, 1941; Vinogradov, 1949; Zakharov, 1958; L.A. Popov, 1958, 1959, 1960). Geographic Range outside the USSR (Reconstructed) The southern boundary of walrus distribution in the Atlantic Ocean out- side the waters of the USSR has also varied considerably. Judging from the finds of walrus remains in excavations, this animal penetrated far southward at one time. In the nineteenth century, its remains were found in Denmark, England, France, and on the east coast of North America in New Jersey, Virginia, and Carolina (Moor, 1952*). Instances are known of the discovery of walrus remains in Maine and Massachusetts. More- over, fragments of the skull and other bones were found in the Gulf of Maine (G.M. Allen, 1930; Palmer, 1944). In the first half of the sev- enteenth century, walrus hunting prevailed on Sable Island (44° N lat.), close to the Canadian coast north of the Gulf of Maine (G.M. Allen, 1930) and on the Magdalen Islands (Mansfield, 1959). 33 32 In the middle of the nineteenth century, stray walruses were caught on the Shetland and Orkney islands (Moor, 1952*). At the end of the nineteenth century, walruses were perhaps permanent inhabitants on the coast of Finmarken in northern Norway, found on the coasts of Scotland, and in the Gulf of St. Lawrence. In general, at the end of the last century, walruses were widely distributed in the northern part of the Atlantic Ocean and in the Arctic Sea. It has been assumed that in the eastern part of the Pacific Ocean, walruses penetrated south of the Aleutian Islands, reaching Shumagin Island and even the Alexander archipelago in the eastern part of the Gulf of Alaska. _ In the middle of this century, walruses were known to inhabit the northern part of the Atlantic Ocean, being recorded in the waters of the Canadian archipelago (Southampton and Devon islands, Fox Basin, and Baffin Bay), the Labrador coast, Davis Strait, the west and east coasts of Greenland, Spitsbergen, and were sometimes spotted near Iceland (Fig. 16). In the Pacific Ocean, walruses inhabit the eastern part of the Bering Sea, from Bristol Bay in the south to the Bering Strait in the north (includ- ing the St. Lawrence Islands, Nunivak, and St. Matthew), and extend along the north coast of Alaska into the Chukchi Sea up to Cape Barrow. The northern boundary of the range of walruses in the Barents and Kara seas runs beyond 80° N lat. In April, 1957, a walrus was noticed in the breeding grounds on Franz Josef Land north of 81° N lat. (Vaigachev, 1958). In the waters of the Pacific Basin, the area 72 to 74° N lat. can be regarded as the regular northern boundary of the range of walruses, but the animals can be found even more northward in favorable years, depending on the situation of the ice floes. (V.A.) Geographic Variation Usually, three subspecies are recognized. All of them inhabit the waters of the USSR. 1. Atlantic walrus (O. r. rosmarus (Linnaeus, 1758)) (syn. arcticus, ? obe- sus). This is the smallest form. The maximum body length of males is 375 cm, of females 338 cm; the corresponding averages are 345 and 293 cm respectively (Chapskii, 1963). The condylobasal length of skull in males is 256-379 mm @ = 369), in females 303-342 mm (х = 314); the maximum width of skull in males is 268-291 mm, in females 222-257 mm (х = 234.2). The length of male tusks along the curvature from the edge of the alveolus to the 33 $11550} зпем JO SPUlJ JO $215—ф ‘5э5ПИЕМ Jo $риц Аер-1и2$5924 Jo $213—$ Чоицхо MOU ‘эле зэ5пем ЭША Ul $401891—2 ‘э8иел рэ12п11$1099эл1 Jo Алерипоа— т ‘(лэ.иэзгу “W'A) Smipuisos хпизаоро ‘зпцем ay} JO понпанзта ‘9т “314 С5 34 34 tip is 34-38 cm, in females 27-33 cm (Ognev, 1935). The tusk tenet in one male was 52.5 cm (Chapskii, 1963). In the USSR it inhabits the Barents and Kara seas; outside the USSR, the North Atlantic Ocean to the Canadian archipelago in the west, inclusive. 2. Laptev walrus (O. r. laptevi Chapsky, 1940). Somewhat larger than the Atlantic form. The maximum body length of males is 410 cm, of females 370 cm; maximum length of the tusk is 65 cm in males, in females 58 cm. The maximum circumference of the tusk in males is 21 cm, in females 14 cm (L.A. Popov, 1960). This form inhabits the Laptev Sea, the western part of the East Siberian Sea, the Lena Delta, and the Novosibirsk archipelago. It is more numerous near the coast of eastern Taimyr, where it is mainly confined to the coastal region and the shallow waters; it is rare in the western part of the East Siberian Sea in the region bordering the Laptev Sea. Not found outside the USSR waters. 3. Pacific walrus (О. г. divergens (Illiger, 1815)) (syn. cookii, ? orientalis). Largest form of the species. The maximum body length of males is 450 cm (х = 336), of females 367 cm (х = 283). The skull is more massive than in other forms and the frontal section considerably broader. The condylobasal length of the skull in males is 383-428 mm (x = 396.8), in females 315-357 mm (x = 332); maximum width of the skull in males is 290-333 mm (х = 309), in females 219-265 mm (x = 245). The length of the tusk in males is 46-80 cm, in females 40-60 cm; its width in males is 66-84 mm, in females 41-53 mm (Ognev, 1935; Nikulin, 1941; Freiman, 1941). It inhabits the Bering, Chukchi, and eastern part of the East Siberian seas. Some contacts between the Pacific, Laptev, and East Siberian Sea walruses are possible. Outside the USSR, it is found in American waters of the Bering and Chukchi seas. (V.A.) Biology Population. As a result of prolonged hunting, the walrus population has declined steeply throughout its range. The population of the Atlantic walrus has suffered the most and only a few stray herds are now known. The walruses of the Kara Sea suffered the highest destruction and those remaining now do not exceed a few thousand. Small groups are confined to Franz Josef Land 35 34 35 and Spitsbergen. The population in the eastern part of the Canadian archipelago is split into a few groups. In the northern part of Hudson Bay and in the region of Southampton and Cox, some 3,000 are known (Mansfield, 1959). Population figures for other regions are not available. The population of the Laptev Sea walruses has been less affected by exploitation, as it has always been relatively small. Nevertheless, hunting has had its impact here as well, and the present population barely exceeds 5,000 - 6,000. The Pacific walrus populations are best protected. Aerovisual estimates, aerial photographs, and studies of coastal breeding grounds and rookeries on the icy coasts revealed a summer population of 30,000 - 35,000 in the USSR. According to American and Canadian authorities, some 15,000 walruses inhabit the US waters in summer. The total population thus is 50,000 (Fay, 1957; Fedoseev, 1962; Krylov, 1968; Gol’tsev, 1968). Habitat. An outstanding feature of the habitat is the relatively shal- low water and abundance of benthic mollusks and partly of crustaceans. Drifting ice is common in these sections and, in the winter months, wal- ruses inhabit only the ice (Fig. 18). In the absence of ice in summer, walruses form coastal rookeries mainly on sandy or pebbly shoals on the coasts of the continent or islands. Food. There is no information on the winter food of walruses, nor on seasonal food variations. Our knowledge is limited to a list of animals retrieved from the stomach of walruses in the summer months. Fig. 17. A group of walruses оп an ice floe. Chukchi Sea (photograph by V.M. Bel’kovich). 36 Fig. 18. Adult females and young walruses (left). Chukchi Sea (photograph by V.I. Krylov). In the first two years juveniles survive on the mother’s milk and take to independent feeding only in the third year. By then, the tusks have grown sufficiently large to enable the walrus to independently scrape food from the bottom of the sea. The main food items of adults comprise bottom-dwelling invertebrates (Table 1). Throughout its extensive range, the walrus primarily feeds on various mollusks, which in the species composition of its food occupy first place. These are followed by crustaceans, of almost equal dietary importance. All the other food items, i.e., worms, echinoderms, ascidians, and fish, can be regarded as secondary. In the stomach of an Atlantic walrus, over a hundred polar cod were found (Chapskii, 1936); fish remnants were few in the stomach of Pacific walruses (V.I. Krylov); only remnants resem- bling otoliths were found in the stomach of walruses in the Canadian archipelago (Mansfield, 1958). In addition to the food items listed, the stomach of one walrus (site of find not mentioned) contained Tridacna (Moor, 1952*). Remnants of pinnipeds and even whales were found in the stomach of some walruses in all the regions studied. | The walrus seeks its food from the sea floor. It is assumed that it digs the bottom with its tusks, selects mollusks, breaks the shells, and eats the molluskan bodies. 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The depth to which the walrus dives while feeding can be as much as 180 m (Mansfield, 1959). Some walruses prey on seals and, occasionally, on birds. Seals usu- ally avoid the area inhabited by a walrus-predator, but will return there if the predator is killed (V.A. Arsen’ev, 1935). Two categories of preda- tory walruses are known: the first group feeds on uncommon foods occa- sionally when invertebrates are not available for some reason, while the second category comprises regular predators. The latter have long, thin, sharp-tipped tusks; these are lone males (Fay, 1955*). Home range. There are no separate sections for individual walruses or even for groups of them. In the absence of ice in summer, walruses form coastal rookeries; however, it is still not clear whether the same animals return to the same sections. At the rookeries in the Laptev Sea, the same walruses evidently return to their own sections (L.A. Popov, 1958). Daily activity and behavior. No systematic diurnal activity has been established among walruses. The Atlantic walruses of all ages and sexes gather on the beaches _in August and September since there are almost no ice floes in the USSR waters in the regions of their summer habitat. There they remain, sometimes for 1.5 months, often going into the sea to feed. The animals driven away from the shore by stormy waves return to the beach as soon as the weather clears. Fig. 19. A male walrus with a broken tusk. Chukchi Peninsula (photograph by V.I. Krylov). 39 37 40 The Laptev walruses also form mixed coastal rookeries in which females with pups (Fig. 20) and juveniles live along with the large males. They lie in a definite sequence—females with pups close to the water, juveniles and the remaining adults farther away. They rest crowded together compactly in various postures, but mostly on the side. Yearlings and juveniles sometimes lie atop the adults. Fights are quite common between large bulls, who put their powerful tusks to good use. At all times, some animals are moving toward the water while some are returning to the rookery; thus the rookery is never quiescent (L.A. Popov, 1958). In September and October, the coastal herds disperse and the coast remains empty until the next season. The Pacific walruses spend much of their time on ice floes. Two types of colonies are distinguishable. One consists exclusively of adult males and the other of females with pups, among which a small number of males are sometimes observed. In calm weather walruses sleep soundly. The resting animal does not even deign to look at a ship approaching the ice floe and, when disturbed, takes to water rather reluctantly. Walrus herds have often been observed resting on ice floes within 100 т of freshly killed animals. They did not react even to gunshots, the clatter of operating winches on a ship, nor to any other noise. Yet, in general, their hearing is better than their sight. Fig. 20. Suckling walrus pup on an ice floe. Chukchi Sea (photograph by V.I. Krylov). 41 Exclusively male haunts are usually small but mixed herds run into several hundreds. The rookeries are mostly organized along the edges of drifting ice floes. The animals rest on permanent floes, whether smooth or hummocky. Frightened animals literally dive into the water, but soon surface, gather in herds, and begin groaning loudly. The herding instinct is strong among walruses (Dunbar, 1955). An injured animal is helped by others to float on the water surface. Pups that tire of swimming, often climb atop their mothers or any other walrus. In autumn the Pacific walruses form coastal rookeries in some other regions as well (Fig. 21); these may be permanent or temporary. The latter can be classified as regular or occasional. Only two permanent rookeries are preserved at present: at Rudder and Meechken in the Gulf of Anadyr. Only males, mainly 7 to 11 years of age, gather here (Fig. 21). Juveniles (mainly 4-6 years old) and older males come to the beach in small numbers. Females of all ages remain confined to ice floes during autumn. A few thousand walruses gather in each rookery, lie packed close together, often in two tiers. The more mobile young walruses, lying closer to the water, leave the shore more 38 Fig. 21. Coastal rookery of male walruses. Chukchi Peninsula (photograph by P.G. Nikulin). 40 39 42 often than the older males resting farther away; then the older walruses become less densely crowded and rest for longer intervals and more peacefully. Permanent rookeries are formed at the end of August and function until the end of September -October when, with the appear- ance of ice floes, the walruses depart for them. During the period of walrus abundance on the Chukchi Peninsula there were many permanent rookeries, but now almost all of them are deserted. Temporary rookeries serve as resting sites for walruses migrating from the Chukchi Sea to the Bering Sea. Such rookeries are visited by animals of both sexes and of all ages, including females with pups. The period of their formation and the population of animals depend on the ice conditions each year. Sometimes temporary rookeries contain 1,000 or more walruses, but animals may leave within 2-3 to 7-10 days. Regular temporary rookeries are formed every year at the same places. Some 15 such were counted in recent years on Arakamchechen Island, in the region of villages Dezhnev and Ue’len, on Capes Intsov and Serdtse-Kamen’, on Idlidlya and Kolyuchin islands, at village Vankarem on Karpkarpka island, and a few on Wrangel and other islands. Finally, occasional rookeries are formed in the warm, less snowy years at the most unexpected places (sometimes even in villages). Wal- ruses, tired of long sojourns in the water, form such rookeries, which Fig. 22. Marking of a walrus. Chukchi Peninsula (photograph by V.I. Krylov). 40 Fig. 23. Marked male walrus. Chukchi Peninsula (photograph by V.I. Krylov). are extremely short in duration. Having rested on the coast, the animals quickly return to the sea, continuing their migration. On the coasts of the Chukchi and Bering seas, over 30 permanent or temporary rookeries have been recorded in various years. At present, 15 have completely vanished, with no walrus recorded in them in recent decades (Gol'’tsev, 1968). Walruses have often been observed sleeping on the water. Having filled its air sacs, the animal assumes a vertical posture with its head and the blown-up sacs of the neck jutting out of the water, and sleeps in that position (Nikulin, 1941). Seasonal migrations and transgressions. Walruses perform regular seasonal migrations (Fig. 24) but the details of such migrations are not very clear. The Atlantic walruses inhabiting our waters spend the winter-spring months in the southeastern part of the Barents Sea. In October, in the Kara Strait region, coastal rookeries are formed, but with formation of 9 In 1963, 500 Pacific walruses were marked for the first time in the Rudder rookery using a specially designed tag shaped like a large button with a base diameter of 3 cm and stem length of 5 cm. The tag was attached to a long pole and then jabbed into the skin of the walrus (Krylov, 1965). The animals permitted the markers to approach within the requisite distance (Krylov, 1965). No tags have been returned thus far (1967). 41 41 44 Fig. 24. Migrations of the Pacific walrus, Odobenus rosmarus divergens (V.A. Arsen’ev). the ice cover, the animals return to the ice. In June, as the ice floes begin to drift, the walruses usually abandon this region and enter the Kara Sea by two routes: through the Kara Strait or by encircling Novaya Zemlya from the north. The animals passing through the Kara Strait in the second half of July and in August inhabit the ice along the western coast of Yamal Peninsula from Belyi Island to Sharapov Spit. The easterly winds of August drive the ice away from here and the walruses migrate with it. By early October these animals are seen on the southern coasts of Novaya Zemlya close to the Kara Inlet, where they form coastal rookeries that function until formation of the ice cover. Another group of walruses, together with the ice floes, migrates to the northern extremity of Novaya Zemlya and spends July-August on ice floes in the coastal waters. By September, as the drifting ice moves away from the Novaya Zemlya coasts, the animals begin to congregate in the immediate proximity of the coasts, mostly on the northeastern extremity of the island, and to form beach rookeries in a section from the Oransk Islands to Cape Sporyi Navolok. By October, ice has usually 42 45 begun to form afresh on the northern coasts of Novaya Zemlya and the walruses abandon the coastal waters and move onto the drifting ice to reach the Barents Sea. They move along the western coasts of Novaya Zemlya and by the end of the month have reached their winter habitat in the southeastern part of the Barents Sea (Chapskii, 1936). The migrations of the Laptev walruses have not been studied. In August-September they form coastal rookeries, mainly on the south- western strip of the sea, on Faddei, Andrei, Preobrazhen’e, Begichev, the Peschan Islands, and in Pronchishcheva Bay. In the autumn, as the ice floes appear, the walruses move onto them and migrate northward, spending the winter in ponds of open water in the ice and in cracks fairly close to the coastal rookeries. In August, 1951, some 400 walruses were sighted at 87° N lat. and 140° E long. (Uspenskii, 1958), which perhaps could be regarded as having strayed beyond the range. The Pacific walruses spend the winter in the shallow waters of the southeastern part of the Bering Sea, reaching the area of compact ice floes in Bristol Bay. In March-April, they begin moving northward on the ice floes, approaching the coasts of Chukchi Peninsula in the Provi- deniya Bay-Cape Chaplin region. In May, walruses are seen even in the - Bering Strait, and by June have emerged, together with the floes, into the Chukchi Sea, reaching Wrangel Island. During July, however, walrus herds continue to move through the Bering Strait and spread out in their summer grounds, from the eastern part of the East Siberian Sea to Point Barrow in Alaska, only in August. In mid-October, when intense formation of new ice occurs and polar ice starts to drift southward, the walruses move to the Chukchi coasts. They swim mostly through clear waters for a distance of 50-100 km from the coasts to the Bering Strait and, entering the Bering Sea, proceed to their winter grounds. Large walrus herds swimming to the Bering Strait, even among compact ice floes, have been sighted in October, evidently struggling to escape the fast-freezing sea (P.G. Nikulin). Some walruses remain in the Gulf of Anadyr in summer on the ice floes of Cape Bering and Krest Bay until the ice has completely cleared, after which they remain in clear water. In July-August, coastal rookeries are formed in the Gulf of Anadyr (Rudder) and, at this time, some wal- Tuses move in groups or singly in the clear water along the Chukchi coasts eastward, forming new rookeries in the Bering Strait (Arakam- chechen) and even at its confluence with the Chukchi Sea (Inchoun). They remain there until the end of September or October, after which the animals move onto the ice. Rookeries are also found on Wrangel and Herald islands (Belopol’skii, 1939; Nikulin, 1941; Kleinenberg et al., 1964). 43 46 Walrus finds outside their usual range have been reported for both the Atlantic and Pacific populations. Almost every year single walruses are sighted in summer and autumn in the inlet of the White Sea and in Mezensk Bay. In 1956, a walrus was killed on the eastern Murman coast (Bel’kovich and Khuzin, 1960). Walruses have been sighted on the coasts of Norway, mainly in the northern areas. A walrus was found in 1902 and 1903 in G’esver Fjord, in 1904 close to Trondheim, in 1917 around Kirkenes, and in 1931 in Grosbaken Fjord. Two walruses were sighted in 1942 at Finmarken, and in 1953 one was found lying on the beach at Makkaur lighthouse (Lund, 1954). An interesting journey of a walrus along the Norwegian coasts has been described. In October, 1926, in the southern part of Norway (Haugesund), a male supposedly from the coasts of Scotland was sighted. The same male was seen on November 11 on the coast of Holland, at the end of November on the northern coast of Denmark (near Skaagen), and finally on January 9, 1927, was killed in Bokhuslan region in Sweden. An even more amazing journey was performed by another male in 1954 (assuming that it was the same animal throughout). In early January, a male was noticed at the northernmost tip of Norway in Bakkeby region and then sighted on Trena Island on February 3, having traveled 600 km in 26 days. From there, the walrus took to the coast on March 15 and traveled another 700 km south into Batalden region, from where it entered Swedish waters. It soon returned northward and, on March 27, beached around Sul (about 62° М lat.), then traveled still northward. In mid-April, it was sighted on Helligwer Islands (67° М lat.), again near Trena, and then at Lofoten. In mid-June, the male was seen at 69° N lat. at Sandesundver (Lund, 1954). The following journeys of Pacific walruses are known. In 1931, a herd was sighted in Korf Bay (60° N lat.) in Kamchatka, in the summer of 1935 about 500 of them inhabited Natalii Gulf (61° N lat.), in 1939 they were sighted on Karaginsk Island (59° N lat.) and on Cape Paklan (59° 40’ N lat.), and in August of the same year they surfaced on the coast of Verkhoturov Island (59° N lat.) where two of them were killed. In July, 1969, 20-25 walruses were sighted. Some walrus skulls were found on the Commander Islands while in the summer of 1969 three carcasses were found. One of them, a female 3 m long with 60 cm long tusks, found on June 17, was torn to shreds by polar foxes (some 40 of them gathered), which lends credence to this animal having been killed on the coast (Chugunkov, 1970) (Fig. 25). In May, 1940, close to Yamsk Islands (northeastern part of the Sea of Okhotsk), not far from a rookery of fur seals, a young female was killed (without embryo, length of tusks 20 cm) (Moiseev, 1951). 43 47 Fig. 25. The carcass of a female walrus found on the coast of Mednyi Island. А polar fox is seen on the carcass (photograph by D.I. Chugunkov, 1969). Possibly, this was one of the pups which hunters often took onboard ship in the Chukchi Sea. However, in the summer of 1966, on Yamsk Islands, four adults were again seen (G.A. Fedoseev). In April, 1954, a large female was killed on Kad’yak Island in the Gulf of Alaska (Fay, 1957). Reproduction. Information on walrus biology is predominantly col- lected in the summer months during the hunting season and hence data on breeding are extremely scant. Data on the Pacific walrus are somewhat more comprehensive. The growth of follicles in the ovaries commences in the first half of April and mature follicles are seen in early May; such follicles are present in gestating females as well. In the first half of June, the growth of follicles ceases. In mid-May, ruptured follicles are replaced by corpora lutea, indicating the fertilization of egg cells. Mature spermatozoa form in males in early April; by the first ten days of June spermatogenesis has ceased. The maximum quantity of mature sperm is observed from mid-April to May end. Thus the mat- ing of walruses occurs during May and to a lesser extent in early June. By this time, some females are already with embryos in the early stages of growth (Fig. 26). 44 44 48 Fig. 26. Embryo of the Pacific walrus, Odobenus rosmarus divergens. Bering Sea (photograph by V.I. Krylov). Pups are born in about the same period as mating—from April end to May end. Individual instances of birth are known in early June as well. The period of whelping extends for a month or more. Hence gestation extends for almost 12 months. Whelping duration can be ascertained from the differences in the sizes of embryos found at a given time. Thus the length of embryos investigated on July 30 varied from 13 to 21 cm, on August 30 from 27 to 37 cm, on September 14 from 34 to 47 cm, and on September 23 from 43 to 53 cm. The embryos of Pacific walruses measured in the same period were 5-11 cm longer than those of Atlantic walruses and 4-7 cm longer than those of the Laptev Sea. This suggests either a very early whelping in the Pacific walrus, or that the much larger size of this subspecies is determined even during embryonal growth (Krylov, 1966, 1969). The reproduction tempo of walruses is the slowest among all the species of pinnipeds and the whelping pattern is highly complex. Among the Pacific walrus females (285 studied) mothers were seen suckling pups born in that year and again gestating; suckling yearlings and gestating; and some gestating but not lactating; i.e., females at different stages of reproduction were encountered. The following relationships have been established between groups of females with different reproduction rhythms (Krylov, 1968): 49 Females, whelping annually 4.5% Females, whelping once in two years 12.3% Females, whelping once in three years 42.2% Females, whelping once in four years 41.0% The reproduction rhythm of Laptev walruses is similar to this pattern (L.A. Popov, 1960). Young females have a more frequent reproduction rhythm com- pared to older ones. Most females with an annual whelping cycle fall in the age group 6 to 11 years; those with a three-year cycle from 12 to 18 years; and those whelping once in four years or more are over 15 years of age (Krylov, 1968). The annual population growth is 8% since the average of pups per mature female is 0.35 (Mansfield, 1959). 45 According to other data, the annual population growth is 11.2% (Krylov, 1968). ' The prevailing view regarding polygamy among walruses (Allen, 1880; Nikulin, 1941; Freiman, 1941) has not been confirmed by recent investigations. During the period of reproduction walruses do not form harems but live in family groups of three to six animals comprising the male, female, and pups of different ages (Tikhomirov, 1964c; Krylov, 1968). Ice is not a suitable substratum for organizing a harem, whether of walruses or other pinnipeds (М. Smirnov, 1937*). Growth, development, and molt. Reliable information on the growth of walruses (and other pinnipeds) became available only after the devel- opment of a method for determining the age of each animal. Age is deter- mined from the annual depositions (rings) in the dentine of the teeth (Fig. 27). This method has been verified in many species of marine and land animals and is generally accepted (Tikhomirov and Klevezal’, 1964*). Differences in the tempo of growth of males and females are percep- tible already in the juvenile and persist throughout their lives (Table 2). Pups of both sexes grow very fast. The tempo of growth slows down 46 at two years of age, evidently due to a changeover from suckling to inde- pendent feeding. Again, a reduced tempo is seen in females of 7-9 years and males of 8-9 years. This lag coincides with the period of sexual maturity, which in females begins at 6-8 years with the peak in the 7th year, and in males at 7-9 years with the peak in the 8th year. This is followed by a fairly uniform increase in body length in females up to 11-14 years and in males up to 17-20 years. Very slow growth continues up to 20 years in females and up to 23 years in males; thereafter body growth ceases. Females older than 25 years enter a climacteric period and old males also evince no interest in mating. Among females whose 45 50 Fig. 27. Annual layers on a polished section of а cheek tooth of the Pacific walrus, Odobenus rosmarus divergens (photograph by V.I. Krylov). age was determined, the oldest was 30 years, and among males 43 years (Krylov, 1966, 1967, 1968). The body growth curve of walruses of the Canadian archipelago is very similar. Here, by the beginning of the third year, the body length of walruses averages 2 m and weight 340 kg; the average length of adult females is 2.6 m and weight about 505 kg; the corresponding values for males are 3 m and 750 kg. The maximum weight of females is 725 kg and of males 1,270 kg. Females cease to grow in the 15th and males in the 20th year (Mansfield, 1959). Simultaneous with increasing body length the tusks grow in length and thickness. The newborn pup has no teeth and the canines begin to cut a few months later. By the end of the first year, the canine mea- sures 2.6 cm (average of five measurements; V.I. Krylov) or 2-9 cm (Mansfield, 1959). Slowing down of the tempo of tusk growth is com- mensurate with that of body length. Tusks grow throughout the animal’s life but detecting their growth in adults is impossible because they wear down constantly, thus decreasing in length. The tips of male tusks diverge sideways, while those of females are slightly proximate. This difference is distinctly visible in the frontal view. Female tusks are thinner and some- what crescent-shaped, while those of the male are stronger and almost Straight. The sex ratio in newborns is close to 1:1. Suckling extends for about two years but the stomach of pups older than a year often contains mollusks along with milk; the stomach of those older than two years 51 45 Table 2. Change of body length with age in the pacific walrus (V.I. Krylov) Age Male Female No. of Mean No. of Mean animals length animals length Newborn 4 138 8 129 1-4 months 3 158 5 146 Year-old 5 198 11 185 2 years 4 238 5) 218 3 years 6 247 3 238 4 years й 260 7 258 5 years 11 281 11 268 6 years 13 292 6 276 7 years 24 297 И) 276 8 уеагз 25 306 6 278 9 years 30 306 11 286 10 уеагз 49 322 13 290 11 years 32 337 8 307 12 years 45 338 18 308 13 years 24 338 8 298 14 years 30 352 17 307. 15 years 35 354 25 299 16 years 43 351 18 308 17 years 34 358 22 304 18 years 35 358 15 312 19 уеагз 31 362 8 315 20 years 29 370 10 311 21 уеагз 19 371 8 312 22 years 12 370 4 328 23 years 17 375 7 326 24 years 14 371 8 321 25 years 14 369 4 321 26 years 13 372 4 321 27 years 4 367 28 years 12 370 29 years 7 371 30 years 12 363 31-32 years 6 370 33 years 5 372 34-38 years 7 374 sometimes contains milk. Females under parturition every year suckle two pups simultaneously—the newborn and the yearling. The first molt occurs soon after birth. In the first few days the body of the pup is covered rather densely with grayish-brown hairs, which are gradually shed and become sparse 1-2 months later. Adults have large bald patches and the hairs are small and sparse in other places; however, 52 they molt every year. The period and duration of той have not been established but evidently molt extends over a few months. In July and August, most of the walruses studied in the Bering and Chukchi seas had hairs that could easily be pulled out, while some animals had already grown strong new hairs by early July (Nikulin, 1941). In the Kara Sea in July to October, hairs of “various stiffness” were observed (Chapskii, 1936). Enemies, diseases, parasites, mortality, and competitors. The enemies of walruses are killer whales at sea and polar bears on ice (rarely also in coastal rookeries). The killer whale more often attacks the young, smaller animals but can be dangerous even to a large animal. In 1936, in the Gulf of Anadyr, whales attacked a herd of walruses under observation. Some 15 killer whales encircled a group of 60-70 walruses. Two whales broke into the center of the herd, split it into groups of 10-12 animals, and totally destroyed one such group. Meanwhile, the rest of the walruses in the compact group rapidly swam to the coast. The stomach of a killer whale caught on August 11, 1936 in the Gulf of Anadyr revealed the skin, blubber, and pieces of flesh of a walrus (Zenkovich, 1938). Only a large male can keep the polar bear at bay; however, this predator is not a significant threat to walrus herds. The only species of lice known among the ectoparasites of walruses, 47 Antarctophthirius Boh., is found in large numbers in the whiskers, in the skin folds on the sides, in the hind flippers, around the anal opening, etc. Up to 10 lice per cm? of skin were counted here and there (Ass, 1934). Eleven species of endoparasites are known. The trematode Ohdneriella rossica Skrjabin, parasitizing the hepatic ducts of the liver, has been reported only in the walrus. Orthosplanchus fraterculus Odhner, a parasite of the gall bladder, has been found in the walrus, the bearded seal, and the sea otter. The cestode Diphyllobothrium cordatum Leuckart, which inhabits the small intestine of the walrus, has also been found in the bearded, common, and Greenland seals; Diphyllobothrium latum has been reported in many species of pinnipeds and whales, in land carni- vores, and in man. Diphyllobothrium romeri Zschokke has been found in the small intestine of only the walrus. The nematode Anisakis (Anisakis) rosmari Baylis is found only in the walrus stomach. Contracaecum oscu- latum osculatum Mosgovoy and Ryjikov localizes in the stomach and small intestine and is found in many species of pinnipeds of the North- — ern and Southern hemispheres. Terranova (Terranova) decipiens Krabbe, parasitizing the stomach and intestine, has a wide range of hosts among pinnipeds and cetaceans. Three species of acanthocephalans parasitize the intestine. Corynosoma strumosum Rudolphi has been found in seven species of pinnipeds and two species of cetaceans; Corynosoma semerme 48 53 Forssell in three species of seals, in guinea pigs, and in many birds. The third species is Corynosoma valdum van Cleave. Ten of the 11 known species of helminths of walruses have been reported in the Atlantic walrus, but only five in the Pacific walrus, per- haps due to the thoroughness with which helminths were studied in the former (Margolis, 1954; Delyamure, 1955). The diseases and natural mortality of walruses have not been studied. It has been suggested that mortality may reach 18-20% of the annual population increment (Chapskii, 1936), which is rather doubtful. Prob- ably, the mortality of pups during the first two years of life is extremely insignificant because of the two-year lactation cycle and the absence of a large number of enemies. Adult mortality is mainly the result of hunting by man. Mortality, mainly of females and pups, occurs in coastal rookeries due to suffocation by much larger animals. On St. Lawrence Island, in 1936, a large herd of walruses was chased to the coast by killer whales and more than 20 of them were crushed as a consequence. In July, 1949, 54 dead walruses (mostly females) were found on Punuk Island 5 km from St. Lawrence Island (Schiller, 1954). In November, 1951, many walrus carcasses were found on the coast of St. Lawrence Island. All of them were greatly decomposed, their heads severed, and the limbs hanging from the skin. More than 50 carcasses lay on the beach and many more were afloat in the sea. Based on age and sex composition (as far as could be determined from such remnants), a whole herd had perished. The investigator felt that the animals had been killed by a massive wave (possibly caused by an underwater explo- sion) close to the Siberian coast, where they sank, then floated after decomposition, and were brought by winds and currents to the coast of St. Lawrence Island. The deaths were evidently not due to infection, as the Eskimos fed this meat to their sledge dogs and used it as bait for foxes (Schiller, 1954). The cause of mortality of this herd remains unestablished. The hearded seal, which also feeds on benthic invertebrates, competes with the walrus for food to some extent, as mollusks and crustaceans are of great importance as food for this seal (Pikharev, 1941). There are no animals which compete with the walrus in habitat selection on the coast or on ice. Population dynamics. The walrus population throughout its range has greatly decreased, mainly as a result of hunting. At present, in the Soviet Union, the USA, Canada, and Norway, hunting has been banned. The inhabitants of the Chukchi Peninsula and Alaska are permitted to hunt walruses, however, to meet their personal requirements. 49 54 Information is not available on the population dynamics of wal- ruses of the Atlantic Ocean and Laptev Sea. The population dynam- ics of the Pacific walrus is as follows: 1850-1860, 200,000; 1860 - 1880, 150,000; 1880 - 1910, 80,000; 1910-1950, 60,000; and 1950 - 1956, 45,000 (Fay, 1957). Some attempts have been made in recent years to deter- mine the population by different methods: aerial photography, aerial observations, observations from sealing vessels and in coastal rookeries. The total walrus population in the waters of the USSR was roughly esti- mated at 30,000. Considering that approximately 20,000 walruses were counted in American waters during the summer months, the entire wal- rus population may be estimated at 50,000 (Zenkovich, 1938; Collins, 1940; Buckley, 1958; Fedoseev, 1962; Gol’tsev, 1968; Krylov, 1968). Field characteristics. The walrus is a large animal with a huge thickset body, relatively small head, and broad blunt snout, with two huge tusks directed downward, which are absent in all other pinnipeds. These tusks are not visible in one-to-two-month-old pups, or they are so small as to be indistinguishable. The skin lies in large wrinkles and folds, and the hairs are sparse and coarse in adults. Walruses form large herds on ice or beaches, on which they are densely packed (Fig. 28). The animals swimming in water present a char- acteristic picture of a round head with long tusks. (V.A.) Economic Importance Walrus hunting is not important to the overall economy of the country but is significant in the economy of some regions. In the Chukchi Penin- sula, the walrus is one of the main sources of raw material for the local inhabitants. Products of walrus hunting have served as food for the peo- ple and sledge dogs, and material for building canoes and even houses (‘уагапраз”) until quite recently. Shoes, dresses, harnesses for dogs, etc. are made from this raw material. Walrus hunting is done by two methods—by boats (now banned) and on the coast. Sealing at sea was done from specially designed small wooden schooners. As the schooner approached a breeding site on the ice, motorized whaleboats were lowered from the sides and hunting commenced. Occasionally, walruses were shot directly from the schooner. Walruses were killed using firearms, for which reason crews of the hunting vessels always included some sharp shooters. The skin with the blubber was removed from the animals killed on the ice and the animals then hauled in parts onboard the schooner. Sometimes walruses swimming in water are killed (this practice has been .banned recently). In this method, a manual harpoon is hurled, 48 55 Fig. 28. А coastal rookery of walruses in the Gulf of Anadyr (photograph by A.V. Yablokov). piercing the body of the priorly injured or killed animal. The harpoon is connected by a short line to floats (inflated seal skin or various types of artificial floats) which prevent the carcass from sinking. Hunting in water has invariably been wasteful because no less than 40% of the killed animals sink and are lost (Р.С. Nikulin and V.I. Krylov). According to Canadian investigators, such losses have been no less than one-third of the kill (Mansfield, 1959). When hunting from a ship, the hide with a layer of blubber is prepared and salted in the hold and brought to the processing plant on the coast. The carcass is sometimes brought to the coast and given to the locals but quite often simply left behind. Only recently has the bringing of the carcass to the coast been made compulsory. In all meth- ods of hunting the tusks are invariably removed and used in the making of various articles. Local hunters use rifles in hunting walruses in their breeding sites on ice floes. The sea is surveyed constantly from high ground and when walruses drifting on floes are sighted, the hunters enter the sea. Hunting is done by a team of 7-10 men from motorized whaleboats or indige- nous canoes (now rare) fitted with outboard motors. The whaleboat approaches the floes as closely as possible and then the hunters open 56 50 Fig. 29. “Warts” on the skin of an adult male. Chukchi Peninsula (photograph by A.V. Yablokov). fire. The body of killed animals is cut into large pieces, which are stacked in the whaleboat and brought to the coast. Wounded walruses are har- pooned and then shot. Hunting in coastal rookeries is of little significance. It is resorted to only by the locals for whom some areas are reserved. Killing is done using special pikes mounted on long poles. No sound is permitted. Hunters follow a strict sequence of killing, conforming to traditional practices geared to conservation. Only some and not all the walruses are killed. It is believed that a colony ceases to exist if all the walruses present in it are killed. Campfires and other types of contamination of the rookeries are prohibited as such scare the animals. The tradition prohibiting the killing of all animals is based on the assumption that the same walruses return to the same region year after year. The number of coastal rookeries in the Chukchi Peninsula and their populations have decreased considerably. If at the beginning of the 1930s there were more than ten permanent rookeries, only two are known at present. Thus walrus hunting in the coastal rookeries has lost its earlier importance. At present, walrus hunting is done in summer when herds | 3)// 50 Table 3. Walrus hunting in the Chukchi Peninsula (Krylov, 1968) Year Total killed — Of which Used Losses (40%) 1932 5,180 3,750 1,480 1935 9,730 6,950 2,780 1938 11,570 8,264 3,306 1941 5,043 3,602 1,441 1944 4,654 3,324 1,330 1947 4,410 3,150 1,260 1950 5,642 4,030 1,612 1953 3,815 2.725 1,090 1956 8,140 5,814 2,326 1959 4,456 3,183 1273 1961 3,602 2,573 1,029 51 Fig. 30. Skeleton of an eared seal (Steller’s sea lion) (figure by N.N. Kondakov). migrate on ice floes close to the coasts or in regions of their summer habitat on ice floes. 50 During this same period, in Alaskan waters an average of 2,200 to 2,600 walruses were killed, of which some 1,300 were used (30-50% loss). Thus about 10,000 Pacific walruses were killed annually. Attempts 51 52 58 to shoot the much larger animals led to the preponderance of males in the catch which disturbed the natural sex ratio (Fay, 1957; Scott and Kenyon et al., 1959; Krylov, 1968). After the banning of state hunting, the catch of walruses in waters of the Chukchi Peninsula reduced: an average of 2,000 animals were hunted here until an annual limit was imposed, initially at 1,000 and later raised to 1,500. The slow tempo of reproduction determines the extremely slow restoration of depleted herds of walruses. In order to conserve their stocks, measures aimed at conservation of animals and hunting regulation are necessary. Primarily, restriction on hunting is necessary for each individual herd, so that the killing will not exceed the annual increase in population and, consequently, not deplete the stocks. The prospects of exploiting the Pacific populations may be based on the following premises: total population 50,000; sexually mature, 70% of the herd; at a 1:1 sex ratio, mature females form 35% of the total population or 17,500. About 8.5% of the females do not take part in reproduction, i.e., only 16,000 females are productive. The pups born per annum number 5,600. Keeping in view not only the need for popu- lation conservation, but also the need to reinforce it steadily, the annual catch in Chukchi and Alaskan waters should not exceed half the annual population increment or 6% of the population, i.e., about 3,000. Further, measures should be taken to cut down the irretrievable hunting losses. (V.A.) SUPERFAMILY OF EARED SEALS Superfamily OTARIOIDEA Smirnov, 1908 Family of Eared Seals Family OTARIIDAE Gill, 1866 Males are stocky and large animals while females are of moderate size with a lighter build. The head is elongated, narrowed exteriorly, and proportional to the body size. Small pinnae are present. The neck is long and movable. The limbs are very long, the fore flippers usually are not shorter than the hind ones and terminate in skin-cartilaginous tips; their undersurface is bare. Claws on the fore flippers are either absent or rudimentary. The hind flippers are capable of bending forward under the trunk and help in movement on land. The claws on the three middle digits of the hind flippers are small but well developed and disposed far from the outer margin. Claws are absent on the two extreme digits. 52 59 Color varies from black to straw-yellow in the various age and sex groups. The guard hairs on the neck of adult males are long and form a fairly perceptible short “mane”. The adipose layer is insignificant. The testes are in the scrotum. There are four teats. The width of the skull above the canines is 1/2 - 1/3 that over the ear openings. The nasals are separated posteriorly by an acute projection of the frontals (Fig. 31). The supraorbital processes are well developed. The bony palate is relatively flat. The structure of the tympanic bul- lae is complex, flattened, and relatively small. Articular, mastoid, and paroccipital processes are fused into an extremely massive projection, jutting far downward and sideways beyond the margin of the tympanic bulla. The teeth are well differentiated into incisors, canines, and molars. The upper and lower canines are roughly identical in size, with the upper ones normally developed (not transformed into large tusks), and the molars conical. The distinct milk teeth are well developed and are shed a few weeks after birth. The dental formula is: 3 1 4 2—1 I 5 С т’ Р д М хе 30 are The scapula is stunted but broad, the humerus and often the ulna is shortened, but the elements of the hand are extremely elongated and form a large flipper. Sexual dimorphism is very pronounced, while age-related dimor- phism is insignificant. The family is characterized by distinct polygamy and large harems are formed in the summer rookeries. The animals survive mainly on certain Fig. 31. Nasal portion of skuli. A—Steller’s sea lion, Ewmetopias jubatus; B—northern fur seal, Callorhinus ursinus (figure by N.N. Kondakov). 1—nasal bone; 2—premaxillary bone. 54 60 types of fish and cephalopods. Eared seals undertake regular seasonal migrations. These seals inhabit warm and temperate waters of the Northern and Southern hemispheres, entering cold seas only in the summer months. In the Northern hemisphere they inhabit only the Pacific Ocean and the surrounding seas; in the Southern hemisphere they live along the Pacific coast of South America and then along the coasts of Australia and New Zealand. They are found in the Atlantic Ocean along the coasts of South Africa and South America and also on many subantarctic islands. They are not found in the northern half of the Atlantic Ocean (Fig. 32). The systematics of the family have not been properly worked out. Eared seals have sometimes been grouped into one family with the wal- ruses and assigned the rank of a subfamily; at other times the two groups have been regarded as independent families. The majority of authors attribute a common origin to them. | This fact and the similarity of several vital morphological features justify combining the two families into a single superfamily, Otarioidea Smirnov, contrasting them jointly with the family of true seals, Phoci- dae, considered under the superfamily Phocoidea Smirnov. In general, however, within the order the family is regarded, quite justifiably, as the least specialized (primitive); some authors have compared their origin with that of the family of bears (Ursidae) of the order Carnivora. The family comprises 13 genera, of which seven are extinct and six extant. Members of four of the extinct genera (Allodesmus, Neotherium, Desmatophoca, and Pithanotaria) first appeared in the Lower Miocene, the genus Dusignatus in the Upper Miocene, and the other two (Ponto- lis and Pliopedia) in the Middle and Upper Pliocene. All the fossils of Otariidae have been found on the Pacific coast of North America and for this reason this area may be considered the center of origin of the family (Simpson, 1945; Scheffer, 1958; Mitchell, 1966). Among the extant genera, Arctocephalus is known from the Pliocene of South America and the Pleistocene of New Zealand, i.e., within the distribution range. Two other extant genera (Zalophus and Eumetopias) are known from the Pleistocene, also within their contemporary range. The extant fauna includes 6 genera with 12 species: the genus of southern sea Поп, Опа, with one species, О. byronia [= О. flavescence]; the genus of Steller’s sea lion, Eumetopias, with one species, Е. jubatus; the genus of Californian sea lion, Zalophus, with one-species, Z. californianus (with three subspecies); the genus of Tasmanian sea lion, Neophoca, with two species, № cinerea (Australia) and №. hookeri [= Phocarctos hookeri| (New Zealand); the genus of southern fur seals, Arctocephalus, with six species: A. pusillus (South Africa), forstert Fig. 32. Range of the family of eared seals, Otariidae (V.A. Arsen’ev). 62 (New Zealand), doriferus (Australia), gazella (Kerguelen), australis (South America; includes three subspecies), and philippii (possibly two subspecies) [most authors recognize three additional species of Arctocephalus: townsendi (Guadalupe Is.), tropicalis (subantarctic) and galapagoensis (Galapago Is.); and the genus of northern fur seals, Callorhinus, with one species, C. ursinus (with three subspecies). Some authors (Scheffer, 1958; and others) are inclined to divide the extant family into two subfamilies: Otariinae Boetticher with the gen- era Otaria, Eumetopias, Zalophus, and Neophoca, and a more specialized Arctocephalinae Boetticher with the genera Arctocephalus and Callorhi- nus. The second group includes species having fur with underfur (fur seals), while those of the first group have no fur. Of the six contemporary genera, two (Callorhinus and Eumetopias) inhabit the northern half of the Pacific Ocean, three (Arctocephalus, Otaria, and Neophoca) the Southern hemisphere, mostly the southern parts of all the three oceans including subantarctic waters [Arctocephalus townsendi inhabit islands off Mexico in the North Pacific], while the last genus, Zalophus, is found in both the Southern and Northern hemispheres. Two genera are found in the fauna of the USSR: Steller’s sea lion, Eumetopias Gill, and the northern fur seal, Callorhinus Gray. These ani- mals live mainly in the coastal belt of the Far Eastern seas and in the northern part of the Pacific Ocean. The economic importance of the various genera varies. The genus of northern fur seals (and, to a much lesser extent, the genus of southern fur seals) is of immense economic value for its extremely valuable fur. The rest of the genera of the family are not exploited and have no economic significance. (V.A.) Key to Species of Eared Seals (Otartidae) [of the USSR] Identification Based on External Features 1 (2). Hand of fore flipper, both on dorsal and ventral surfaces, without hair (totally naked). Narrow tips of lateral (I and V) digits of hind flippers not broader than the three middle ones (II-IV) and Slightly shorter. Hair coat on trunk with dense, soft, and silky underfur, better developed in young animals. Length of whiskers in adult males 30-38 cm, in females 20-25 cm. Body length of adult males from tip of nose to tail end in a straight line (Lcv)!° 135-200 cm, of females 110-150 cm. Color 10 Not along the dorsal curvature 2 (1). 3 (4). 4 (3). п 2 (1). 63 of adult males dark brown, young males and females silvery-gray, adult females darker with gray streaks. ......................- позу POTS! Northern fur seal, Callorhinus ursinus (р. 98) Hand of fore flippers not entirely naked dorsally, metacarpal region covered with hair. Narrow tips of extreme (I and V) digits of hind flippers slightly longer and broader than the middle ones (П-ГУ). Hair coat on trunk either lacking underfur or latter scanty. Body length of adult males more than 240 cm (up to 340). Head massive, with broad and high snout; forehead of males not steeply raised. Whiskers thick and long (up to 50 cm long and 2 mm in diameter). Color of adult males varies from straw-yellow (in summer) to brown (in winter); females yellowish-brown......... ОНА Steller’s sea Поп, Eumetopias jubatus (р. 68) Body length of adult males less than 240 cm, of adult females less than 185 cm. Head with pointed, longish snout, in males with steeply raised forehead. Whiskers very thin and short. Color of trunk dark, cinnamon-brown to sepia. .....................--- инж Californian sea Поп, Zalophus californianus (р. 92) Identification Based on Skull Features Distance [diastema] between fourth upper premolar and first molar (IV and V) teeth behind canine considerable, approximately twice distance between each adjoining pair of premolars. In males, gap equals 5-8% of condylobasal length of skull. Supraorbital process massive, fairly squarish, its upper outer edge bent slightly acurately outward. Sagittal crest in adults high and long; length of rostral part of skull measured from anterior end of premaxillae to anterior margin of orbit longer than distance from anterior margin of orbit to upper posterior zygomatic process. Condylobasal length of skull in adult males 360-404 mm, in females 290-327 mm. AS giver). ао topes Steller’s sea lion, Eumetopias jubatus (p. 68) Distance between last upper premolar and first molar (ТУ and V) teeth behind canine not more than between remaining teeth in row, constituting not more than 4% of condylobasal length of skull (males). Supraorbital process triangular with tip turned backward. Arcuate flexure of outer side not seen. Sagittal crest well developed, undeveloped, or extremely small. Length of ros- tral part of skull same or greater than the distance from anterior margin of orbit to apex of upper posterior zygomatic process. Skull size small. 56 64 3 (4). Length of rostral part of skull measured from anterior end of premaxillae to anterior margin of orbit more than that from ante- rior margin of orbit to apex of upper posterior zygomatic process. Nasal processes of premaxillae at level of anterior part of nasals not broadened and gradually taper toward upper end. Sagittal crest in adult males well developed, long and high. Condylobasal length of skull in males up to 330 mm, usually up to 300 mm, in females пр 25 о: > Californian sea Поп Zalophus californianus (р. 92) 4 (3). Length of rostral part of skull from anterior end of premaxillae to anterior margin of orbit equal to distance from anterior margin of orbit to apex of upper posterior zygomatic process. Nasal process of premaxillae at level of anterior part of nasals considerably broadened. Sagittal crest absent or very poorly developed. Condylobasal length of skull in males 220-260 mm, in females 180- 200, MM. т: лоне Gene heed: pest eee: wae ee Coe ae ae Northern fur seal, Callorhinus ursinus (p. 98). (V.H.) Subfamily of Sea Lions Subfamily OTARIINAE Boetticher, 1934 Genus of Steller’s Sea Lion Genus Eumetopias Gill, 1866 1866. Eumetopias. Gill. Proc. Essex. Inst. 5, p. 7. Arctocephalus monte- riensis Gray = Phoca jubata Schreber, 1776. (V.H.) This is the largest member of the family. Males are massive (body length over 3.5 m) and females half as large. The pinnae are small. The fore flippers are covered with fur beyond the metacarpus. The margins of the flippers are represented by a thick, compact coriaceous edge, which is totally bare. All digits of the fore flippers are devoid of claws; the tips of the digits are in the form of round disks (rudiments of claws). On the hind flippers, the two outer digits are without claws (with thickened horny disks); there are three middle digits with well-developed claws (Fig. 33). The hair coat consists mainly of guard hair; the underfur is either absent or extremely scanty. The skull is large, in adult males massive, with large crests; the latter are absent in females. Length of the skull in males exceeds 350 mm, in females up to 300 mm. Anterior portion of the skull at the level of the canines is relatively broad. Suborbital apertures are relatively large. The bony auditory tympana [tympanic bullae] are small and flattened. The 65 56 Fig. 33. Fore and hind flippers of Steller’s sea Поп (figure Бу М.М. Kondakov). premaxillae gradually narrow toward the apex. The posterior section of the palatines is almost straight or semiarcuate. That part of the jaw bearing molars is relatively massive (Figs. 35, 36, 37). Transverse notches are present on the masticatory surface of the medial upper incisors into which the cusps of the lower incisors fit. The lateral incisors are almost as large as the canines, which are thick and massive. The premolars and molars have simple roots and cusps. » iit “А их | у: К д‘ “it И hia! 57 Fig. 34. Nostril of Steller’s sea lion, Eumetopias jubatus (figure by N.N. Kondakov). Si 57 66 The genus Eumetopias occupies within the subfamily of sea lions (Otariinae) a position at the commencement ofa series of specializations. This is a genus neighboring Опа (southern sea lion), which commences the series of eared seals and thus stands at the base of the series of pinnipeds in general. Being a fully specialized typical form of the order in all respects, this genus of sea lions still bears on its skull the features of land carnivores. This is the most primitive of pinnipeds of the USSR fauna and one of the most primitive in the world fauna. Steller’s sea lion is distributed in the warm [cold] and temperate waters of the Northern hemisphere and inhabits the coastal waters of the Pacific Ocean along the Asian and American continents roughly from 33° to 65° N lat. In the west it is found from the coasts of the Korean Peninsula in the south to the northern coasts of the Sea of Okhotsk, eastern coasts of Kamchatka, the Commander Islands, and the Bering Strait. On the eastern coast of the ocean, these sea lions are found from Pribilov Islands, St. Matthew, Nunivak, Aleutian, and other islands in the north to the Californian coasts in the south. The genus consists of a single species, Steller’s sea lion, Eumetopias jubatus (Schreber, 1776). pt 7 № i = й ges > > Nie vad Zama Fig. 35. Skull of adult male Steller’s sea lion, Eumetopias jubatus [dorsal view] (figure by N.N. Kondakov). 67 57 Fig. 36. Skull of adult female Steller’s sea lion, Еитеор!ах jubatus [dorsal view] (figure by N.N. Kondakov). 57 Fig. 37. Skull of young Steller’s sea lion, Eurnetopias jubatus [dorsal view] (figure by N.N. Kondakov). 58 The economic importance of sea lions is insignificant. In spite of their relative abundance, their hunting is not well organized and only a few are caught in the USSR as well as in other places in the North Pacific Ocean. (V.A.) STELLER’S SEA LION Eumetopias jubatus (Schreber, 1776) 1766. Phoca jubata. Schreber. Die Sagetiere, 3, p. 300, Table 83B. Bering | Island. 1811. Phoca leonina. Pallas. Zoogr. Rosso-Asiat., 1, p. 104. Non Lin- naeus, 1758. Japan, Kuril’sk islands, Kamchatka. ; 1828. Otaria stelleri. Lesson. Dict. class. Н.М. 13, р. 420. Bering Island. (V.H.) Diagnosis Only species of the genus. Description The males are large, massive, and heavily built; the females are usually a meter shorter than the males and more slender in appearance. The snout is broad and blunt with a slightly upturned nose. The whiskers are long (up to 30 cm in females, and up to 60 cm in males) and thick (up to 2 mm), on average 69 to 71 [?] cm in length. The neck is long and movable, thick in males, and relatively thin in females. The thick neck of the males is the result of skin folds and long guard hairs on the nape, which perform a protective function during fights with other males. The hair coat consists mostly of guard hairs. The underfur is sparse, of poor quality, and almost totally absent inold animals. The color of the hair coat varies depending on the sex and age of the animal and the season. Newborn pups possess a soft pelage. The upper part of the body is dark brown or sandy, gradually turning brown toward the sides; the color is a monochromatic dark chestnut-brown on the belly. There are no color differences between males and females at this stage. The color turns perceptibly lighter after the first molt. Immature juveniles are a light brown. Adult females and males are almost identically colored, the back creamy, the belly dark umber with a creamy tinge, more intense in males. The nape of males is a dark purple but in females creamy. The color of the belly is noticeably darker than that of the back. The winter fur is darker than the summer fur. In winter old males are predominantly chocolate or brown, almost black on the belly. The color gradually becomes lighter in winter; in the summer months (before molt) the upper part of the body is straw-yellow (Nikulin, 1937; AS. Perlov). For the skull description, see the description of the genus. The condylobasal length of the skull in males is 367-404 mm (x = 389.7), in Plate I. Steiler’s sea lion, Eumetopias jubatus Schreber. In the center are an adult male and female. Top—movement of the animals on land and diving into water; bottom—postures of mating and playful behavior, and a group of these animals on a rock in the sea (Kamchatka, Cape Shipunsk, 1973) (figures by V.M. Smirin). 59 69 females 308 - 320 mm (x = 314.7); zygomatic width in males 211-261 mm (х = 239.8), in females 176-185 mm (х = 180.9); maximum width of the skull in males 198-238 mm (х = 223.7), in females 154-164 mm (x = 160.2) (Ognev, 1935; Nikulin, 1937; Chapskii, 1963). The average body length of males is 320-330 cm, of females about 230 cm. Males weigh 700-800 kg (sometimes over 1,000 kg), females up to 320 kg. The average weight of internal organs (for 12 specimens) is: heart 3,233 g, lungs 12,439 g, liver 18,829 g, spleen 526 g, stomach 7,687 g, intestine 21,473 g, kidneys 1,667 g, pancreas 1,420 g, and mesenteries 5,708 g (A.S. Sokolov et al.). (V.A.) Taxonomy See under the characteristics of the genus. Geographic Distribution The Steller’s sea lion mostly inhabits the coastal belt of the North Pacific Ocean where its distribution is very extensive. The range of the species has undergone no significant change. Geographic Range in the USSR Stray animals and small groups are seen throughout along the coasts of the Sea of Japan, eastern coast of Sakhalin, on Shantar and Kuril islands, northeastern coasts of the Sea of Okhotsk, eastern coast of Kamchatka, and Koryak Land. Rookeries are formed every year on [ony Island in the central part of the Sea of Okhotsk, on Ol’sk and Yamsk islands at the inlet into Shelikhov Gulf, on Kunashir Islands, Iturup, Urup, Simushir, Russhua, Onekotan, Srednev Hills, and Kamennye Lovushki, and on some other islands of the Kuril range; in the Bering Sea at some points on the eastern coast of Kamchatka (on Shipunsk, Kozlov, and Navarin capes), on the Commander, Karaginsk, and Verkhoturov islands. Some rookeries function throughout the year, while some are inhabited only in summer. In the northern part of the Bering Sea animals reach up to the Gulf of Anadyr and the Bering Strait (Fig. 38). Steller’s sea lion has been reported from neither the northwestern part of the Sea of Okhotsk (from Tauisk Bay to Ayan Bay) nor the upper reaches of Shelikhov Gulf. Geographic Range outside the USSR (Fig. 39) This species is found on the western as well as eastern coasts of the Pacific Ocean, on the coast of North Korea, and in the Pacific waters of Japan (Hokkaido Island_and the northernmost part of Honshu Island). 70 59 Fig. 38. Range of Steller’s sea lion, Eumetopias jubatus, in the USSR (V.A. Arsen’ev). On the Japanese islands, within 35 -37° М lat., rookeries of the Steller’s sea lion were recorded in the past (Ognev, 1935) but no longer exist today. In the waters of the Korean Peninsula and Japan the Steller’s sea lion was observed only in the winter months. In the eastern part of the Pacific Ocean, Steller’s sea lions are dis- tributed from southern California (roughly 33° N lat.) along the entire coasts of the USA and Canada up to the Gulf of Alaska and the Aleutian Islands, and in the eastern part of the Bering Sea from Bristol Bay to the Bering Strait. Permanent rookeries are found on many islands of the Aleutian range (Attu, Kiska, Amchitka, etc.), on the islands of the Gulf of Alaska, on the Pribilov Islands and on nearby Vancouver Island, =. ed } fj А к $. - > \\ р а yt = 4 09 re x "6 A : ~ ~~, д = с к =. 22777 777 77 as Ages Л. 2 bo A рее _ => 17755. = я и . :- =~ SALE Fe “8 й 5 === Е т Me eee Se SG 233 Br nth A ` и Е ИА: я г iu к Seine ed’ tins. Gon haa Qe Apelor у 235 Fig. 144. Skull of the common seal, Phoca у. vitulina (figure by М.М. Kondakov). 234 310 geographic distribution of the population and its affinity to the pagophilic or pagophobic group. These seals can be divided into two main types—very dark and very light-colored. The dark type is most often encountered in the group of Atlantic common seals and also among the pagophobic™ populations of the Pacific Ocean part of the range. The light-colored type is common among the pagophilic Pacific populations, 1.е., the larga. Among the dark-colored animals, the main background is either very highly pigmented or very densely covered with additional, very large spots. One way or the other, there is little space for the light-colored sections on the dorsal side of the body. Such sections are seen sometimes only in the form of narrow and sinuous light-colored streaks, more often as short, as if broken streaks in the form of light-colored dabs on a dark- colored background or as indistinct rings. Dark-colored animals are also encountered with large contrasting rings and an abundance of minute ringlets interspersed with spots of diverse shapes, sizes, and colors (usu- ally with an indistinct contour and often superposed on each other). These are colored different shades of brown-olive, cinnamon, dark gray, and black. The ventral side of such specimens is also usually dark-colored although the color of the spots on it is somewhat fainter (see Plate III). The main color of the light-colored animals, on the contrary, forms a light-colored background on which dark gray spots admixed with black ones are scattered quite densely. The black spots appear as though super- posed on the dark gray spots and rarely as dabs. This significant diversity of color acquires a very definite and distinct character in the background of geographic variation (See pp. 323 -330). The age-related color changes have not been adequately traced (Millais, 1904; Havinga, 1933; and others) due to extensive individual variation of all the elements constituting the skin pattern. Among the under-yearlings of the European common seal, a broad, dark, and more monochromatic (brownish) band extends along the middle of the back from the head to the tail. This band is only slightly interrupted by dark- colored specks and gaps which are usually smaller than those among older animals. This is also a characteristic of the Pacific Ocean forms, including the larga. The ventral side is silvery-white, with extremely rare spots in most cases. Sometimes, however, even young animals are encountered with innumerable spots on the belly, as noticed among the pagophobic form (Fisher, 1952) and the arctic larga (Chapskii, 1967). The spots on the body flanks are more diverse in number, brightness, and configuration. The color of the head is lighter than among older animals. 6 Breeding not associated with ice floes. (лохериох ‘мм Aq ainsi) (1еэл эц1 ип) ‘Teg 2840] вийпил (0э20ца) DIOY_ ‘eBIe] рие чехии] 55иэйииху вицпил (DIOY_) DIOY_ Чеэ$ плау 1O pues] Ш эа we 311 With age, the dark-colored dorsal band gradually becomes mottled with white gaps, often in the form of fairly distinct oval rings. As a result, the median longitudinal band on the back, which is distinctly monochro- matic among juveniles, ultimately disappears altogether. Further, such a process of age-related color changes is evidently not seen among all underyearlings. Animals are encountered among them (Moore, 1955*; Heinroth, 1956*; and others), which are almost indistinguishable from the adults either in differences in mottled clear spaces medially on the back, or in the development of innumerable light-colored oval rings on the dorsal side of the body, or in the color of the head. Young animals, which have yet to reach adult size, are generally sim- ilar in color to the latter but nevertheless differ in a more monochro- matic color of the back (Ognev, 1935). Fully adult and older animals have a more vivid and contrasting spottiness in which the dark and light components of coloration are fairly evenly represented and uniformly alternated, creating a bright mottled, speckled pattern usually lost on the light-colored ventral side. This is relevant, however, only to a def- inite type of spottiness that is especially characteristic of the Atlantic forms and the arctic larga. Coloration is highly diverse and very light- colored animals are also encountered together with brightly mottled and very dark, sometimes almost wholly black animals. The dry skin of the light-colored animals appears almost white from a distance. The hair coat of the under-yearlings is softer, somewhat denser, with a better developed “layer” of thin, tender, and extremely short hairs, almost like underfur. The coat of adults is considerably coarser; the seta- ceous guard hair (base of the coat) is thicker and dominates sharply over the underfur, which is difficult to distinguish among this coarse guard hair (Havinga, 1933; Fisher, 1952). The color differences between males and females have not been thor- oughly distinguished; some regard the spotted pattern as more intensely manifest among males (Havinga, 1933); others (Millais, 1904) consider the back of the female to have more dark spots while the ventral side is covered less densely with spots than in the male and the color of females is therefore considerably paler. The ringed pattern is better developed among male largas than in females and the color in general is brighter and the pattern better contrasted (Chapskii, 1967). Information on the seasonal color variation requires verification. It was pointed out (Millais, 1914*) that before every molt the general color shade, as well as the intensity of spots, turns much lighter and duller as though faded. According to other authors (Havinga, 1933), the color of the hair coat turns a dirty yellow before molt. 235 236 312 Two main types are noticed in the structure of the skull (Fig. 144) as also in the color of the hair coat. These types are associated with the ecological and taxonomic grouping of the species into two forms as pagophilic and pagophobic.’? Among the animals of the pagophobic form represented in the Atlantic Ocean, the nasal processes of the maxillary bones usually do not reach the nasal bones or sometimes only slightly contact the latter without wedging deeply posteriorly between their outer edge and the maxillae. The length of the anterior part (rostrum) of the nasal bones in most cases somewhat exceeds one-half their total length. The uncinate processes of the pterygoid bones are in the form of club-shaped thickenings or hooks slightly flattened laterally and not bent outwardly. The bony nasal septum in the choanae does not extend posteriorly farther than the anterior edge of the palatine bones. The posterior edge of the bony palate is distinctly notched, often with slightly curved sides and usually with an additional angular notch medially. The occipital foramen [foramen magnum] usually has a high angular notch on the upper edge. The premolars, except the first, are large; the second and the third, with rare exceptions, are disposed obliquely in relation to the tooth row in such a way that the anterior edge of the last tooth runs somewhat inward beyond the posterior edge of the preceding tooth. The anterior articular edge of the zygomatic bones is somewhat longer than the total of their posterior edge measured between the most prominent points. The body length of adult seals’! caught on the Dutch coasts and measured in a Straight line (Lcv) was around 150 cm in most cases, the largest of them reaching 160-165 cm” (Havinga, 1933). The maximum length evidently not measured in a straight line, but along the dorsal surface (Lc) among the Norwegian seals is 180 cm (Collett, 1911 - 1912). On the German coasts the length of the largest animals (measured evidently along the dorsal curvature) does not exceed 175 cm (Moore, 1955*). According to one view in the literature, the Pacific Ocean seals of this species are larger than their Atlantic counterparts (N. Smirnov, 1929; Ognev, 1935; Bobrinskii, 1944*, 1965*). Such an assumption, however, is justified only on an indiscriminate comparison with the latter of all The skull description is based on specimens from the North and Barents seas (nominal form). For the skull characteristics of the Far Eastern form, see under “Geographic Variation”. 71 Various authors cite the sizes of seals in their own way and do not always clearly state how the animal was measured—in a straight line or along the dorsal body curvature. 72 The values cited by Havinga expressing the body length along a straight line but up to the tip of the hind flippers have been converted using his own conversion factor (13%). 313 the seals belonging to the various Pacific Ocean forms, although their taxonomic structure is quite varied. In fact, the pagophobic seals of the Far East, for example from the Kuril Islands, are perceptibly larger: the body length (Lc) of the largest animals is around 200 cm or even more (Belkin, 1964). The larga, however, belongs to the pagophilic forms and its maximum dimensions are no different from those of the Atlantic seal at maximum length (Lc) 175 cm (S. Naumov, 1941) to 182 cm (Chapskii, 1967). Under the conditions of intense oppression by man, the small size of the European seal is perhaps wholly to be expected; at the same time, natural factors too could have been responsible for the size differences.” The length of the adult male larga averages 173 cm (Lc), of the female 162 cm. Two very large female adults caught on the southern coast of the North Sea weighed 76 and 105 kg; the maximum weight of two males was 100 and 114 kg (Moore, 1955*). Two male Okhotsk largas in the spring weighed 92.5 and 96 kg while an adult female also caught there weighed 82.5 kg (Wilke, 1954); the weight of male Bering largas in the spring reached 100 kg and that of females 88 kg. The condylobasal length of the skull of adult animals of both sexes from various parts of the range varies from 190 to 255 mm. Among adult males, the skull on average is 10 mm longer than that of adult females. The width at the zygomatic arches exceeds the width at the mastoids: among adult largas, the former measured 110-138 mm and the latter 112-125 mm; the rostral width varied from 31-45 mm. The relative weight indices of the internal organs among male largas in the spring at an average body weight of 73.5 kg are as follows: heart average 6.3%., lungs (with trachea and larynx) 13.3%., liver 26.2%., kidney 2%., and stomach 9.2%. (Sokolov, Kosygin and Tikhomirov, 1966). The length of the os penis among adults is 13.5-14.0 cm (for more detailed body and skull dimensions, see “Geographic Variation”). (K.Ch.) Taxonomy The seals: of the subgenus Phoca are usually compared with those of the subgenus Pusa, and especially with the ringed seal (Phoca hispida), from which they have inherited many features. However, the structure of the cheek teeth reveals significant differences and these along with 3 Data on the Atlantic seals are extremely scant and fragmentary and there is no accurate information on their actual ages. Moreover, the oldest arfd the largest rarely attract hunters. 237 314 some other differences probably justify the placement of these species in different but nevertheless very close genera. The taxonomic structure of this species in its broader interpretation, as adopted in this publication, has been reviewed several times and even today evidently cannot be regarded as conclusively established. In spite of describing several forms from the Pacific Ocean (see under synonyms), its taxonomic structure in the waters of the USSR has long led to two subspecies: Atlantic—Phoca v. vitulina L. and Pacific—Phoca v. largha Pall. (N. Smirnov, 1929; Ognev, 1935). However, the ecological popula- tion and morphological heterogeneity of the Pacific population of the common seal (Chapskii, 1960) detected after the 1930s necessitated a review (see p. 158). As a result, the efforts of researchers attempting to reflect the diversity of the Pacific Ocean seals by describing new species and subspecies had to be evaluated from a new viewpoint. Of particular interest at present are the attempts of Allen (1902) who described a new species of seals from the Commander Islands (Ph. steinegeri) and Inukai (1942) who pointed out that it be regarded as a special subspecies of the larga (Phoca kurilensis). This tendency again intensified in the 1960s fol- lowing the collection of extensive new data on the ecology, morphology, and distribution of the Pacific Ocean seals. As a result, suggestions were made to regard the Kuril “island” seal (Belkin, 1964; McLaren, 1966) and the pagophilic form of the larga (Chapskii, 1966, 1967; McLaren, 1966) as independent species. It is possible to interpret differently the ranks of these seals but the following premises in any case are beyond doubt. Firstly, the Pacific Ocean members of the subgenus Phoca s. str. cannot be regarded as a single form whatever the level be, species or subspecies, assigned to it. The ecological and morphological features of the larga are so sharp that they run beyond the limits of even the so-called good subspecies. Secondly, there is greater relative similarity between all the pagophobic populations of the Pacific and Atlantic than between the Pacific Ocean forms of different ecological types (i.e., between the pagophilic larga on the one hand and the pagophobic, island or Kuril, seals, and Richard’s seal on the other). As a result, the species of the common seal, Phoca vitulina, according to some authors (Chapskii, 1966a, b, 1967; McLaren, 1966) should be divided into two species: 1) common seal—Phoca vit- ulina L., and 2) larga—Phoca largha Pall. The composition of the second species (larga) thus does not include the pagophobic Pacific seals, possi- bly deserving in turn the rank of an independent species. In the present work, however, according to the note on p. 158 and the general attempt to resist extreme division of the species, the larga has not been regarded as an independent species. (K.Ch.) 239 315 Geographic Distribution This includes the waters of the continental shelf of the temperate and subarctic belts of the Atlantic and Pacific oceans with the congruent lim- ited regions of the North Arctic Ocean. The overall range, interrupted by the arctic seas and land fringes of Asia and America is distinctly divided into two isolated sections: North Atlantic and North Pacific. These rep- resent a typical-example of interrupted amphiboreal distribution. Geographic Range in the USSR Commensurate with the above character of overall distribution, the seals belonging to this group inhabit, on the one hand, our western marine boundaries and, on the other, the Far East (Fig. 145). On the western coasts of the USSR the Atlantic seal is encountered almost only in the Murman region in the east up to the inlet and in rare cases up to the isthmus of the White Sea. It is not found in the central basin and the White Sea bays. In the middle of the last century (Ber, 1862*), it was reported in Novaya Zemlya but no one found it there subsequently. In our territorial waters of the Baltic, the distribution has not been established with certainty either on the southwestern continental coasts of the Estonian Soviet Socialist Republic, on the Khiuma and Sarema islands (Aul, Ling, and Paaver, 1957), or in the coastal waters of the Lat- vian and Lithuanian Soviet Socialist Republics. References to the rather frequent encounter of this seal on the Baltic coasts including even Esto- nia (Lewis, 1885*; Grosse and Transehe, 1929) should be regarded as erroneous. In general the correct view of the distribution of the com- mon seal in the Baltic Sea was established even in the 1930s, when it was regarded as an inhabitant of only the western and southern fringes of the sea (Freund, 1933). However, in the adjoining parts of the sea, especially in Gdan’sk Bay, the common seal was without doubt encoun- tered in the recent past (Ropelewskii, 1952). Its incidental find there- fore is wholly possible on the coasts of Kaliningrad region although it may not be a regular inhabitant there. Its presence even more north- ward along the Lithuanian and the adjoining sections of the Latvian coasts is also possible. There is no doubt of its absence in the Gulf of Finland. Thus in the USSR waters of the Atlantic portion of the range the seals of this species are distributed almost exclusively at places where there is no formation whatsoever of a stable ice crust. This pattern of distribution wholly corresponds to the ecological nature of the European ` ~ “30 Fig. 145. Distribution of the common seal, Phoca vitulina, in the USSR (К.К. Chapskii). 238 317 populations of the species pertaining to the pagophobic form though some animals and groups can be encountered on ice floes.” In the USSR waters of the Far East the seal, essentially belonging to a different ecological-taxonomic (pagophilic) form, is distributed very widely from the USSR boundary with the People’s Democratic Republic of Korea to the coasts of the Chukchi Peninsula and the southermost part of the Chukchi Sea adjoining the Bering Strait.’> In the Sea of Japan the larga’ is common in the Peter the Great Gulf and is encountered north of it along the coast up to Tatar Strait where its concentrations are considerable on the continental coasts, Sakhalin coasts, and on the ice floes away from the coasts. In the Sea of Okhotsk this seal is distributed all along its periphery but unevenly and not uniformly in different seasons. In spite of adapta- tion to breeding on ice floes, in winter the larga does not remain close to the continental coasts and islands surrounded by dense, massive, gen- erally stationary ice floes, but prefers instead the more pelagic regions of the sea with drifting ice floes abounding in many pools of open water and washed-out holes. The main regions in which the larga is concen- trated in the snowy period of the year in the Sea of Okhotsk, accord- ing to the latest data (Fedoseev, 1970; С.А. Fedoseev) are: the western part of the sea (opposite the mid-northern part of Sakhalin Island and also in Terpeniya Bay), northwestern coast of Kamchatka, in Yamsk and Tauisk Bays, and in the region from Cape P’yagina to the Kola Решп- sula. In summer, with the thawing of ice floes, the entire population of the Okhotsk larga is wholly concentrated in the coastal belt of the sea. The animals are not dispersed uniformly all over the territory, however, but gather mostly in the regions that provide the most favorable food conditions. The estuarine sections of innumerable rivers are extensively used and the larga transgresses quite far from the estuary into many such rivers. The uneven distribution becomes even more perceptible closer to autumn as a result of seasonal concentrations of the population in cer- tain fixed sites year after year where coastal rookeries are formed. They are concentrated in three very important regions: (1) in the western part of the sea (Shantarsk Islands), (2) in its northeastern fringe (including Shelikhov Gulf and Taigonos Peninsula), and (3) in the coastal belt of 74 Three specimens of this species were caught in such an environment on the Murman coasts, between Kil’din Island and Nokuev Bay, in the spring of 1901 (Smirnov, 1903). 75 The larga hardly spreads along the polar coast of Eastern Siberia into the west beyond Long Strait although there are references (Rutilevskii, 1962) that it perhaps reaches even the estuary of the Indigirka River. 76 For accurate definitions of “larga” and “Kuril or island seal,” see under “Geographic Variation”. 240 318 western Kamchatka. In the first of these zones rookeries are formed on the mainland coast where they are generally few, as well as on the islands. In the western part of the Gulf of Sakhalin small rookeries and stray haunts are encountered on individual reefs of Capes Litke and Mofet (S. Naumov, 1941). Some rookeries are seen in Konstantin Bay (western- most corner of Akademii Bay). A rookery exists in Nikolai Bay. The exis- tence of rookeries has not been confirmed in Ul’bansk Bay although the larga is found there in autumn. In Tugursk Bay rookeries are known in Mamga and Kumchai bays and also near the isthmus separating Tugursk Bay from Konstantin Bay. In the Shantarsk archipelago rookeries are found exclusively on the small islands: Sivuch’i Kamni (northern and southern), Utichii, Ptichii, Srednii, and on the reefs close to Bol’shoi Shantar Island (Pikharev, 1941; S. Naumov, 1941). In the region of Taigonos Peninsula, where the larga is preferentially confined in summer to the estuary sections of the more southern rivers of Penzhinsk and Gizhiginsk gulfs, beach rookeries were noticed in the early 1930s mainly in the proximity of Capes Verkholamsk, Naklonnyi, Krainii, Povorotnyi, and Pupyr’; on Tretii and Krainii, Khalpili, Rechnaya Matuga, and Uikana islands; and also in Taigonos and Dorozh’ya bays. Farther to the southwest, the larga was similarly distributed all over the region and its rookeries were found in the same places year after year, the most important of which are Cape Ostrovnaya, estuarine sections of the Berezovka and Ireta rivers, and several points on the P’yagina and Kona peninsulas (Freiman, 1936). In Tauisk Bay rookeries were found mainly in Motyklei, Tokhar’, and Stanyukovich bays, and on Nedorazumeniya Island; they were also seen in Nagaeva Bay and on the open coast from it to Kholkhotlya (Freiman, 1936). In the early 1960s, rookeries in Tauisk Bay were seen on Capes Polonsk and Amakhtonsk, in Melkovodnoi Bay, and at some other places (Tikhomirov, 1966). Recently, Babushkina, Ushki, Shel’tinga, and Penzhinsk bays, and also sections on the western coast of Kamchatka have also been cited (G.A. Fedoseev). In the Tigil’sk region of western Kamchatka, right in the early 1930s, rookeries were detected (Lun’, 1936) at 10 places on the coast: (1) 20-25 km north of Lesnaya River, (2) 10-13 km more to the north, (3) оп Cape Kinkil’sk, (4) 40-50 km north of Amashino River, (5) on Cape Babushkina, (6) on Tal’nichnaya Islet (15 to 20 km south of Cape Babushkina), (7) near Moroshechnaya River, (8) near Cape Utkholoksk (Kavachinsk rookery), (9) on Cape Yuzhnyi (Utkholoksk rookery), and (10) in the other Moroshchechnaya River (south of Cape Khar’yuzov) (Lun’, 1936). 241 S19 Six rookeries were detected in 1958 roughly in the same section of the western Kamchatka coast: two in the lower courses of Moroshchech- naya River, one on Cape Khar’yuzovo, one on Cape Yuzhnyi, and the rest near Cape Utkholoksk and Cape Babushkina (Tikhomirov, 1966a). No definite data whatsoever are available on the concentrations of larga on the continental coast of the Sea of Okhotsk to the southwest of Okhotsk; however, as at other places, here too the larga is found in the summer-autumn season but more isolatedly, and evidently without forming regular rookeries. On Sakhalin, it is common not only along the western coast, espe- cially in the northern part commencing roughly from the threshold of Nevel’sk Strait to the Gulf of Sakhalin inclusive (Tyk Strait, Cape Lak, etc.; Gakichko, 1931), but also on the eastern side. In the northern third of Sakhalin largas are confined in larger numbers than in the more south- ern regions, on the open beaches as well as in Nyisk, Daga, Chaivo bays, and are particularly abundant in Pil’tun Bay, forming here and there fairly regular rookeries (Ambroz, 1931). The Kuril range is inhabited almost wholly but the true larga inhabits mainly the northern and southern islands, being almost wholly absent in the central part of the range inhabited by the pagophobic island or Kuril seal, which is more widely distributed there, on no less than 28 islands including the Malaya Kuril range (Belkin, 1964; Marakov, 1968). The coastal strips of the southern half of Kamchatka, western as well as eastern, fall in the normal range of the larga; the island seal, however, inhabits predominantly the sea coast. Both forms are encountered on the Commander Islands, predominantly on Mednyi Island in Zabobrovaya and Zapalata bays in the northwestern extremity (Marakov,* 1966) form- ing regular rookeries there. The majority represent the pagophobic form of seal; the larga is encountered there comparatively rarely. The summer-autumn rookeries on the coasts of southern Kamchatka are generally few. They are noticed particularly near the estuary of the Mutnaya River, on Capes Khodzhelaika and Senyavina, and also in the region of Vakhil’ River; about a hundred seals were counted in each area (Ostroumov, 1966). In Karaginsk Gulf, the larga is common and even abundant at times in winter and summer and there is even a rook- ery along the coasts of Karaginsk Island (N.N. Gerasimov). The animals inhabit other points too, including Avachinsk Bay (V.F. Muzhchinkin). Groups of them are found in the region of Kronotsk Bay from July through October in the estuarine sections of the Kronotskaya, Tikhaya, and Mutnaya rivers (in water as well as on sandy shoals) (R.M. Vik- torovskii). The range in the Bering Sea encompasses the entire coastal 320 belt from Kamchatka to the Bering Strait. The larga is evidently encoun- tered almost exclusively (arctic form of the subgenus) in all of this extensive expanse, covering the region of Parapol’sk Dolo and Kara- ginsk Island, Olyutorsk Gulf, Koryak coast, and Gulf of Anadyr, as well as the Chukchi Peninsula coast. This coastal zone is unfavorable to the habitation of the island pagophobic seal as it is blocked by the winter ice floes with the exception of only the southernmost part; this very factor is responsible for the seasonal distribution of the larga. As in the Sea of Okhotsk, it is confined here in winter to the areas far away from the coasts in the sea beyond the limits of the shore ice and highly cohesive but broken ice practically outside the continental zone. Only at places along the Chukchi Peninsula coast where the shore ice is not much and open pools are formed, is it seen there from time to time even in winter, thus providing a basis for affirming that it is confined there year round (Freiman, 1936b). In the winter-spring season the range of the larga in the northern © part of the Bering Sea enlarges roughly up to the edges of drifting ice floes whose disposition is unstable. The range can extend even beyond the limits of the continental shelf by the time of maximum ice floe formation. The main collections of the larga at this time are seen along the extensive edges of the ice floes (at some distance from the very fringe deep onto the ice floes), in the region of St. Matthew Island in the northeastern part of the sea, and almost up to the Pribilov Islands. Another part of the population at this time is usually found in the zone of drifting ice floes in the Gulf of Anadyr. With spring thawing of the ice floes and also to some extent with the movement of their masses under the influence of the prevailing southeastern current, the “pelagic” part of the range gradually recedes northward and by June end (when ice floe remnants are seen only in the northwestern corner of the Gulf of Anadyr) is confined to a nar- row strip fringing the contour of the mainland. In this summer-autumn period isolated stray sections of the range inhabited temporarily by small local populations are formed around the St. Lawrence, St. Matthew, and Pribilov islands. This situation, however, is extremely hypotheti- cal. On the southern Chukchi coasts, although rare, the larga is encoun- tered everywhere. In the Chukchi Sea west of the Bering Strait, distribu- tion of the larga extends in the form of a small tongue into the coastal belt, evidently only to Kolyuchinsk Bay (P.G. Nikulin; Tikhomirov, 1966b), which thus represents the western arctic boundary of the Pacific Ocean part of the range (see footnote on p. 317). 321 243 Geographic Range outside the USSR Coasts of Japan, Korean Peninsula, and China up to the Yangtse. In the territorial waters of the Korean People’s Democratic Republic and South Korea, some stray or a few animals are seen here and there, and probably not every year; they penetrate on the west coast up to Yonilman Bay (Nishiwaki and Nagasaki, 1960)77 (Fig. 146). On the Japanese coasts the larga is more common on Hokkaido Island. It is carried on drifting ice to the Okhotsk coast and the pago- phobic form enters from the Kuril Islands. It is also encountered on the Pacific Ocean side and on the coasts of the Sea of Japan. Evidently it transgresses south of Sangarsk Strait, along the coast of Honshu Island and up to Cape Inubo and possibly even more southward (Nishiwaki and Nagasaki, 1960). The range for the western side of this island has been shown in extremely general terms (Moore, 1965*). Sometimes the animal enters even up to Kyushu and Osumi islands. Young animals are seldom seen every year in spring on the Chinese coasts of the Yellow Sea (on Shantung Peninsula) (Leroi, 1940*; Schef- fer, 1958). Yet the seals enter even the Yangtse estuary (Allen, 1938). On the American side of the Pacific Ocean and the Chukchi Sea the seal is distributed along the entire coast from the northern Alaskan coasts through the Bering Strait and the eastern coastal section of the Bering Sea (including the Aleutian Islands). It then extends in a narrow belt all along the Pacific Ocean coast including the Alexander archipelago, and south to the Californian Peninsula (Mexico) at 28°12’ N lat. (Cedros Island). Along the northern coast of Alaska the seal reaches east beyond Point Barrow, entering the estuary of the Colville River (Bee and Hall, 1956) although it is extremely rare there; it was noticed even on Herschell Island (near the estuary of the McKenzie River). In the European part of the northern Atlantic it extends to the extreme southwestern regions of the Baltic Sea (along the Swedish coasts in the north not farther than Gotland Island), in the Denmark Strait, and the adjoining regions of the North Sea. From here, one branch of the range extends north and northeast along the Norwegian coasts up to Murman; another branch runs along the western side of Jutland, along the coasts of the Federal Republic of Germany and Holland up to Pa-de-Kale. Here the range branches again: the smaller and weaker П The authors undoubtedly committed an error by assuming that the ringed seal (Phoca hispida) inhabits within the boundaries outlined by them. Equally.erroneously, they point to the distribution of this species in the south and along the eastern coast of Honshu Island. ‘(ysdeyD УМ) рийпла о20ц4 Чеэ$ чош@оэ эцз до э8ие1 рэ12п1150099] “OPT “BIJ 2 091 323 branch runs along the French coasts, encircles Brittany, and is lost in Biscay Bay. Rarely, stray animals sometimes reach the Spanish coasts and extremely rarely even the Portuguese coasts (41° М lat.). In the north, from La Manche, the range includes the coastal waters of the British Isles and Ireland and also the west and north of Scotland: Hebrides archipelago, Orkney, Shetland, and Feeroe islands. The coastal waters of Iceland represent an isolated section of the Atlantic part of the range; here the common seal is distributed almost everywhere and is regarded as the most abundant representative of the Pinnipedia (Semundsson, 1939*). Along the eastern coastal waters of Greenland, the range extends north up to the Polar Circle and along its western coasts moves roughly up to 73° N lat. up to Upernavik (Mansfield, 1967) although some stray animals are encountered from time to time even more northward (Vibe, 1950). The American part of the North Atlantic range mainly encompasses the Canadian coastal waters north of Mann Strait including Nova Scotia, Gulf of St. Lawrence, Newfoundland Island, and the Labrador coast. Far- ther north and west, the seal is distributed along the eastern, southern, and southwestern coasts of Baffin Island and along the entire coast of Hudson Bay. Lancaster Strait can be regarded as the westernmost point of records of this seal in the Canadian archipelago (Mansfield, 1967). South of Canada, along the eastern coast of the United States, it is encountered rarely and only some stray animals reach the state of North Carolina at times. (K.Ch.) Geographic Variation Early in this century concepts regarding the species and subspecies of seals of the subgenus Рйоса $. str. were quite confusing because of the extremely incomplete descriptions that were often difficult to compare. At least three species and three subspecies belonging to the same subgenus were proposed for the Pacific part of the range alone (Allen, 1902). Later, attempts to recognize the infraspecific differences (N. Smirnov, 1908; Ognev, 1935; S. Naumov and N. Smirnov, 1936) tended to return to the viewpoint of Nordquist (1882*), according to whom the subgenus has only one species, Ph. vitulina, and only one subspecies, Ph. у. largha, vicariating with the European Ph. у. vitilina, which inhabit the entire Pacific Ocean Basin. Even the latest revision of the seals of the genus (subgenus) Phoca adopted by Doutt (1942) did not alter the prevailing situation. Moreover, Doutt affirmed that he was not able to detect any characteristics which could distinguish the animals on the American coast of the northern part 324 of the Pacific Ocean from their counterparts on the Asian coast. It was equally impossible to distinguish the Atlantic European common seals from the American seals. Doutt notwithstanding, differences between some populations of the Pacific and Atlantic members of the subgenus Phoca were once again confirmed while concomitantly the differences noticed within the Pacific populations remained unexplained (Chapskii, 1955). With further study of the systematics of the latter, the reason for the heterogeneity of the Pacific populations of the subgenus was found in their ecological dif- ferences (Chapskii, 1960). With the growing collection of specimens and greater information, the taxonomic heterogeneity was demonstrated more fully (Belkin, 1964; Moore, 1965*; Chapskii, 1965, 1966, 1967; McLaren, 1966). It was possible to differentiate these seals into two groups based on ecological as well as morphological features: pagophilic seals breeding on ice floes and aigialoid or pagophobic seals breeding on land. According to some contemporary scientists (McLaren, 1966; Chap- skii, 1966, 1967), who assign an independent species status to Phoca largha Pall., the larga is almost the only form belonging to the first group of pagophilic seals. Seals of the second group, of which five forms have been described, wholly represent a different proximate species, Phoca vitulina L. As pointed out above, in this publication the larga has been regarded as a subspecies. Three subspecies of one or the other types are encountered in Soviet waters. In general, however, the geographic variation of the common seals in our waters has not yet been adequately studied. 1. Larga, mottled seal, Phoca (Phoca) vitulina largha Pallas, 1811 (syns. ochotensis, macrodens, pallasii, petersi). Body length (along the dorsal curvature, Lc) varies from 150- 180 cm and condylobasal length of the skull, 185-230 mm. The color (Fig. 147) is relatively light, mottled: the main background on the underside of the body is whitish, light silver, while the upper part is mostly dark with a fairly dense network of rather small whitish oval streaks or rings and with small but vividly colored spots (brownish to black) scattered over the body. The newborn sports a white furry coat that lasts through the lactation period. The transverse profile of the tympanic bullae reveals a steep and high drop to the base of the shortened and rounded lobe of the external auditory meatus (Fig. 90). The less stable skull features are as follows: the nasal processes of the maxillaries extend far posteriorly along the nasal bones, while the frontal part of the nasal bones is shorter than one-half their total length (Fig. 92); the posterior edge of the zygomatic bone has 245 245 325 Fig. 147. Typical coloration of an adult pagophilic larga, Ph. у. largha. Verkho- turov Island, Kuril range, August, 1971 (photograph by S.V. Marakov). an arcuate notch (Fig. 91); the bony nasal septum in the choanae runs posteriorly beyond the anterior edge of the palatine bones to the middle and farther than the latter; the posterior edge of the bony palate has an oval outline or even an additional arcuate notch at the center, and the uncinate processes are usually flattened and turned outward (Fig. 93); all the premolars are located directly one behind the other and not inclined relative to the general line of the tooth row. This subspecies inhabits the Far Eastern seas from Peter the Great Gulf and Pos’et Bay to the limits of the range in the Chukchi Sea, includ- ing Tatar Strait, Sea of Okhotsk, northern and southern islands of the Kuril range, the Bering Sea, and partly the Commander Islands. Outside the USSR it is found in the coastal waters of Alaska (in the Chukchi Sea in the east to the limits of its range), Aleutian, Pribilov and other islands in the eastern part of the Bering Sea, south to Bristol Bay. South of the USSR border, it is found up to the northern coasts of Japan and the Yellow Sea inclusive. Two races (natio) were identified within this form (S. Naumov and М. Smirnov, 1936). One of them, п. pallasii, distributed in the Sea of Okhotsk and in the northern part of the Sea of Japan, is a very small form according to S. Naumov and N. Smirnov (condylobasal length 188-228 mm). The other race, п. /argha, inhabiting the Bering Sea, 246 326 differs from the preceding one in much larger craniometric dimen- sions’ (condylobasal length 191-238 mm). The differences between these races are artificial since the Bering larga, according to the above, repre- sents a mixture of the typical larga with the pagophobic populations (Kuril or island seal). It is possible that the Okhotsk largas in fact are somewhat smaller than the Bering counterparts and differ from them in other features, but-they should be compared exclusively with the pagophilic kin. These “races” are not consistent in the proposed form. 2. European common seal, Phoca (Phoca) v. vitulina Linnaeus, 1758 (syns. canina, variegata, scopulicola, littorea, Птпае, thienemannit). Does not differ much from the larga in size. The body length (along the dorsal curvature, Lc) varies from 175-180 cm; the condylobasal length of the skull of adult males varies from 203-217 mm and that of females 190-205 mm, average 205 mm (Ognev, 1935). The color is more or less dark. The main background on the upper side is dark, quite often almost black, interrupted by white, uneven cel- lular streaks; the underside is lighter; dark-colored spots on the general dark-colored background are less noticeable and less contrasting. Pups do not sport a white fur coat as it is shed before or at the moment of birth. The transverse profile of the tympanic bullae is flattened with an insignificant short step-like drop to the base of the elongated lobe of the external auditory meatus which is almost straight anteriorly and asym- metrically pointed at the tip. The posterior edge of the zygomatic bones has an angular notch; further, the upper portion of the fork has a slightly convex contour (Fig. 91). The rest of the skull features likewise stand in contrast to the corresponding ones of the larga (See its characteristics оп pp. 323-326). This subspecies inhabits the southernmost part of the Baltic Sea and from St. Nos in the west to the state boundary in Murman. Outside the USSR it is encountered throughout the rest of the European section of the range. 3. Island or Kuril seal,’® Ph. (Phoca) у. curilensis Inukai, 19457? (syns. chorisii (?), stejnegeri, nummularis,®° macrodens, and insularis; the name richardi was also used sometimes). 78 Known on the Kamchatka and Commander Islands from the end of the nineteenth century under the local name “antrus” or “апиг”. 79 The first name for this form should be stejnegeri J. Allen, 1902 (editor’s note). (У.Н.) 80 See the note on р. 307 (synonyms). 247 327 This is the largest form of the species with a body length (along the dorsal curvature) ranging from 140-204 cm; the condylobasal length of the skull varies from 208-247 mm (Belkin, 1964). Color (Figs. 148-151) varies intensely but an extremely dark, often almost black, main background is a characteristic feature. The ventral side is often very light in color and interrupted by numerous ring-like clear spaces, mostly isolated and dispersed unevenly. Among other ani- mals, the body underside is as dark in color as the upper portion. A broad and obtuse snout is a characteristic feature. Newborns, as in the case of the European common seal, shed the embryonic coat in the mother’s womb. The transverse profile of the tympanic bulla has the same short projection to the base of the lobes of the external auditory meatus as in the common seal, the bony lobe itself being long, set sideways and often pointed. The posterior edge of the zygomatic bones has an angular notch; the longitudinal bony septum in the choanae is usually very poorly devel- oped; a characteristic feature is the angular flexure of the upper contour of the profile in the zone of interorbital constriction. The second and third premolars are usually set obliquely relative to the tooth row. The articular fossae are broadly exposed, flattened. In other skull features, Fig. 148. Dark-colored Far Eastern island seal (“antur’’), Ph. у. kurilensis. Iturup Island, Kuril, July, 1966 (photograph by S.V. Marakov). 328 247 Fig. 149. Color details on the vent:al side of the female island seal, Ph. у. kurilen- sis, Iturup Island, July, 1966 (photograph by S.V. Marakov). 248 Fig. 150. Variation in dark coloration of the adult island seal, Ph. у. kurilensis, Iturup Island, July, 1966 (photograph by S.V. Marakov). 248 249 329 Fig. 151. Island seal (“antur”), РИ. у. kurilensis, Mednyi Island, July, 1972 (pho- tograph by S.V. Marakov). there is only a shift in the specific features or they stand between the characteristics of the larga and the Atlantic common seal. This seal is found on the Kuril and Commander Islands and on the eastern coast of Kamchatka. Outside the USSR it is found on the Aleutian and Pribilov islands and Japan and probably on the coasts of southern Alaska. The tendency to combine the island seal with the subspecies Phoca vitulina richardi found on the American coasts (Moore, 1965*) cannot yet be regarded as substantiated for the simple reason that the skin coloration of these seals differs sharply. Outside the USSR three more pagophobic subspecies are recognized (Scheffer, 1958; King, 1964): (1) Ph. v. richardi Gray, 1864—American waters of the Pacific Ocean from the Alaskan Peninsula to California, and eastern part of the Bering Sea, including the Aleutian and Pribilov islands; (2) Ph. у. concolor De Kay, 1842—western Atlantic American 330 and Greenland waters; and (3) Ph. v. теПопае Doutt, 1942—Lower and Upper Seal Lake on Ungava Peninsula (eastern Canada). Craniologically, Ph. v. richardi is somewhat closer to the island (Kuril) seal and correspondingly differs sharply from the larga, but the skin coloration of the adults and semiadults of the two is often similar.. The range of this seal evidently does not extend in the north beyond Bristol Bay and should thus be limited to the regions free of ice floes (Moore, 1966*; McLaren, 1967*). Insofar as the other two forms are concerned, their morphological features require further study. (K.Ch.) Biology Population. It has not yet been possible to determine precisely the pop- ulation of the common seal (including the larga) in the USSR waters in spite of several efforts in this direction. In some sections of the Sea of Okhotsk a visual count has been made in the rookeries but this has not helped to assess the total reserves of even the larga in the Sea of Okhotsk. Some local populations have been estimated more accurately. Thus in the rookeries on the Kuril Islands the population was put at 5,250, of which 2,000 - 2,500 constituted island (pagophobic) seals (Belkin, 1964). Very similar figures were recorded even later but without differentiation into island and larga seals: 6,000 on the Kuril Islands, 1,000 on Bering Island, 1,000 on Mednyi Island, and 100 on Seal Island (Marakov, 1970). Insofar as the total population of seals of the subgenus Phoca s. str. in the Pacific Ocean is concerned, it has very approximately been estimated in the range of 20,000 - 50,000 (Scheffer, 1958) to almost 400,000 - 450,000 (Chapskii, 1966). Attempts were made toward a more rational census of the seal pop- ulation. Based on the average area of the ice floes inhabited by lactating animals and their average density (1 to 3 newborn/km’, area of “rookery” 150-200 km?, and their number in the Sea of Okhotsk and the Bering Sea over 2081), the maximum number of pups is approximately 15,000. This figure is evidently much less than the actual number. Different figures were arrived at by the end of the 1960s to the early 1970s. According to aero-visual counts on the spring ice floes, the total larga population in the Sea of Okhotsk was put at 170,000 (Fedoseev, 1971). A similar method had been used sometime before to estimate the larga population in the Sakhalin section of the range. The results were: about 10,000 (with pups) in the region east of the mid-northern section of Sakhalin, about 4,000 in Terpeniya Bay, and 8,000- 11,000 in Tatar Strait (Fedoseev, 1970; G.A. Fedoseev). 81 This calculation was based on the data of Tikhomirov (1966b). 250 331 А very large proportion of the total population is concentrated in the Sea of Okhotsk. There is a view that the larga is second to the ringed seal in “natural level of population” (Pikharev, 1940*; Fedoseev, 1966) while its total reserves in this sea are not less than 20-25% of the total population of all species of seals (Tikhomirov, 1966a). The larga is most abundant in the following areas in one season or the other: (1) southwestern Sakhalin part of the sea north of the southern Kuril Islands; (2) western corner of the sea from the Gulf of Sakhalin to Uda Bay, including the Shantarsk archipelago; (3) northeastern part of the sea including Shelikhov Gulf; (4) the region adjoining northwestern Kamchatka; and (5) the coastal belt west of P’yagin Peninsula up to the estuary of the Tauya River. The larga population elsewhere in the continental expanse between Tauisk Bay and the Shantarsk archipelago is evidently much less than in all the regions cited above. Such a large number of summer-autumn coastal rookeries as found along the coasts of the Sea of Okhotsk is not found anywhere in other parts of the Soviet Far East. The maximum number is concentrated there in the western corner of the sea (mostly in the Shantarsk archipelago and Yamsk-Siglansk region) and along the western coast of Kamchatka. The most detailed information on the distribution and population of the rookeries is available for that part of the sea from the Gulf of Sakhalin to Uda Bay, including Shantarsk Island. The concentrations of the larga here along the mainland beaches are few and rather small, with the exception of Konstantin Bay; at one time there in the 1930s, up to eight rookeries with a total population of 900 to 4,000 seals were reckoned (S. Naumov, 1941; Pikharev, 1941). The population of the larga is not high in the western part of the Gulf of Sakhalin (S. Naumov, 1941). In Nikolai Bay in 1929, one source put it at 100 or a little more and another source in 1938 at 1,000. The larga is not very rare in Ul’ban and Tugursk bays in autumn. In Uda Bay there are obviously no large congregations. The population is some- what more on the island rookeries in the Shantarsk archipelago, espe- cially on the Sivuch’i Кати, Srednii, Ptichii, Utichii, and also Мау Shantar islands. In 1929, the total population of this region was put at 2,000 (S.P. Naumov, 1941) and in 1938 and 1939, at 16,000 to 18,000 (Pikharev, 1941). The population was the same in 1932 but later dou- bled (27,500 -35,000) (P.G. Nikulin). In some very large rookeries оп the Shantarsk Islands, especially on the Sivuch’i Кати! (Shantarsk archi- pelago), 6,500-7,000 (Pikharev, 1941) or even more (P.G. Nikulin) were reported in the 1930s. In 1958, the total population of the larga in these rookeries was put at 1,000- 1,600 (Tikhomirov, 1966). 250 251 332 Fig. 152. Гагра, Ph. у. largha (left) and island seal (“antur’’), Ph. у. kurilensis, Mednyi Island, June, 1969 (photograph by S.V. Marakov). Considerable concentrations of the larga were noticed in 1929 - 1930 in the Yamsk-Siglansk region in the fore-estuary expanses at Ireti and Yama and very significant rookeries at Ostrovno, Kamyl, and the Bero- zovka River estuary. Not even a rough estimate of the population there was ventured. The largas were fewer in Tauisk Bay than in the preceding region; coastal rookeries were few and small in the 1930s (albeit large numbers occurred in the western part of the bay) (Freiman, 1936). In the 1960s, the number of largas in Tauisk Bay was small, barely 2,000. The low population is explained by the relatively high coastal habitation and possibly low food availability (Tikhomirov, 1966). In terms of larga population, Shelikhov Gulf is next after the Yama region and the western coast of Kamchatka (Fig. 154) but the rookeries there, as already mentioned, are few, especially in the region of Taigonos Peninsula although the data on the former population have been given for only two—Uikan rookery (up to 500) and Cape Verkholamsk rookery (up to 400) (Freiman, 1936). No less than 12,000 seals were counted in the 1930s in 10 rookeries in the Tigil’sk region on the western coast of Kamchatka (from the Moroshchechnaya River to the Tigil’ River). The population in some 333 \ 251 Fig. 153. A herd of largas, Ph. v. largha, Karaginsk Island, Bering Sea, July, 1968 (photograph by N.N. Gerasimov). 252 Fig. 154. A section of the rookery of the larga, Ph. v. largha, western coast of Kamchatka in the estuary of the Utka River, September, 1967 (photograph by D.I. Chugunkov). rookeries went up to a few thousand (Tal’nichnoe, 15-20 km south of Cape Babushkina, 3,000-4,000 as also on Cape Utkholoksk) (Lun’, 1936). In 1958, six rookeries with a total population of about 6,000 were found in the same region (Tikhomirov, 1966). The total population of the larga in the western part of the Bering Sea has not yet been determined. It is known thai it is not rare along eastern Kamchatka and Koryak Land (from Cape Olyutorsk to Cape 252 334 Navarin). It is quite abundant also in the coastal waters of the western and southern regions of the Gulf of Ападуг; by the early 1930s, the proportion of the larga there constituted up to 15% in the local hunters’ catch but dropped to a few percent on the Chukchi coasts where this seal inhabits in maximum numbers the Pinkegnei, Tkain, and Privideniya bays (region of rivulets on bald patches) (Freiman, 1935а*). The population on the- Commander Islands, as already noted, is extremely small; it is usually represented by the settled pagophobic form of island (Kuril) seal; the larga is, however, encountered there as a rare find. While only one rookery was known there in the early 1930s with a hundred seals or slightly more (Barabash-Nikiforov, 1936), evidently there are now no less than 15 rookeries (although no exact figure is available). On the Kuril Islands, the island seals have been reported on 28 islands, with pups included on 11 of them (Velizhanin, 1967). During their residence on drifting ice floes, the largas are much more numerous in the central and eastern parts of the sea, especially in the region of St. Matthew Island, in the zone of ice floes south almost up to the Pribilov Islands, and northeast of the latter. Here, in the spring of 1963, over 90% of the seals encountered on ice floes were largas. A nearly similar picture was noticed even in the preceding season; in the more northern regions of the sea, between St. Matthew and St. Lawrence islands, the larga accounted for only a little above 20% (Kosygin, 1966a*; Tikhomirov, 1966). Larga in the Bering Sea has been reported as con- stituting not less than 30% of the total seal population inhabiting there in the zone of drifting ice floes far from the coasts, and occupies sec- ond place after the ribbon seal in terms of population (Tikhomirov, 1966b). In the other regions of the Far East, considerable concentrations of larga are seen in Tatar Strait although these groups are much smaller than in the Sea of Okhotsk and the Bering Sea. In Tatar Strait the larga predominates markedly over all the other species of seals. The reserves in the Sea of Japan are small; it is evidently maximum on the threshold of Tatar Strait and in Peter the Great Gulf. It has, however, not been possible to express the population numerically. The population of the common seal in our waters of the Atlantic Basin is extremely small. These seals are particularly rare in the south- eastern Baltic Sea and relatively few on the Murman coasts. It is signifi- cant that the coastal people so far have not given their own name to this species.*? Only some stray animals were caught in the Murman region after several years, at the very end of the last century (N. Smirnov, 1903). 82 At the beginning of this century, this seal was known in Murman as “kamenka”. 253 253 335 The seals of this species were not caught in nets, which was the form of hunting in vogue in the 1930s. Habitat. The Atlantic common seal and the Pacific pagophobic forms inhabit selected sections of the coastal zone in a fairly settled manner. Within the southern Baltic, on the Polish and German coasts, the seals emerge onto land in the uninhabited sandy or rocky coasts, small islands, spits, and bald patches in the river estuaries and bays. On Murman, they are evidently confined to the bays and estuarine sections of rivers, espe- cially of the Voron’ya; they probably take advantage of the rocky coasts and islets. The Far Eastern pagophobic populations (island seals) select for their rookeries reef ranges and groups of individual rocks, as well as sandy-pebbly coastal sections which offer them protection from the surf (Fig. 155). Such conditions on the Kuril Islands are provided by reefs, small coastal islets, sections of very large uninhabited islands, low rocky ledges, niches between rocks, and coastal shoals (Belkin, 1964а*; Velizhanin, 1967). Some animals (males) at places, for example on the Lovushki Islands, rest on small isolated rocks almost fully submerged in the strait and overgrown with algae (Belkin, 1964b*). Fig. 155. Typical habitat of the island seal (“antur”), Ph. у. kurilensis, on Mednyi Island. Commander Islands, Zapalata Bay, June 6, 1962. Sea otters also live here (photograph by S.V. Marakov). 254 336 The island seals on the Commander Islands (especially on Mednyi Island) select diverse biotopes for rookeries. These may be sections of the low coast, pebbly or rocky talus zones at the foot of a precipitous cliff, scarps, sloping projections of rocks, etc., especially under reefs pro- tected from the surf. For temporary residence, these seals inhabit large exposed rocky mounds that have been rounded by intense wave action and smoothened by surf abrasion (Marakov, 1968, 1969). The Bering Island population is confined mainly to the region of the Barrier Reef (Marakov, 1967b). The pagophilic seals, 1.е., the largas, associated with the coastal land are not permanent settlers since they abandon the beaches jammed by ice floes in winter and spend the winter-spring season on drifting ice floes. There is a distinct difference in the site selection of the larga for whelp- ing, lactation, and molt. Nevertheless, it is important for it that the ice floes be quite stable, clean, without large hummocks, and at the same time not excessively compacted, with an abundance of water pools to provide access to the water. Largas avoid stationary shore ice or a highly compacted mass of broken ice floes. According to some observers, they exhibit a distinct preference for ice floes along the edges (Tikhomirov, 1964) since they are very sparse; according to others, they are encoun- tered not only close to the edge but also far from it, inside drifting ice floes that are highly compacted (Kosygin, 1966). The larga begins to be seen directly on the coasts immediately after the thinning of the ice cover and access to them becomes available. Evidently it appears earliest and in some numbers close to the river estuaries where the ice floes disintegrate usually more rapidly and where the food conditions are more favorable. Immediately after coming onto the beaches, initially in small numbers and later en masse, the seals begin to move on land to form temporary rookeries. At the beginning of summer, these rookeries appear disorganized and tentative and the animals rest here and there without confinement to any one particular site for long. The islets and spits in the river estuaries that dry up in low tide often serve as rookery sites; where such sites are not available, they are confined right on the gentle beaches in the lower courses of the river itself (Tikhomirov, 1966b). When attracted to the fish migrating for spawning, they themselves enter the estuaries and the lower courses of rivers and even ascend tens of kilometers up the estuary of some rivers. Transgression into rivers is a regular phenomenon. Evidently there is no single sufficiently deep river abundant in fish in the spawning period which has escaped the notice of the larga. Its long transgres- sions, sometimes hundreds of kilometers, into rivers such as the Amur and the Anadyr, have long been known. In the former river the larga 254 255 98 Fig. 156. Largas on a drifting ice floe in Litka Strait, Karaginsk Island, Bering - Sea, early June, 1968 (photograph by D.I. Chugunkov). was seen even 400 km away from the estuary (Nikol’skii, 1889). In the past the larga ascended 60 km away from the sea along the Sakhalin rivers Poronai and Тута. Transgressions for 10-15 km were noticed in the Tyma River comparatively recently (Ambroz, 1931). The larga trans- gresses regularly into many rivers of western Kamchatka such as the Voyampolka, Khairyuzov, Belogolovaya, Tigil’, Kavacha, Utkholok, and Sopochnaya, ascending 5-35 km along them (Lun’, 1936; Ostroumov, 1966). These seals also enter some rivers on the southwestern coast of Kamchatka. — The larga transgresses for considerable distances into the rivers of eastern Kamchatka. In the 1960s, they were found 35 km from the estuary of the Kamchatka River; a few decades before that they transgressed even farther, up to Klyuchi village and even up to Kozyrevsk, 250 km from the estuary. The larga transgresses for distances of 25-30 km into Zhu- panov, Vyvenka, Kultuchnaya, and Apuka. In many other rivers, however, these seals are concentrated close to the estuary, often a whole herd, or transgress only into the lowest courses (Ostroumov, 1966). Instances of distant transgressions of the common seal, episodic in nature and regarded as exceptional events, have been known in the remote past even in the rivers of Western Europe, especially in the Elba, 338 in which they were sighted at a distance of 646 km, almost 700 km, and even 757 km from the estuary (Moore, 1955*). By autumn, sightings in the rivers become increasingly rare and the bulk of the well-fed largas concentrate on the sea coasts close to their favorite haunts, where they form regular (permanent) rookeries. Groups of isolated rocks smoothened by water and close to the coast or the rocky shore protected from the surf represent preferred sites in the Sea of Okhotsk. In any case there is a distinct preference for land sections submerged in high tide. Sections with an intensely rugged coastline and abounding in reefs and tiny islets meet these conditions to a very large extent. Some dependence could even be established, other conditions remaining comparable, between the degree of ruggedness of the coast and the nature of the rookery: the more rugged the coast, the more rocky it is and the more abundant the reefs on it, the larger the sites suitable for rookeries. Here the seals are more numerous but the number of animals in each rookery is small. Hunters distinguish three main types of rookeries (Tikhomirov, 1966a). One is the rookery located in a highly rugged locality .where sections of large pebbles are interspersed with rocky hummocks and large isolated rocks. Such, for example, are the rookeries on the Sivuch’i Islands. The second type is found on exposed sandy or pebbly laidas (low coastal plains dissected by tortuous rills), islets, spits, gently descending into the water (rookeries on Cape Polonsk, in Tauisk Bay in Melkovodnaya Gulf, and in western Kamchatka in the Moroshchechnaya River). The third, and most common type of rookery, is represented by coastal reefs and rocks, for example on Capes Khairyuzovo, Yuzhnyi, Utkholoksk, etc. (western Kamchatka). At several places the larga inhabits land together with or in the neighborhood of pagophobic island seals. Such instances are particularly numerous in the northern and southern parts of the Kuril Islands; in the southern parts of the range, largas are even permanent settlers (Belkin, 1964). The biotopes of both forms of seals there are almost identical. A somewhat similar situation is noticed on the Commander Islands where the larga is, however, far less numerous, constituting a rare find in the rookeries of island seals. Food. The common seal in general is a distinct fish-eating animal but not specialized for a limited number of any particular species and feeds on quite diverse, mainly large fishes available on the coasts at a given time of the year. At the same time, various invertebrates constitute a significant proportion of its food ration. For nearly two-thirds of the year, from spring to autumn, the larga is assured of abundant fish food. In eastern Sakhalin, in spring, it takes 256 339 advantage of the approaching herring; in summer it feeds partly оп herring and also on humpbacked salmon [pink salmon, Oncorhynchus gorbuscha] and Siberian salmon [chum, O. keta]. When the summer run (migration) of salmon slackens, the larga takes to consuming river fish—“kundzha,” [Sakhalin char, Salvelinus leacomaenis] rudd etc. Some seals while chasing these fishes reach far up the rivers. The larga takes to capelin during its en masse availability in the first half of July (Ambroz, 1931). In western Kamchatka, in late spring and early summer, the larga feeds on char and sea trout which descend from the rivers into the fore- estuarine regions of the sea, and later on smelt and capelin which form spawning schools on the coasts. Later, from the middle and second half of June, the larga takes to feeding on herring which comes close to the coasts in large numbers. At the end of the herring season, usually from early July, it takes to chasing salmons—chum, red salmon, pink salmon, etc. (Lun’, 1936). On the opposite side of the Sea of Okhotsk, on the Shantarsk Islands, it takes to navaga (Pacific navaga [saffron cod, Eleginus gracilis) in spring as supported by the coincident en masse approach of this fish with the increasing population of the seal (Lindberg and Dul’keit, 1929). In Schast’ya Bay (western part of the Sea of Okhotsk) a similar relation has been reported between the gathering of the larga and the availability of smelt in shoals (S. Naumov, 1941). In Tatar Strait, at the end of May, numerous largas were noticed; they were evidently drawn there by the availability of herring. In the snowy period, when the adult seals are confined to the regions of drifting ice floes far removed from the coasts, the food regime of the seal is somewhat different. Thus in the southern part of the Sea of Okhotsk, not far from the Hokkaido coasts (Wilke, 1954), the larga feeds in April mainly on pollock (Theragra chalcogramma) followed by herring, while cephalopod mollusks, especially the octopus (Octopus daeffeni), account for a much smaller quantity of food intake. The ratios between these food groups are hardly constant. This seal readily feeds on whatever species are available at the time of foraging for food. The larga concentrations in the southeastern part of the Sea of Okhotsk, on the Kuril Islands, feed in the spring-summer period quite intensively, consuming roughly equal proportions of fish and cephalopod mollusks. The following fishes were detected in the stomach of largas: Atka mackerel, rockfish (Sebastodes sp.), walleye pollock, and saffron cod; octopuses from among cephalopods; and squids of the genus Gona- tus to a much lesser extent (Panina, 1966). At the end of the reproductive cycle, quite often even after incom- plete molting, the larga migrates into the coastal zone, concentrating 257 340 mainly in the estuarine sections and the lower courses of rivers where salmon migrate in large numbers for spawning. The seals reach there either at the time of high tide or at a much later period or are even confined to the fore-estuary or bar sections. On the western coast of Kamchatka, in the region of Tigil’ River, even brooklets to which the pink salmon go for spawning are quite often literally blocked with the seals, which are caught there in tens or even more. Not being able to penetrate the brooklets, the seals at times even crawl in high tide to the estuary, thereby blocking the entry of fish until their hunger is satiated. Tens of larga transgress into very deep and broad rivers and in rivers such as the Tigil’ even 200-300 or more may be seen (Lun’, 1936). The same pattern of transgression is seen at other places also. While chasing the fish, the larga can dive under water without respite for up to 400 m, gaining a speed of over 4 m/sec, or it can dart several meters into the air (Chugunkov, 1969). For 1.5-2 months (August and even September), the seals are not satisfied with the fore-estuary sections but transgress even into the lower courses Of salmon-bearing rivers and brooks. This transgression ceases abruptly only in October. Only rare seals, not yet fully fed, hunt for food at this time even in the largest rivers. The affinity of the larga for coastal sections and rivers abundant in salmon, consuming much and damaging the commercial fish (often snatching only the daintier and meatier portions of the back), is respon- sible for its reputation as a carnivore inimical to the fishing economy. However, because of incomplete data, it is yet difficult to establish the exact extent of damage inflicted by the larga. In addition to fish and cephalopod mollusks, crustaceans occupy a definite place in the diet of the larga. Among these crustaceans, the main ones are shrimps and partly amphipods. In Tatar Strait common shrimps of the genus Sclerocrangon and amphipods of the genus Gammarus have been found time and again in large quantities in the stomach of seals caught at the end of April; in some cases the stomachs were literally crammed with shrimps (Freiman, 1936b). How important shrimps are as food items for the larga in this region is deomonstrated by the following data. Of 36 seals caught on May 12-14 in Nevel’sk Bay, the stomach of 21 animals was filled with shrimps; of the remaining 15 caught on May 15-19 in the environs of Cape Lak, the stomach of nine seals also contained only shrimps while the remaining six seals contained shrimps with fish remnants (S. Naumov, 1941). In the spring-summer period the adult and semiadult Bering largas feed mainly in the morning (up to 9:00 a.m.) and in the evening (after 341 4:00 p.m.) and consume a large amount of mixed food although fish con- stitutes the main item. at this time. Slightly less than one-half of the seals feed on a single type of food at this time of year: only fish 29%, only crustaceans 11%, and only cephalopods 7% (Gol’tsev, 1969). In Tatar Strait (Gol’tsev, 1971) the larga in spring (in March - April) feeds mainly on pollock, octopuses, navaga, squids, and sand eels. The food of the larga in Peter the Great Gulf has not been adequately studied. Evidently in Spring (in March), navaga constitutes the main food followed by floun- der (Liopseta sp.) and thirdly by perches (of the family Scorpaenidae); in addition to these, the larga there also consumes shrimps (Pandalus sp.) (Goltsev, 1971). In the gulfs of Anadyr and Karagin, where studies were carried out for three years (Gol’tsev, 1971), the larga feeds in spring on three groups of animals: (1) different species of fish, mainly the polar cod, pollock, sand eel, goby, and to a lesser extent navaga, armed bullheads (Ago- nidae), stichaeid blennies, etc.; (2) octopuses; and (3) crustaceans (mainly the shrimp Pandalus goniurus and more rarely the following species: Spirontocaris macarovi, Eualus gaimardi, and Е. fabricii). Groups of crus- taceans, such as amphipods, crabs, and hermit crabs are very rarely found in the stomach (usually in not more than 2%). The following aspects could be set down with respect to the food of the larga: (1) evidently there are no total or prolonged seasonal absti- nences in the feeding of the larga even at the time of molt; (2) the “food spectrum” of the larga is quite broad. Evidently it is this phenomenon that helps them to survive under diverse biotopical conditions which, in the ultimate analysis, explains the unusually extensive distribution of this species, extending from the Chukchi to the Yellow seas; and (3) eurytro- phy in turn is ensured by the fact that the large can get at food not only in the shallow and surface sections of the sea, but also at considerable depths, of the order of 300 m or more. Some other feeding patterns have also been pointed out (Gol’tsev, 1971), especially the daily feeding rhythm: the animals of this species feed mainly in the morning and evening hours. Some age-related changes of food intake were also noticed; the juveniles after lactation feed initially on amphipods, shrimps, and schools of small fishes while the proportion of pelagic fishes (navaga, polar cod, and sand eel) as also cephalopod mollusks increases later. Evidently bits of algae and stones and sand accidentally enter the stomach along with the food and are totally unin- tentional. Amphipods disappear from the food of adult largas and the spe- cific proportion of benthic organisms, fishes, and crustaceans (decapods) 258 342 correspondingly increases. Among the benthic and demersal fishes con- sumed are the flounder, halibut, goby, stichaeid blennies, and some poachers; among decapods the snow crab and others. The quantum of cephalopod mollusks consumed also increases considerably. Although there are no clear ideas so far on the seasonal changes in the feeding intensity of the larga, it is not entirely uniform. During reproduction and lactation feeding is most intense and weakens sharply during molt although food intake does not cease altogether, as shown by the data given above. However, after the end of the snowy period, feeding is again intense. As the seals become well fed, the intake again decreases. In any case, the Okhotsk larga at the end of summer is so well fed that its food hunting impulse weakens sharply. It is at August end and early September that the larga loses interest in food in spite of the continuing arrival of salmon at places and the availability of other food. The larga then prefers to rest on the beach rocks or shoals in the interval between high tides. Evidently there do exist some minor interruptions in feeding caused by seasonal massive availability of one or the other food items, i.e., short interruptions in their en masse availability, when the numbers of some species decrease and the migration of others has not yet peaked. The Far Eastern pagophobic (island) seal differs vitally from the larga in its food regime. Being a more settled animal, it relies on various food items available in the region of its habitation which, on the Commander Islands, is restricted to just a mile-long coastal strip (Marakov, 1968). Such a relative stability of habitation of the local populations of island seals is evidently due to the extremely abundant benthic biomass, up to 20-30 kg/m on the coasts of the Commander Islands (Marakov, 1969). Invertebrates play a relatively important role in the food of these Commander Island seals although the seals feed there on fish too, especially the smooth lumpsucker Cyclopterichthys ventricosus, and, when it is no longer available, on sculpins (genus Сойи5) and greenlings (Hexagrammidae); from among the invertebrates, they consume predominantly cephalopod mollusks, crabs, even amphipods, gephyreans, etc. (Barabash-Nikiforov, 1936). Instances were also noticed of the intake of rockfish (Sebastodes sp.) and mysid. In the winter-spring season the Commander Island population survives almost exclusively on invertebrates; these food components, it should be assumed, occupy an important position in different seasons of the year. There are other indications too that smaller fish are consumed by local seals. For example, these seals are not seen in Saranna Bay at the confluence of the Saranna River, which has the most abundant salmon reserves in the Commander Islands, although hundreds of these seals regularly inhabit 259 343 the coasts 10 km away from its estuary, оп Саре Tonk (Marakov, 1967). Neither were seal concentrations observed around the estuaries of other rivers in spite of the availability of salmon in them. The corresponding Kuril populations feed on crustaceans (shrimps), cephalopod mollusks (squids) Pacific ruff, bullheads, and other marine animals of the coastal zone (Velizhanin, 1967). Another list of food items has also been given for these same regions: greenling, rockfish, saffron cod (navaga), goby, walleye pollock, flounder, cephalopod mol- lusks (especially octopuses, more rarely squids) and also diverse types of shrimp (Crangon dalli, Sclerocrangon sp., and Lebeus polaris) (Panina, 1966Ъ). The nearest kin in the American territorial waters of the Bering Sea and in the more southern territorial waters of the USA and Canada, Richard’s seal, also feeds on extremely diverse foods. On the Pribilov Islands, as far as can be judged from the limited data (Lukas, 1899*), it feeds in summer mainly on octopuses (Octopus punctatus) and to a lesser extent on crabs. On the Aleutian Islands (on Amchitka Island), its food mainly consists of greenling and octopuses (Kenyon, 1965). The more southern seal populations, living beyond the Bering Sea limits, consume mainly fish, including cod, herring, salmon, flounder, rockfish (Sebas- todes sp.), goby, and even skate; they also consume lamprey, cephalopod mollusks, and crustaceans (Scheffer, 1928*; Scheffer and Sperry, 1931; Imler and Sarber, 1947; Fisher, 1952). In the season of spawning migra- tions of river smelt the seals feed almost exclusively on this fish but, when available, herring becomes their main food item; likewise salmon, when available, is consumed in significant quantities. Nothing is known about the food of the common seal on the Murman coasts. On the British and Dutch coasts, and generally on the coasts of the North Sea, the food of these seals includes herring, flounder, cod, goby, eel, and occasionally salmon. Shrimps are consumed but less often. The quantum of the daily food intake, according to the data of zoological gardens, averages 5.5-7.5 kg of fish depending on the age of the animals and the nutritional value of the food; the annual intake thus amounts to 1,800 kg (Moore, 1955*). Taking into consideration the fairly prolonged intervals when there is no feeding in some other seasons, this norm cannot be extrapolated to the food ration under natural conditions. At the same time, the experience of feeding seal pups weighing 17.5 kg at the time of capture on the western coast of Kamchatka, and in captivity at Utkin, confirms the phenomenal appetite of these animals. They do not feed regularly or even daily, but in captivity consumed 1.8-2.6 kg per day of different types of salmon, 1.е., roughly 15- 19.5% of the pup’s own body weight. The young seals voraciously consumed various types 260 344 of local salmon: Chinook, masu, pink, chum, silver, arctic, and “Kam- chatka steelhead” (with preference for small live fish). On catching a large fish, the larga kills it first by crushing the head between its teeth while small fish are swallowed whole. Larger fish are torn to bits using the teeth aided by the fore flippers. When satiated, the seal desists from fish pursuit (Chugunkov, 1967). Home range. The distinctly settled population, characteristic of the | Commander and Kuril (island) seals, as already mentioned (p. 335), occu- pies relatively small sections exposed to very little seasonal variation. The Commander Islands are inhabited by individual herds which form extremely close-knit rookeéries at various places and remain almost sta- tionary year round. Their precise boundaries have not been determined and establishing them is not easy since the animals in search of food or impelled by other causes can be dispersed quite widely and encountered almost everywhere on the coast (Marakov, 1966*). The herding tendency of the seals varies sharply in the different seasons while the area covered by their habitation in a given season undergoes wide variation. A herd of 50-100 seals inhabiting an area of a few hundred square meters, entering the water at high tide, is confined within some 3-5 km. But in the winter-spring season the population is “dispersed” in an extensive expanse of drifting ice floes far away from the coasts. This process of animal scattering peaks by spring when the mating pairs are often seen at a distance of several kilometers or many hundreds of meters from each other. The density of largas at the time of whelping in some regions of the Sea of Okhotsk averages 1.2 animals per km? (G.A. Fedoseev). At the end of lactation and mating, they again form groups. Initially these are small and are not as coherent as the much later beach rookeries in which the seals are often disposed shoulder to shoulder. The herd begins to grow only in summer when the animals arrive from every direction and form the final rookeries. The density of such colonies is dependent on the total local population concentration of food items and the nature of the coastline. Daily activity and behavior. Alternation of periods of wakefulness (activity) and sleep (rest) is mainly guided by the need for feeding and at places, especially in the formation period of the beach rookeries, by tidal conditions. The periodicity of daily activity under the influence of the latter factor is very sharply manifest among the larga of the Sea of Okhotsk in the beach rookeries on islets and reefs that are inundated. Right at the commencement of low tide, the animals gather around an exposed shoal in anticipation of rapid food availability. As soon as the first patch of their favorite land opens up, they rush to occupy the site right at the water’s edge. As the water recedes, new animals crawl to 345 the newly exposed sections of the shoal ог reef, forming а large herd т a comparatively short time. Land occupancy and the first residency of animals on exposed land close to one another is accompanied by loud discordant noises, restless movements, and sometimes even scuffles. Ulti- mately the animals become quiet and begin to rest. They lie somnolently even under unfavorable conditions—frost, rain, or snow—being driven from their place only by a high tide (Lun’, 1936). The animals in the beach rookeries sleep very lightly. There are no special guards in the rookery. From time to time, one or the other seal awakens, raises its head, peers about, and on sighting no danger, drops its head and sleeps again (Tikhomirov, 1966). A reverse process occurs with the commencement of high tide as the land becomes flooded. The water steadily drives the animals from the peripheral sections and they quietly move sideways in order to gather again at the former site when the tide recedes. When sleeping or quietly resting seals are disturbed by danger, say the intrusion of man, the entire rookery is instantly enlivened and the animals rush to the water pell-mell like an avalanche. As soon as the danger has passed, they gradually return to their former places on the coast. Opinions on which of the sense organs are better developed in the larga are somewhat contradictory and the matter still not wholly resolved. According to some, sight and smell (Tikhomirov, 1966) and, according to others, sight and hearing (Marakov, 1966*) are the best developed. Whichever, the larga is one of the most cautious seals. It is difficult to discuss the daily regime of the larga in winter because of inadequate data. Before the period of reproduction commences, the animals are evidently quite scattered, far away from the coasts among drifting ice floes, and take advantage of open water pools for respiration. The larga usually does not make air holes in the ice. With the onset of the mating period, the activity of the males naturally intensifies. Since ultimately only one male remains for long alongside the female, and the males and females are numerically equal, animal pairing can be assumed as relatively peaceful. The activity of whelped females is subordinate to the lactation rhythm; in the intervals between suckling her pup the female has sufficient time for rest although she has to ensure the safety of her pup and also feed herself. At the end of the feeding period the adult animals split into groups of some tens each, sleep long periods on a large ice floe or simply rest in a somnolent state. In spring immature animals remain apart from the adults but no information is available on their behavior at this time. 261 346 At the conclusion of molt the larga again becomes highly active, partly because of the preceding period of relative abstinence from food. The herds break up and abandon the ice floe. Further, in the period of intensive feeding, the animals attracted by schools of fish again gather into groups. Now, once again, their activity alternates with hours of rest. The pagophobic (island) seals on the Kuril and Commander Islands are quiet and peace-loving animals with an even more distinct herding tendency than the larga. Their lairs consist mostly of 20-50 animals though some contain even up to a hundred. Quite often the same lair also holds one or more single seals without animosity. Similarly, They exhibit no enmity toward other warm-blooded animals. Scuffles are pos- sible only between the males in the mating season. Affinity for a given site is more pronounced in them than among the larga. While the lat- ter usually inhabit the lower coasts and advance toward the coastline with the receding waterfront during low tide, the island seals can be seen 30-60 m away from the waterline (Velizhanin, 1967). In spite of all their quiescence, however, the screams of the Commander Island seals, similar to those of bitterns, can be heard during the period of lactation; the voice of a female calling her pup in case of danger sounds the same (Marakov, 1966*). Seasonal migrations and transgressions. These aspects have not been adequately studied. Unlike the harp and some other species of seals, the common seal is usualiy regarded as a settled animal, albeit this is not entirely correct. Only the Atlantic common seal and the Pacific island seal can be regarded as fairly settled animals. Contrarily, the larga, except for the not so numerous Kuril population, should be regarded as a migratory animal. At least the Okhotsk and Bering populations are such. Although pagophilic, the larga nevertheless avoids compact ice floes. It therefore abandons coastal regions covered with stable ice floes in winter and returns to them only after the floes break up. The period of its appearance on the coasts depends on the time they are freed of ice floes. In the Sea of Okhotsk, in the Yamsk-Siglansk and Tauisk regions, it approaches the coast for the first time usually at the end of May (Fig. 157). In the Tigil’sk region (Kamchatka) the first, but still very rare, predominantly old, animals are sighted in April. On the contrary, in the Shantar Sea, where the ice floes prevail for a much longer duration, the larga approaches the coast only in July and at some places only in August in some years. Until recently, it was not clear where the largas, especially, the Okhotsk form, spent the winter and early spring months, having abandoned the coast. Some authors assumed that they probably 262 261 347 winter close to the coasts among the drifting ice floes (Freiman, 1936). Others held that some part of the Okhotsk population, perhaps even a significant percentage, winters in the seas of Japan and Kamchatka (S. Naumov, 1941). The second view is supported by the total disappearance of the larga in winter from the Shantar Sea, from Sakhalin Bay, and later even from Tatar Strait. Wintering of the larga in the Sea of Okhotsk in the 1950s and 1960s was demonstrated by direct observations. It was assumed that having left the coastal sections for wintering, the larga localized “in certain regions confined to the sites of their coastal rookeries” (Tikhomirov, 1961). Mapping of the main concentrations of the larga showed that by early spring they are at least distributed in “patches” along the periph- ery of the Sea of Okhotsk, but far away from the coast on the edges of drifting ice floes in the central sections of the basin (Tikhomirov, 1966). In fact, in the Sea of Okhotsk the seals of most local populations spend the winter-spring period quite far from the land but nevertheless in a “traverse” opposite the coastal sections where the beach rookeries are formed in the middle or end of summer. The animals forced to abandon their coastal sections are attracted toward them in winter and, moving away from the shore ice and in general the stationary coastal ice floes in the zone of drifting ice floes, they continue to remain as long as the ice Wy о Гм [п и 2 Fig. 157. Figure showing the distribution of the larga in the Sea of Okhotsk during whelping and the main migratory directions to the molting sites in 1969 (by G.A. Fedoseev). 348 conditions suit them in such places as are closer to the coast abandoned by them. Whether or not the food factor influences the winter migrations of these animals is not known for certain. Thus the largas scattered in winter and early spring among drift- ing ice floes in the Kamchatka belt of the Sea of Okhotsk evidently represent the Kamchatka population. The most active among them are sighted in spring along the coasts in the very first favorable conditions. The larga population remaining on the ice in the northeastern fringe of the Sea of Okhotsk is evidently confined to the coastal rookeries of. the Yamsk-Tauisk region. The distances separating the closest beach rook- eries, according to winter finds of their populations, are at least 150 km in a straight line, which the animals traverse twice a year. Perhaps even more distant migrations also occur. There is a view that the larga population forming innumerable summer-autumn rook- eries on the Shantarsk Islands and around them depart for wintering and for reproduction far southward and winter in the southwestern- most corner of the Sea of Okhotsk, toward the northeastern coast of Hokkaido between La Perouse Strait and the southern part of Sakhalin, on the one hand, and the southern islands of the Kuril range on the other (Tikhomirov, 1961). The autumn-winter courses of the Shantarsk populations to the sites for wintering and reproduction are not known. The spring migrations, however, have been confirmed as running along the eastern coast of Sakhalin (Tikhomirov, 1961, 1966). Thus the large congregations of “whelped larga” noticed in April in the above region in the extreme south of the sea, begin to migrate northward in May along the eastern coast of Sakhalin. In June the largas are concentrated in the region of the northern extremity of Sakhalin and remain there as long as the ice floes prevail. The thawing of ice floes and the disappearance of the seals along with them correspond in time with the arrival of the larga in the Shantarsk region (Tikhomirov, 1961). Thus, in order to reach its summer-autumn sites, the Shantarsk pop- ulation (assuming that the pattern of its migrations is accurate) has to traverse a distance of nearly a thousand kilometers. It has to negotiate the same distance back with the approach of winter. The graphic variants of the larga migrations in the Sea of Okhotsk (Fig. 157) agree to some extent with these accounts. The seasonal migrations of the other populations of the Okhotsk larga are considerably shorter. They are the shortest in the case of some small local groups concentrated in summer оп the northern and southern islands of the Kuril range. These groups of larga lead an almost settled way of life, remaining for more than three-fourths of the year near their 263 349 beach rookeries. They abandon them only at the time of reproduction, which takes place in the neighboring ice floes (Belkin, 1964). Most of the larga population of the Bering Sea undertake migra- tions in the same manner as the Okhotsk larga and are dependent on the very same factors—formation of compact stationary ice floes in the winter-spring months on the coasts and migration to the reproduction sites away from the coasts. These sites represent massive drifting ice floes with a fairly large number of open water pools and shore ice, protected simultaneously from storm waves. Most of the Bering population is dis- posed in the winter-spring months in the eastern part of the sea; the larga from Soviet waters also reaches there. Seasonal migrations on one scale or another exist among all the pop- ulations of the basin of the Sea of Japan. The larga begins to migrate southward from Tatar Strait in November and disappears from there finally in December. It reappears there again in spring only after the thinning of ice floes (Dorofeev, 1936; S. Naumov, 1941). In Peter the Great Gulf the number of larga increases noticeably in autumn and decreases in spring while it abandons these sites almost altogether in summer (Ognev, 1935) and is evidently dispersed widely in the coastal waters to the north and south; some remain in their original sites. The Far Eastern pagophobic populations of the Kuril and Comman- der Islands, eastern Kamchatka, and some other sections of the range not exposed to prolonged winter glaciation behave altogether differ- ently. These seals lead a fairly well-settled way of life and undertake no significant migrations whatsoever from their coastal sections. Such behavioral characteristics are evidently typical of the local groups of pagophobic seals inhabiting the Aleutian and Pribilov Islands, coastal sections of Alaska, British Columbia, and the USA. The Atlantic seal belongs to the same type as its migrations occur as a rule within the confines of a limited expanse inhabited by various populations. These seals migrate under the influence of weather and human persecution, food availability, need for more isolated sites for reproduction, and other factors, as exemplified by the behavior of the Dutch populations (Bem- mel, 1956). Reproduction. The pagophilic (larga) and the pagophobic (island) seals mate at different times. Among the former, this period sets in ear- lier. In the Sea of Okhotsk and the Bering Sea the larga mates soon after whelping. The presence of an adult male hanging around almost each suckling female is the first sign of mating activity. These observa- tions have been supplemented by a study of the genital organs (includ- ing the presence of sperm in the vagina). The total period of mating 350 in the northern part of the Sea of Okhotsk and in the Bering Sea con- tinues for about one month, from April 20 to mid-May; the bulk of the females, however, mate in the period from May 1 to 10. Mating pairs start to form roughly one week or 10 days before whelping and break up by the commencement of the period of molt (Tikhomirov, 1964, 1966).83 In the more southern regions of the Soviet Far East, in the southern parts of the Sea of Okhotsk, and in the Sea of Japan, mating occurs early, corresponding to the much earlier periods of whelping, i.e., in March and April (Tikhomirov, 1966a). The island seals reproducing on the coasts, in the Kuril and Com- mander Islands and on the coasts of eastern Kamchatka, have not been adequately studied in this regard. Mating of the Commander Island pop- ulations occurs at the end of May (Barabash-Nikiforov, 1936) or more probably in June-July (Marakov, 1964*, 1966*); it occurs obviously at May end and in June on the Kuril Islands (Inukai, 1942; Belkin, 1964). Information about the mating season of the seals on the Murman coasts is not available. It is also not available for the population entering the Soviet Baltic waters. In general, however, the European, especially the Scandinavian, populations of the common seal have long been thought to mate in the autumn months: in September (Holmgren, 1865; Lilljeborg, 1874) or even early October (Collett, 1911-1912). The mating of seals on the southern coasts of the North Sea has been placed in August (Havinga, 1933; Moore, 1952*, 1955*). Despite the earlier views followed by sev- eral Soviet authors (Ognev, 1935; Bobrinskii, 1944*; Vinogradov, 1949; and others), the European common seals mate at the end of July and in August (Harrison, 1960). Among the American (eastern Canadian) pop- ulations of the Atlantic common seal, mating occurs not in the autumn, as was thought earlier, but immediately after lactation ceases, i.e., June end or early July (Fisher, 1954). As in the case of other seals, there is no distinct polygamy among common seals albeit the bulls in one way or another claim the right to mate with a particular female. Furious and prolonged scuffles evidently do not occur among them although marks of seizure in the form of 8 In the light of these data it is difficult to agree with the earlier viewpoint (Sleptsov, 1943) that the Okhotsk largas mate late, from June through the first half of August (peak at July end), when most of these seals have already completed molt. S.P. Naumov (1941) places the mating of the Okhotsk larga as late as August-September, restricting it thus to the coastal rookeries in which it is assumed to occur. E.S. Chuzhakina (1955), using this very information, contradicts herself a few lines later by correctly pointing out that mating precedes molt, which occurs from April end to early June. 265 351 scratches and scars are often seen on the skin of the Atlantic as well as the Pacific pagophobic seals (Allen, 1880; Havinga, 1933; Averin, 1948). Among the larga, fights for the possession of a female are evidently few. The literature adduces neither direct evidence on scuffles among males, nor indirect evidence. The skin of the males reveals no injuries inflicted by competing suitors. In general, the period preceding the formation of mating pairs, noticed from the commencement of lactation, proceeds imperceptibly among largas. Direct observations of the act of mating of the larga have also not been reported in the literature. On the coasts of eastern Kamchatka, it was observed in water on July 7, 1942 (Averin, 1948). The region, as also the date of observations, indicate that the mating seals in this case were probably pagophobic island seals. On the coasts of Scotland, eastern England, and Holland, mating of the common seal was invari- ably observed in water (Havinga, 1933; Venables and Venables, 1957; Harrison, 1960). Mating pairs of island seals (the male pursuing the female in water) can be seen on the Commander Island coasts in calm weather in June and July. Both animals loudly whip the water incessantly with the hind flippers, often dive, and on surfacing sniff each other’s snout (Marakov, 1966* ). A study of the genitals and vaginal smears convincingly demonstrated that ovulation of the Bering larga commences even around the 20th of April, with peak mating in the first 10 days of May (at this time, only virgin females and those that have not whelped in the current year mate), while the total duration of the mating season extends from April 20 through May 15; by this last date almost all the females are inseminated (Tikhomirov, 1964, 1966, 1970). These periods apply to a great extent to the Okhotsk larga also. No unanimous opinion was available until recently about the dura- tion of gestation. The assumption of late (autumn) mating and dif- ferences of opinion regarding the existence and duration of a latent period gave rise to controversies in determining the duration of gestation also. It was assumed as seven months (S. Naumov, 1941), nine months (Sleptsov, 1943; Chuzhakina, 1955; Venables and Venables, 1955), close to ten months (Havinga, 1933; Scheffer and Smith, 1944*), or 11 months (Moore, 1952*, and others). At present, a fairly long (two to four months) lag in the implantation of the blastocyst is recognized among the pago- phobic seals (Ph. v. richardi) and largas (Fisher, 1954; Harrison, 1960). Thus, of the nearly 10.5 to 11 months, on average, from the time the female mates up to her parturition, active embryogeny lasts only seven to nine months. 352 Twins have not been reported among largas while they are not alto- gether rare among the European common seals. Thus, of the 12 births recorded from 1933 through 1940 in the Bremerhaufen zoological gar- den, two were twins (Moore, 1955*). Of the 70-80 births occurring annu- ally in nature in these same years on the coasts between Vezer and the Elba, about 10 were twins (Juncker, 1940). In the Far East the larga becomes capable of reproduction at three years of age at the earliest but the majority about one year later. Some animals even after attain- ing maturity lag behind until the fifth and even the sixth year of age (Fedoseev and Shustov, 1964*; Tikhomirov, 1966). Immature females at four years of age constitute 7% and at five years 6%. All males are mature from the fourth year of age (Tikhomirov, 1968). The island seals have not been adequately studied in this respect; it has been assumed that maturation of the Commander seals sets in at the age of two to three years (Marakov, 1966*). The European common seals are capa- ble of reproducing in the third or fourth year of age (Havinga, 1933; Heinroth, 1958). By the time maturity is achieved, the testes of the larga weigh at least 100 g (without appendages), rising to 185 g by the mating season (Tikhomirov, 19664*). No data are available on the time of whelping of the common seal inhabiting the Murman coasts. In the Norwegian waters, however (in the southern as well as northern parts), the seals whelp roughly at nearly the same time, from the first half of June to July 20, predominantly at the end of June (Collett, 1911-1912; Eines, 1964*). Along the southern and western extremities of the North Sea and in the southwestern Baltic Sea, pups begin to be seen in most cases in the second half of June to mid- July though much earlier as well as much later dates of birth are known (Havinga, 1933; Wagner, 1936; Venables and Venables, 1957; Harrison, 1960; Moore, 1955*, 1965*; and others). In the Pacific Ocean part of the range, the pagophobic (island) seals of the more southern populations forming rookeries on the Kuril Islands whelp from mid-May through the second half of July (Belkin, 1964, 1966; Velizhanin, 1967); those on the Commander Islands mainly do so in June (Marakov, 1964). The much earlier dates pointed out for the latter region, 1.е., from April end through early May (Barabash- Nikiforov, 1936), are evidently not wholly accurate or pertain to the pagophilic larga. Largas whelp in a much earlier winter-spring season. Those on the southern boundaries of the range in Peter the Great Gulf whelp right in February (Ognev, 1935; Nikulin, 1937) and even at the end of January (Pikharev, 1948). According to the latest data (Kosygin and Tikhomirov, 266 955 1969), the period of whelping in this region extends “roughly from the middle 10 days of February and includes the first 10 days of March”. In Tatar Strait this period shifts to a much later period. A large number of females give birth there in March and the remainder in April; the total period of whelping, however, extends from mid-February to the second half of April. Instances are known of pups being caught there even on May 19 (Dorofeev, 1936; Nikulin, 1937; S. Naumov, 1941; Pikharev, 1948; Tikhomirov, 1966; O.A. Salmin). In the Sea of Okhotsk, the pups appear earlier in the south, mainly in mid-March, than in the north where whelping in most cases occurs in mid-April (Tikhomirov, 1966). In general, however, this period even in the southern part of the sea is also evidently extended since, in the region of the southern Kuril Islands, newborns were sighted on the ice floes at March end to the first half of April (Belkin, 1964). Newborns were noticed at the earliest in the Bering Sea in 1962 on April 3 (Tikhomirov, 1964) and in 1963 on March 26 (Kosygin, 1966). Until the beginning of the middle 10 days of April, however, pups were very rare and most of the adult females caught at this time were gestating. En masse whelping occurs there from the second week of April and extends roughly to the end of that month, peaking in the middle of the month. Newborns were seen in 1963 up to mid-May (Kosygin, 1966). The solicitude of the mother for her pup in the period of lactation is great; she does not tolerate even the proximity of people or ships, considering them a mortal threat to her offspring. The larga usually whelps every year; in any case, barren females hardly exceed 10%.8* In the period of reproduction the maternal population and the bulls are concentrated mainly in the pelagic strip of drifting ice floes 20 km or less wide (Tikhomirov, 1966b) though in the Bering Sea they whelp even deep among sparse ice masses (Kosygin, 1966a). The larga selects for whelping ice floes that are not particularly large and preferably not too hummocky but firm and clean with open water pools among them. They do not whelp on compact stationary ice floes (in the coastal fast ice) and within intensively broken massive ice. Parturition occurs not far from the edge of the ice floe. These seals do not form concentrated. nurseries and whelp far from each other; but in Tatar Strait scattered groups of females, sometimes with up to 20 to 30 pups in the range of vision, have been observed. In the Sea of Okhotsk and in the Bering Sea they remain more scattered, 81 According to Е.А. Tikhomirov (1966), it is roughly 5% of the eligible females; Gol’tsev and Fedoseev (1970) assume it to be 15% for the Okhotsk population and 8% for the Bering population. 267 354 usually at a distance of at least 0.5 km from each other but sometimes even farther apart. Thus no more than two or three suckling females with pups are seen in 1 km? (usually with an adult male alongside). On the whole, the area of the ice floes thinly populated with animals extends for 150-200 km? (Tikhomirov, 19655*). Even the term “rookery” cannot be applied to such thinly scattered seals. The pagophobic Commander Island seals undergo parturition some- times on the coastal rocks or in sand spits which are sometimes even flooded during high tide so that the pups in such cases are “introduced” to the water immediately after birth (Marakov, 1967). The newborn is quite prepared for this: it is born with a short hair coat, having shed the preceding embryonic fur in the mother’s womb. The island seals, also on the Kuril range and outside the USSR waters on the Aleutian and other islands, undergo parturition under fairly similar conditions. The seals inhabiting the more southern regions of the Pacific Ocean coasts of North America whelp on sandy-pebbly shoals (especially in the estuar- ine sections and the lower courses of rivers) and also on small rocky islets and reefs along the continental coast. The Atlantic seals along the-coasts of the North Sea, in the southwestern Baltic, and in all the other parts of the range, including Soviet western Murman, undergo parturition under nearly similar conditions. Growth, development, and molt. Newborn largas weigh 7.5-8 kg, somewhat more, possibly up to 10 kg in some cases. The body length in a straight line (Lcv) varies from 65 to 80 cm (length measured up to tip of tail along the body curvature varies from 75 to 90 cm and pups already somewhat grown up to 110 cm). They have the typical white coat of dense, long, silky hairs of almost pure white color with creamy tones and resemble the white pups of other pagophilic seals, especially of the harp and ribbon seals.®° Their teeth still sit deep in the alveoli and are covered by the gums. Their milk teeth, however, are extremely reduced and may be imperceptible. The subcutaneous adipose tissue has almost not yet developed. The pups of the Atlantic common seal born on the beach almost do not differ in size and weight from the pups of the larga,®° are devoid of the embryonic coat, and are born almost covered with the short, smooth, coarse hair coat that is characteristic of older animals. However, during 85 The hair coat of the newborn in the region of Peter the Great Gulf is a smoky-gray (Kosygin and Tikhomirov, 1969). 8 The maximum of 15 kg (Havinga, 1933) should be regarded as an exaggeration; evi- dently this pup had already completed suckling. 355 the embryonic growth of this seal and other ecologically similar pagopho- bic seals, i.e., Kuril (island) and Richard’s seals, the embryonic white coat is seen, as in the larga. Just before birth, the hair coat loses strength and most often is shed immediately before parturition or right at the moment of birth, more rarely in the first few hours after birth. Only in extremely rare cases do the pups sport this coat for a few days after birth. Yet many such extraordinary cases have been recorded in the last 20 years (Moore, 1965*; Stutz, 1966). Since the newborn pagophobic seals have no embryonic coat, they can enter water within a few hours of birth. As noted above, such pups may even be born right in water; in such cases the pup nevertheless scurries toward land with the mother’s help. The pup of the common seal has a color similar to that of the adult but with a more distinct monochromatic middorsal strip. Suckling of larga pups continues (as far as can be judged from the average birth periods and from the time at which the female abandons her pup) for about one month. For much of this time the mother remains on the ice floe, never leaving her pup and displaying extraordinary solic- itude. In the Bering Sea lactation continues from April 10 to May 10-15 (Tikhomirov, 1964). In this interval the pup adds 12-20 cm and mea- sures 80-90 cm in a Straight line or 95-107 cm along the body curva- ture (Kosygin, 1966) ог 101.5 -111 cm (Chapskii, 1967). The pup grows rapidly during lactation and its weight increases largely as a result of fat deposition. At the end of suckling the initial weight of the newborn quadruples to 30 kg or more. As this period draws to a close, the embryonic coat weakens, is later gradually shed, and soon replaced by a new smooth, short, and very firm coat with sparse hair. Opinion on the duration of the primary embryonic coat is not unanimous. According to some, it remains strong for roughly two weeks or slightly less (Tikhomirov, 1964) and in the Bering Sea up to April 15; according to others (Kosygin, 1966), only 5-7 days, the pups being fully molted at 15 days of age (Fig. 158). Casting of the embryonic coat proceeds in the very same sequence as in pups of other species of seals; after a brief latent period during which the embryonic coat loses strength and the new coat still covered with short and smooth hairs grows, molting commences and the old embryonic fur is shed in clumps on large sections of the head, flippers, and tail. In this state, given the sharp contrast between the dark-colored molted sites and the still preserved light-colored embryonic fur, the pups appear to sport a white fur vest (Kosygin, 1966). Later, the hairs on the back and on the ventral side of the trunk are shed. Most pups of the larga are completely free of their embryonic fur by mid-May (Tikhomirov, 1964). 268 268 356 Fig. 158. Intensely molting white large pup. Bering Sea (photograph Бу С.М. Kosygin). Only on completion of molt does the pup begin to enter water and adjust to the new environment, remaining in it almost up to autumn. The color of the new hair coat after the embryonic coat has been shed is very Similar to that of the adult but is also variable with near total absence of light-colored streaks along the spine. The total duration of lactation of the Kuril (island) seal is consid- erably more extended: 3.5 months from mid-May through August end (Velizhanin, 1967). This is evidently due to the prolonged whelping sea- son. The individual duration of suckling, though undoubtedly short, is evidently much longer than the corresponding duration in the larga. Many newborns of the island seal originally sport a very deep dark, almost black, coat with diffuse light-colored annular spots. Their body length along the dorsal curvature (Lc) varies from 94 - 104.5 cm and they weigh 19-19.4 kg (Belkin, 1964). There is no information on the increase in these values after lactation. The duration of lactation among the European common seal (British population on the east coast) is thought to be three weeks (Harrison, 1960). For seals on the coasts of Holland, the Federal Republic of Ger- many, and the German Democratic Republic, double this duration has 269 357 been indicated. There is basis for preferring the data of direct obser- vations on the duration of suckling of the animals born in zoological gardens, which indicate a figure of six weeks (Heinroth, 1958). The rapid growth and weight increase of pups of all seals, includ- ing those of the species under consideration, over a comparatively short duration of lactation is explained by the very high nutritional value of the milk. The milk of the common seal has 45% fat and 9% protein (Harrison, 1960). Later, the growth and development of the youngster feeding on its own are characterized by slow tempos. It has been indi- cated (Havinga, 1933) that, after the first year, the Dutch seals grow to 95.7-113 cm (length in a straight line, Lcv); this is only 5-10 cm more than the length achieved by the pup of the larga in less than a month of lactation. Further growth of the larga is illustrated in Table 14 (Tikhomirov, 1968). The molting of immature (commencing from yearlings, Fig. 159) and adult largas is mainly confined to the regions of reproduction and only partly to the beach rookeries where probably some adult animals may complete molt, if they have not completed it on the ice floes. In the Sea of Japan, only one molting region is known for certain in the northern part of Tatar Strait. The molting sites of the larga reproducing in Peter the Great Gulf are not yet completely known. It may be assumed that the animals spend a part of their molting period there itself on the ice floes and complete the molting process on the coastal reefs. In the Sea of Okhotsk molting colonies have been detected in the southwestern region of Shelikhov Gulf (in Yamsk Bay), in the northern part of the Sea of Okhotsk itself, west of Shelikhov Gulf to Tauisk Bay, and on the eastern coast of Sakhalin. The local groups perhaps molt also in the proximity of the western coast of Kamchatka. Opinions vary concerning the westernmost part of the sea, 1.е., Gulf of Sakhalin and the Shantarsk archipelago. According to some (S. Naumov, 1941), there are no genuine molting colonies of the larga in the Gulf of Sakhalin; Table 14. Body length of the larga along the dorsal curvature (Lc), (cm) Female Male Age in years Range Average Range Average 1 105 - 136 123 113-136 125 2 120-143 134 132-150 140 3 137-150 143 142-160 151 4 141-155 149 159-161 160 5 140-152 149 148-165 162 358 270 Е. 159. Under-yearling larga. Bering Island, May, 1969 (photograph by I.P. Tomatov). at the very end of the icy season “only insignificant remnants of molted colonies formed in the other regions” are seen there. In fact, throughout June the larga is mainly confined to northern Sakhalin, or even farther away, toward the Gulf of Sakhalin and more so toward—Shantar Sea [Island], which is blocked by heavy ice floes. Only in the second half of this month does the larga localize in a comparatively restricted space between Cape Litke and Men’shikov Island. This region represents the main residence of the larga in spring, on drifting ice floes all along the western part of the Sea of Okhotsk. The seal penetrates west of this region only from the first half of July, by which time most of the animals have completed molting (Pikharev, 1941). Other authors unreservedly include, in addition to the above regions, not only Sakhalin Bay, but also the Shantar Sea [Islands] among the molting regions of the Okhotsk larga (Nikulin, 1937; Tikhomirov, 1966). In the Bering Sea the main concentrations of molting largas are seen in the same two main regions in which whelping occurs: the southeastern fringe of the ice masses (mainly to the north and northeast of the Pribilov Islands), the Gulf of Anadyr and partly the eastern coast of Kamchatka. Molting largas do not congregate in the Chukchi Sea (Tikhomirov, 1966); there are no accurate data whatsoever on the molting of this seal there. 270 359 The information available in the literature on the period of molting varies widely. In Tatar Strait molting occurs at April end and in May (Dorofeev, 1936) or at the end of May (Yu.A. Salmin). In the Sea of Okhotsk, on the eastern coast of Sakhalin, molting larga juveniles were encountered (in twos and threes) on small broken ice floes at the end of May (Nikulin, 1937) and somewhat farther away from Sakhalin but in the same southwestern part of the sea molting adults were encountered at the beginning, middle, and end of June (Pikharev, 1941); further, in 1939, an adult larga caught on June 17 was still in the initial stage of molt while others caught later (up to July 1) were at the peak of molt. In the northern part of the Sea of Okhotsk the first of the molting largas were found among the molting bearded and ringed seals from May 20 (Freiman, 1936). On the whole, the molting period in the Sea of Okhotsk continues, according to some (Tikhomirov, 1961), from the end of April and, according to others (Fedoseev and Shustov, 1964), from around May 10 to mid-July, i.e., roughly оуега period of 2-2.5 months. | In the Bering Sea most of the adult largas generally molt during the same period: from the middle or end of the second ten-day period of May to the middle or even the end of June. It has been pointed out that males and females (which for some reason have not undergone par- turition) begin to molt 10 to 15 days earlier (it is quite possible that some males molt even during the mating season) compared to females that have whelped (Tikhomirov, 1964). Young animals, however, molt somewhat earlier than the adults, from the last 10 days of April (Kosygin, 1966), although the reference to a much earlier period, 1.е., from the first few days of March (Tikhomirov, 1964), is due to an incorrect understand- ing (Or pertains to the southernmost populations). More recent authors (Gol’tsev and Fedoseev, 1970) have indicated that en masse molting of the larga occurs in June. Information on the molting of land-loving forms is extremely scant. Only general references are available on the European-Atlantic seals which undergo molt in summer (Millais, 1904; Collett, 1911-1912); young seals molt earlier, in July, and older ones in August and even early September (Havinga, 1933). The first signs of molting among the pagophobic island seals inhabiting the Kuril Islands are noticed early in July. At the end of this month and in early August, intensely molting animals were noticed on Makaarushi Island; molted animals were caught in mid-August (Belkin, 1964). On the Pacific coast of Canada and the USA (especially on the coasts of British Columbia and Washington state), molting continues from the first half of August to September end (Fisher, 1952). 271 360 Enemies, diseases, parasites, mortality, competitors, and population dynamics. The common seal in the Baltic section of the range is threat- ened by no natural enemies. One of the most important factors responsi- ble for natural mortality there is stormy weather, which can take a toll of newborns. Some pups may perish due to unfavorable birth conditions, as sometimes happens in zoological gardens. Nothing is known about the natural mortality of the Murman populations of the common seal. Information on the larga in this respect is more specific. Although itself quite aggressive, it is attacked at places by carnivores. Bears attack the larga in many regions on the coasts of the Sea of Okhotsk during the formation of beach rookeries. Sometimes bears even stack their quarry in some ravine and cover it with soil. Largas torn apart by bears have been found time and again along the banks of the Moroshchechnaya River in western Kamchatka and in some bear dens along the northern coast of the Sea of Okhotsk, including Amakhton Bay (S. Naumov, 1933, 1941; Tikhomirov, 1966). There is evidence of wolves attacking the larga. A body with the internal organs and brain eaten out was found on the same Moroshchechnaya River with many wolf tracks around it (Tikhomirov, 1966). Even large predaceous birds can peck a newborn larga to death. An indirect proof of this is the hovering of the golden eagle and the bald eagle around seal rookeries. The Kamchatka residents on the coasts of the Sea of Okhotsk speak of seal fights with eagles. On the ice floes close to Sakhalin, pups of the larga pecked to death by eagles were also seen (Inukai, 1942; Wilke, 1954). A similar instance was noticed in the Bering Sea too on the coast of Alaska (Wilke, 1954). Nevertheless, instances of predaceous birds attacking pups, less so fully grown largas, are not very frequent since the breeding sites of the latter are usually quite far from the coasts; further, the adult animals protect their offspring. The shark could be regarded as a natural enemy of the larga though no concrete data are available for recent years. This factor has been cited for the Atlantic common seal (Sutton and Hamilton, 1932*). At places, the larga is victimized by the killer whale (Orcinus orca). On the Japanese coastal whaling base near Abashiri, seals of this species were the most frequent food item found in the stomach of killer whales caught from August through October, 1948 (Wilke, 1954). Abiotic factors are equally responsible for the mortality of the young. Pups on sparse drifting ice floes among extensive open water pools and more 50 оп being transported to the fringe are exposed to dual fatal factors. In highly windy weather the embryonic fur coat sprayed or even drenched by an icy wave simply cannot protect the pup from cold. A powerful wave can sweep a pup off an ice floe or even topple a floe with 272 361 pups into the water. Ice floes quickly break up under wave action and are transported to more southern regions where they soon disintegrate and thaw. Contrarily, during hummocking of ice floes, pups face the danger of being crushed. Every year, from April end to the first 10 days of May, carcasses of larga pups in white fur coat are scattered on the coast of eastern Kamchatka. Particularly large numbers of them were thrown up from May 1 through 5, 1942. On the coast of Kronotsk sanctuary, in Olga Bay, 17 such dead pups were collected (Averin, 1948). These have been regarded as important food supplements to bears awakening from winter slumber. Insofar as the Pacific pagophobic island seals are concerned, espe- cially those inhabiting the Commander Islands, they have practically no natural enemies except the blue fox, which attacks newborns only as a very rare exception (Marakov, 1966*). There are apparently more ene- mies on the Kuril Islands but practically speaking, neither the bear nor the wild dog, nor even the fox can inflict harm since the rookeries are generally located on small islands or on reefs where the seals are inac- cessible to the carnivores. Our literature contains no definite data on the diseases of this seal. Among the external parasites recorded, the louse Echinophthirius hor- ridus localizes mainly on the back, upper side of the tail, and at the base of the hind flippers, often in large numbers (Freund, 1933; Moore, 1955*). This parasite is a carrier of microfilaria in the blood. Information on the helminth fauna of the Atlantic common seal appeared for the first time in the early nineteenth century (Rudolphi, 1819). Later, the helminth fauna of this seal was studied quite thor- oughly (Monticelli, 1889; Reie, 1899*; Stiles and Hassall, 1899; Linstow, 1905; Ransom, 1920; Baylis and Daubney, 1925; Lyubimov, 1927; Delya- mure, 1955; and others) but the helminth fauna of the Far Eastern larga had almost not been studied until quite recently. Only nine species of helminths of this seal were known by the early 1960s (Belopol’skaya, 1960). Much progress has been made in recent years in identifying the helminth fauna of the larga. In 1966-1967, 152 animals were autopsied for helminthological studies: 116 from the Bering Sea and 36 from the Sea of Okhotsk. As of date (M.V. Yurakhno), 29 species of helminths and 4 of their larval forms have been identified in the Atlantic common seal and larga. From among the trematodes these were found: Orthos- planchnus arcticus (infects the liver, gall bladder, and pancreas; from 1 up to 69 specimens were detected in a single animal), Cryptocotyle lingua, Echinostoma acanthoides, Rossicotrema venustus, and Phocitrema fusiforme (parasites of the intestine), and Pseudamphistomum truncatum 273 362 (liver). From among the cestodes these were found: Diphyllobothrium cor- datum, D. hianus, D. schistochilus, D. latum, Diphyllobothrium sp., Pyram- icocephalus phocarum, Diplogonoporus tetrapterus, D. mutabilis, Trigono- cotyle skrjabini, and Trigonocotyle sp. (all these species of cestodes infect the intestine). The nematodes infecting the gastrointestinal tract are: Anisakis sp., Contracaecum osculatum, Phocascaris phocae, Terranova decipiens, T. azarasi, Terranova sp., and species of genera of Anisaki- dae; Skrjabinaria spirocauda infects the heart, blood vessels, and lungs; Parafilaroides gymnurus, P. krascheninnikovi, and Otostrongylus circum- litus infect only the lungs; the nematode Ph. phocae infects the larga more often than other seals. The acanthocephalans infecting the intes- tine are: Corynosoma strumosum, C. semerme, C. validum, C. hadveni, C. ventronudum, and C. osmeri. The helminth fauna of the Atlantic common seal and the larga vary significantly. There are large differences in the species composition of trematodes and cestodes; there is none common among the 14 species and, further, these groups of helminths are more abundantly represented in the larga. The latter is highly afflicted by helminths, especially in the Bering Sea, where all the animals commencing from yearlings were found infected by hundreds and even thousands of C. strumosum (there were more than 10,000 of them in some individuals) and also by a large num- ber of other helminths, among which the cestode Т. skrjabini was the most numerous (from a few tens to some hundreds) (M.V. Yurakhno). The more commonly infected organs were: small and large intestine (80% of the animals), stomach (72.2%), duodenum (66.9%), and гес- tum (60%). The following organs were more rarely infected: liver and gall bladder (7.8%), lungs (6.9%), heart and large blood vessels (5.3%), and pancreas (0.9%). The most pathogenic helminths of the common seal include S. spirocauda, parasitizing the heart, large blood vessels, and lungs.®’ The population dynamics of seals in our territorial waters and in the seas of the Far East has practically not been studied in recent decades. The degradation (which was not very intense) of some beach rookeries of the larga in the 1930s in the region of Tatar Strait and at some points on the coasts of the Sea of Okhotsk as a result of hunting was minimal and intermittent and hence led to no particularly serious consequences (see pp. 364-366). Hunting from ships in recent years in the Sea of Okhotsk 87 Information on the helminths of the species under study published here was specially prepared for this publication by Prof. S.L. Delyamure and A.S. Skryabin, scientists at the Helminthological Laboratory, Crimean State University. 363 and the Bering Sea (see р. 365) has likewise exerted no perceptible influ- ence on the population of the larga. Only some local populations of pagophobic seals, especially on the Commander Islands, have not so far regained their earlier population level due to uncontrolled hunting. The Kuril seal too possibly shares the same fate. The extremely small pop- ulation of Baltic seals is explained as a result of hunting and the high degree of economic exploitation of the coasts, particularly the marginal sections of the range. The small Murman populations at present have clearly gained the maximum levels. While calculating the reproductive capacities of larga populations, the following basic data should be taken into consideration: (1) the quan- titative proportion of the males and females remaining equal, the mature animals in the populations of the Sea of Okhotsk and the Bering Sea average 55.7%; (2) barrenness among the females is not identical in both populations but is generally not high (8% in the Bering Sea and 15% in the Sea of Okhotsk populations). Thus the annual increment is about 25% of the total population; (3) the mortality of underyearlings is high at an average of 42.5% pups and thus the increment in herd is roughly 14.5%; and (4) a comparison of these indices with the number of pups points to an increment of not less than 10% of the popula- tion. However, by the time the newborns become yearlings, the ratio between the additions and losses is equalized (Gol’tsev and Fedoseev, 1970). Field characteristics. The adult common seal on Murman can be dis- tinguished in external appearance from the ringed seal by its much larger size; slightly longer snout; small, almost black speckles on the skin; and the absence of a fairly broad monochromatically dark longitudinal band along the spine. It differs from the harp seal (gray-spotted animals) in the presence of light-colored ring-shaped patches on the back and the body flanks and the highly vivid dark color of the flanks in general; moreover, the harp seal does not come onto the coast. It is easily distinguished from the gray seal by its shorter snout, absence of a convex profile in the interorbital zone, and presence of light-colored ring-shaped patches on the dark-colored background of the skin. Unlike the bearded seal, the common seal is small and, moreover, spottiness is more sharply manifest and streak-like gaps visible. The Far Eastern pagaphobic seals are recognized by their habi- tat (encountered in the coastal regions which do not freeze as a rule) and their ability to inhabit the elevated sections of the coast at a dis- tance of up to 25-30 m from the waterline (the larga is usually found right in the water), by their size (usually larger than the larga), by the broad somewhat puffed up snout, and by bright spotty coloration; quite 274 364 often, contrasting light-colored, predominantly distinct ringlets without blackish-brown fine uneven speckles are scattered on the monochromat- ically dark-colored background. The adult larga during residence on the ice floes (in spring) is confined in groups consisting of a female with pup and an adult male. Later, in the molting period, the seals form hards of dozens of animals. They differ in coloration from the ribbon seal and the bearded seal in a variegated, spotted pattern on the skin, from the ringed seal in fine speckles scattered randomly on a gray background on which the ringed pattern does not form a continuous grid or lattice, formed by the fusion of ringlets, as in the ringed seal. Moreover, the ringed seal is much smaller than the larga and has a relatively shorter trunk and snout. In the beach rookeries largas are readily recognized by their close disposition, their movements (scuffles are frequent in the rookeries), and mainly from the racket they raise, especially at the moment of seizure, and their screams, which sound like a cacophany of barking dogs and bellowing cows (S. Naumov, 1941). The racket can be heard for distances of 2 km or more in quiet weather. Unlike the larga, the island seal is quiet and almost does not raise its voice in the rookery. Only in the case of danger do the females sig- nal the pups with guttural sounds resembling the “boom” of a bittern (Marakov, 1966*). It has been pointed out that the island seal lying on an isolated boulder jutting out not very high above the water usually lies with the head as well as the hind flippers held high, its silhouette resembling a boat with a high bow and stern (Marakov, 1966*). Fur- ther, the larga can also be seen on ice floes with raised hind flippers. (K.Ch.) Economic Importance The European common seal, being extremely few within out waters, is of negligible commercial importance. This seal is generally not caught at all on Murman. The pagophobic populations of the Kuril or island seals have no commercial prospects since their number is negligible, although the local people use a few of them here and there in Kamchatka and at some other places. The larga commands considerable commercial interest. In fact, it is not the only one among our Far Eastern seals whose reserves, in most regions, are being exploited much below the population abundance. The larga has been practically of no interest to the state shipping industry in the Sea of Okhotsk from the early 1930s. It occupies the last and negligi- bly small position: in the Sea of Okhotsk in 1937 to 1939 its proportion 275 365 varied from 4.8 to 8.0% (average 6.5%) of the total catch; the bulk of the catch in these years (as also in the 1960s and 1970s) comprised the ringed seal, Pacific bearded seal, and the ribbon seal. Over two-thirds of the catch of larga came in the summer-autumn hunting season in the beach rookeries of Gulf of Sakhalin, Shantarsk Islands, and partly of Tatar Strait where 500 to 1,000 animals were caught annually. State hunting in the beach rookeries commenced in 1934, was quite irregular, and con- tinued only up to the end of the 1930s in spite of the larga being readily accessible. Thus, in 1934, some 1,500 animals were caught in 22 days (Tikhomirov, 1966). The catch from 1937 to ‘1939 was high although aparently uneven. Hunting was not restored in subsequent years. In spring, the main season of hunting using ships among drifting ice floes in the western regions of the Sea of Okhotsk, the larga accounted for 1.3 to 2.3% (average 1.8%) of the total catch from 1937-1939 (Pikharev, 1941). The reason for this low representation of the larga lay in the characteristic dislocations of its major concentrations on the ice floes in the Sea of Okhotsk as also in its behavioral characteristics. In the early spring the whelped and lactating mothers with pups as also the adult males are concentrated in the Sea of Okhotsk among compact frozen ice floes that are difficult for ships to penetrate. Later, however, when the ice floes become thin and the reproduction period has concluded, the largas gather in the molting rookeries, which are unfavorable for hunting for the sole reason that it is difficult to approach the cautious animals: at the very first rifle shot all the animals disappear into the water where they are inaccessible to the hunters. In the early postwar years hunting of larga in the Sea of Okhotsk intensified perceptibly as a result of enlarging the fleet and the region of its activity. Thus from 1954-1958, the annual catch went up to 3,100 animals, or 4.3% of the total during intense seal hunting. In the next spurt of activity the specific proportion of the larga rose further: from 1959 - 1963, an average of up to 6,000 animals was caught (Fedoseev, 1966) or 6.8% of all the seals killed in these years. From the very beginning of the 1960s, hunting by means of ships occurred in the Bering Sea too, although the larga was caught there in the smallest numbers compared with other seal species: in the early years its proportion in the total catch was only 2.1% in spite of the fact that its population was sufficiently large to support a higher level of kill and to reduce the pressure on the other seal species. In the early half of the 1960s, the total catch of the larga in the entire Far Eastern basin, including hunting by the local people, went up 10 10,000- 15,000 animals. Its proportion in the seals killed by hunting 276 366 using ships did not, however, exceed 9.4% in the best years (Tikhomirov, 1966a). Some intensification of larga hunting with reduced killing every- where of the Pacific bearded seal as well as the ribbon seal and partly the ringed seal is one of the possible methods of rational utilization of the seal resources using ships in the Far East. Intense hunting of the larga would evidently reduce the damage caused by these seals to the salmon reserves. As already pointed out (p. 339), the larga consumes and dam- ages quite a large quantity of Siberian salmon, humpbacked salmon, and other even more valuable types of salmon, which serve as food for the larga in summer. The largas gather in large herds in the fore-estuarine sections, directly in the estuaries, and right in the lower courses of rivers during the period of arrival of salmon for spawning and do not so much consume them as damage them (especially when the fish are abundant), by selectively nipping small bits of flesh along the spine. This adverse role of the larga cannot be ignored, at least at places where fish catching, preparation, and processing have been organized. The Canadian gov- ernment offers a handsome reward for killing even one seal (Phoca v. richardi). In the most important salmon rivers of British Columbia, over 12,000 seals were destroyed over a 10-year period, from 1939 through 1948. From 1941/42 through 1946 alone, some $32,000 were spent for the destruction of 10,000 seals (Fisher, 1952). The loss inflicted by the seal to the fishing industry in Alaska, especially in the Mednaya River region, is nearly 2-3% of the total salmon catch here (Imler and Sarber, 1947). In the spawning rivers of British Columbia during the arrival of salmon, these fish constitute about 30% of the food of seals (Spalding, 1964). The technique of hunting using ships is generally as follows. Motor schooners capable of negotiating among ice floes set out with several motorboats on board, which are dropped in the water in the region of hunting as soon as a sufficiently large number of animals is sighted. The ship heaves to and each boat (with a crew of three in white masks) is assigned a particular direction. The ship maintains radio contact with the motorboats. The teams must not only not lose track of the sighted animals, but also attempt to approach them within rifle range while maneuvering among the ice floes. Depending on the situation, the boats move far away from the ship for several hours or even the whole day. The skin with blubber is recovered from every killed animal while the rest of the carcass (skeleton with musculature) is only carried if space permits. During lactation close approach to the larga depends exclusively on ice conditions. If the ice is dense and the motorboats cannot negoti- ate, the ship approaches the animals. The larga can be shot directly 367 from the ship or experienced hunters climbing overboard and jump- ing from one floe to another can approach the animals singly. Instinct holds the suckling mother close to her pup in spite of fright. The adult male in the proximity of the female with a pup is usually the prized trophy of the hunters. Killing of isolated young (immature) larga dis- persed on the ice floe poses no problem. The situation changes, how- ever, at the end of the lactation period. Then hunting becomes quite difficult. Until comparatively recently, the larga was killed even in the beach rookeries. For this purpose, 15 to 20 hunters in two boats dropped from a ship in the vicinity of a rookery would set out for it taking care not to frighten the animals. Landing away from the rookery and armed with clubs, they would approach the animals as closely as possible by crawling under cover, ready to attack at a signal from the leader. This method was not always successful but often yieled over a hundred animals. In a suc- cessful hunt, from the viewpoint of the hunters, 5 - 10% of the animals in the rookery could be killed thusly, or 500- 1,000 animals throughout the autumn hunting season (Pikharev, 1941). But not all autumn hunts for the larga in rookeries were successful (sometimes the frightened animals escaped before the hunters could reach them); moreover, the killed ani- mals had to be dressed before high tide as otherwise the waves carried it away and recovery was difficult. On the open coasts, where it is practically impossible to approach the rookery, nets are more commonly used to catch the animals. With the onset of darkness, one group of hunters in a boat carrying a sweepnet comes within 50-75 m of the rookery and carefully spreads the net over this area. An auxiliary group of hunters, waiting in another boat at some distance from the rookery, then approaches the net-covered site and both parties draw the ends of the net toward the coast. Between 200-250 seals were caught in one such net in the estuary of the Moroshchechnaya River (western Kamchatka) (Tikhomirov, 1966a). Typical methods of killing the larga in the autumn rookeries are practiced in Tauisk Bay by the Orokhets. On the open rocky coasts the hunters make a preliminary hideout near the rookery, using for this pur- pose a boat priorly cast off near the site. As soon as the seals have become habituated to the hideout, the hunters, taking advantage of high tide, when the animals abandon the rookery, gather in the hideout and wait for the animals to return. Allowing some time for the animals to settle down, the hunters then pounce with clubs and quickly kill as many seals running panic-stricken into the water as possible (Tikhomirov, 1966). 277 368 The Sakhalin Gilyaks have long adopted a very unique but highly ineffective method, using a boat and a long (4.0-4.5 m) flexible pole to which a large ski-like float with a harpoon is attached. Dropping this equipment in the water, the hunter attempts to take it to the closest diving seal and, when successful, pierces the animal with a powerful thrust of the harpoon (Nikol’skii, 1889). In various sections of the vast Far Eastern coast, other hunting meth- ods are known but none are specific for killing the larga. Two methods are very common and extensively used. One is practiced from the end of summer and in autumn. The hunter crawls up to the seal from the coast, shoots it, and rapidly drops a light boat to catch the wounded ani- mal; quite often, however, the seal drowns. In spring the hunters from the coastal villages (mostly from the Bering Sea area) come out in large motorboats and seek the animal among thin coastal ice in quite the same way as done by the hunting parties from a ship in the more pelagic regions. The skin of the newborn larga (white pup) up to five days of age does not command the same high price as that of a harp seal (greenish- white pup) or some other seal born on the ice. The skin of the larga can be used in collars, caps, and other fur goods in a natural state or dyed. However, the number of such skins produced in the early spring hunting season is negligible. Given the prevailing hunting conditions, especially in the Bering Sea, it is difficult to say whether the hunters in one ship can catch over 200 undamaged skins of white pups of this species during the season. The skins of molted pups (killed somewhat later) and juveniles (one to two-three years of age) with a more beautiful, brightly spotted color and relatively thin skin are of much greater value. Because of these qualities and also because of the relative rarity of such skins, the larga has long received particular attention from the coastal people, especially the Chukchis, Koryaks, and Eskimos. They use these skins for making dresses and generally for more delicate work, in particular for fashioning women’s apparel. Until quite recently (1960s), the prevailing fashion for seal skin arti- cles such as caps, jackets, cloaks, shoes, etc. put the seal skins, especially of young ones, in high demand in the fur market. The fashion for short, rigid furs of natural color necessitated catching not the white pups, but the molted pups and the older juveniles. This is all the more rational as such skins are more accessible and hunting is not restricted to the brief spell during which the pups sport the neonatal embryonic fur. Even the skins of the adult larga are perhaps suitable for this purpose through 278 369 their hair coat is slightly sparser and more rigid, the color less attractive, and the hide considerably thicker and heavier. The quantity and thickness of the subcutaneous adipose tissue varies according to the season and the age of the animal. Newborns are almost devoid of it but accumulate a 3-4 cm thick fat layer (together with the skin) in the short lactation period and weigh roughly 15 kg (also with the skin). The skin with the subcutaneous fat (blubber) of animals in a transitional age weighs 15-25 kg. The thickness of the fat layer in adult animals in spring differs little from that of the immature animals but exhibits sharp seasonal changes. The weight of the blubber of the larga in spring (especially after lactation and mating) averages 20 kg; it doubles to 40 kg by autumn (Fedoseev and Shustov, 1964*). The meat of the larga is used as required by the trade. The demand for it has been increasing year by year with the expanding animal farms, particularly in the local coastal collective farms of the Far East. The meat with bones of a young larga weigh 15-20 kg and of semi-adult and adult animals 35-40 kg. Rules governing the utilization of the Far Eastern marine animal resources have not been properly drawn up to date. This applies to the larga also, although there is no need for special controls on its hunting. However, the need to reorganize the hunting activity is quite consider- able. Any seal providing fur (and hence the larga too) deserves the same attention as the fur seals. There is also need for special supervision over the utilization and study of the less abundant island (pagophobic) seals on the Kuril Islands, Commander Islands, and along the eastern coast of Kamchatka, all the more since these seals, as far as is known, do not affect the fishing industry. (K.Ch.) Subgenus of Harp or Greenland Seals Subgenus Pagophilus Gray, 1844 HARP OR GREENLAND SEAL Phoca (Pagophilus) groenlandica Erxleben, 1777 1777. Phoca groenlandica. Erxleben. Syst. Reg. Anim., p. 588. Greenland. 1778. Phoca oceanica. Lepechin. Acta Academ. Petropol., I, p. 259, Tables 6 and 7. White Sea. 1785. Phoca semilunaris. Boddaert. Elen. Anim., р. 170. Greenland, !ce- land. 1811. Phoca dorsata. Pallas. Zoogr. Rosso-Asiatica, I, р. 112. White Sea. (V.H.) 279 370 Diagnosis These are relatively large seals. The body length of the adult reaches 2 m (even more in some cases) up to tip of tail along the dorsal sur- face (Lc). The adult skull is 185-240 mm long. The color of the hair coat varies (depending on the age and to some extent on the sex) from spotted gray (with haphazard but not very densely scattered dark, mostly angular patches in a gray background) to a bright contrasting “wing pat- tern” with two very large highly elongated dark-colored, almost black patches, sharply prominent in the light-colored background. This “wing pattern” is disposed symmetrically on both sides and the anterior tips converge on the back. In the final dress of the animals the color of the anterior portion of the head is the same as that of the “wing pat- tern”. The skull is quite massive, with thick bones. The upper and lower processes of the posterior edge of the zygomatic bones are nearly equal in length; the bony nasal septum in the choanae reaches the posterior edge of the bony palate. The palate has no significant notches but a central prominence, turned backward, is usually seen. The bony lobe of the external auditory meatus is genuflexed forward. Molars and premolars (except the first) have two roots while the crowns, especially of the lower jaw, bear well-developed accessory cusps. (K.Ch.) Fig. 160. A—male harp seal, Pagophilus groenlandica, with the final color of the “wing pattern”; B—adult female harp seal in the “semiwinged” phase of color (incomplete pattern of the “wings” covered in sparse spots) (figure by N.N. Kondakov). 279 371 Description The external appearance is typical of seals of the genus Phoca but the adult animals are perceptibly larger than their counterparts in other species of this genus. The body is almost perfectly streamlined and its hydrodynamic characteristics improve with growth (Alekseev, 1966). The claws on the fore flippers are quite massive, blackish-gray, with a distinct transverse rib formation among adults; on the hind flippers these age- related bands or segments are less prominent or distinct. The length of the digits on the fore flippers decreases successively from the first and the second (which are nearly equal in length) to the fifth. The whiskers are dark gray, flattened, with wavy edges. The labial whiskers are usually disposed in 7 rows; each side of the lowermost row most often has 7 (6 to 9) whiskers, the second to the fourth 9-10 (8 to 11), the fifth 4-9, the sixth an average of 4, and the last an average of 2. Sometimes a lone whisker is seen above the seventh row. The labial whiskers total 46-47 on each side. The supraorbital whiskers number most often 3 each but vary generally from 1 to 4. As a rule, there are two whiskers around the nostrils on each side (Yablokov and Klevezal’, 1964). Two types of hair coat are common among pagophilous seals. These are successively the neonatal (juvenile, white) and the definitive form acquired on shedding it. The first consists of tender wavy fur hairs set very densely in tufts to form luxuriant and concomitantly dense and long creamy-white fur. The definitive hair coat of the harp seal in structure, strength, and notable slant (pile) is the same as in the other seals. It con- sists of uniformly distributed tufts, usually comprising three categories of hairs: guard hair (12.5 mm long), intermediary 1-2 (6.2 mm long), and 4-5 fine and wavy fur hairs (5.5-7.7 mm long). These are disposed in a definite sequence; intermediary hairs anterior to the fur hairs and the guard hairs shifted forward even more, covering the rest of the tuft constituents. A 1 cm? area has 1,700 such tufts with a total number of about 12,000 hairs (Bel’kovich, 1964). The color of the hair coat varies greatly with age (Fig. 161) and the final wing pattern is highly typical and contrasting: one large, long, dis- tinctly contoured, vivid brownish-black patch each on the right and left sides of the trunk on a very light, white, almost pure white, or slightly silvery background. The anterior tips of the patches fuse roughly in the zone of the scapula on the dorsum and with a slight divergence extend backward and downward along the flanks; the patches enlarge percep- tibly in this process with a slight crescent-like form in the midportion. Later, they gradually narrow and disappear in the sacral zone near the 281 372 Fig. 161. Main changes in age-related skin patterns among harp seals. Top to bot- tom: gray form (1-3 years old); transitional phase of female (average 3-5 years old); and final coloration, 1.е., the winged form (5 years and above) (photograph by R.Sh. Khuzin). 280 ‚ 373 base of the tail. On a flat skin they resemble a typical horseshoe ог are somewhat lyre-shaped. The head almost up to the neck, ear open- ings, and up to the anterior front is the same color as the “wing pat- tern”. Such a coloration is acquired by successive transformation which proceeds slightly differently among males and females. The color of the first definitive hair coat (juvenile) acquired on shedding the white coat (neonatal) consists of a light ash-gray main background (quite often, slightly darker on the upper side) and usually a few dark gray or brownish-black spots. These spots vary widely in number, are randomly scattered, often in small groups with small and very large ones intermixed, sometimes with fairly angular outlines. This is the color of the young seal (one to three years). The white coat at birth (neonatal) in males and females is altogether identical (see above). The infantile coloration succeeding it on shedding of the white coat is also typical with no specific differences between males and females. The transitional spotted gray color is sported, almost without change, for several years in spite of annual molt. It shortest life occurs in males: less than half the generation has it for five years and not more than a quarter for six years. As far as is known, older males with a spotted gray coloration do not exist. Females, howeger, are seen much longer in this infantile coloration, 1.е., up to 10 or more years of age; even at eight years, their proportion can be 50% (Khuzin, 1964). Stray spotted gray seals are encountered from time to time even among 14- or even 17- or 20-year-olds (Potelov and Mikhnevich, 1967). The next transitional phase of coloration is the formation of an incomplete “wing pattern”. It reveals a mixture of the original features, 1.е., spotted gray (or grayish) and the final “wing рацегп”.8 The life of this stage too varies in males and females. Only the very beginning of this phase is generally similar in both sexes. Very indistinct, highly diffuse darkening is seen initially at the site of the future “wing patterns”. Their coloration is most vivid on the upper (internal, or dorsal) edge which is very distinct and sharply demarcated from the gray background. The “wing pattern” rudiments turn increas- ingly pale toward the lower lateral edges as though faded and gradually merge with the light gray main background of the body flanks and under- side. This shadow of the wing pattern is covered at places by stray, rare, 88 Our special literature used to refer to such 20-year-old females as the “second phase, with gray wings” (М. Smirnov, 1927) but this phrase was later replaced by the term “‘semi- winged,” first applied to males in which the pattern was not fully formed (Chapskii, 1952). 281 374 small spots that are new and bright as well as the remnants of infantile coloration. After the next molt, the “wing pattern” appears more dis- tinctly in some while the features and the color vividness vary little in others and, as before, remain more distinct only along the upper edge and are covered with spots. At this stage, sex-related differences appear in the very nature as also the tempo of color variations. In the semiwinged phase with an incom- pletely formed “wing pattern,” the color transformation process is usually quite delayed among females. This age-color group commencing from the first five or six years (although relatively few in these generations®’) constitutes in succeeding years a perceptible proportion of almost up to two-thirds of the generation. Even among 17-year-old females, the number of semiwinged animals is obviously quite large. It is quite pos- sible that a significant number of females never attain the final “wing pattern” phase but remain in the “semiwinged” coat to the end of their life (Khuzin, 1964; Potelov and Mikhnevich, 1967). Most females never- theless reach the final color earliest at 6-7 years of age (roughly 10% each of the animals in these age groups). At 10 to 17 years of age the number of females with a “wing pattern” increases perceptibly to around two-thirds of the generation. Detailed descriptions are not available of the successive color gra- dations on transition from the semiwinged to the winged pattern. It can only be assumed that this transition is quite simple and sharp since the first of the youngest females with a “wing pattern” are just a year older than the first of the animals with a semiwinged pattern. It therefore appears that some definite quantitative proportions of females of all coloration phases exist simultaneously in the six- to 14- year-old generations although the representation of the spotted gray variety is the least, semiwinged more (one-fourth to one-third), and the winged variety about two-thirds.” The transitional coloration among males is characterized by some specific features. This transition covers only three groups of ages from five to seven years while seals with the full “wing pattern” are also seen in these groups. It may therefore be assumed that the transition from 89 According to some authors (Khuzin, 1964), the females acquire the semiwinged pattern coloration at six years of age; according to others (Potelov and Mikhnevich, 1967), however, this type of coloration is seen among 10% of even three-year-olds. % Some differences are noticed in these proportions among various populations. The White Sea females preserve the infantile (spotted gray) coloration only up to 10 years of age and the semiwinged pattern up to 16 years; the corresponding ages among the Jan Mayen females are 15 and 22 years; a similar lag of color transformation is also characteristic of the Newfoundland females (R.Sh. Khuzin). 282 575 the spotted gray to the final phase is extremely short among males and often the intermediate semiwinged phase is bypassed. Nevertheiess, the latter phase in turn can be subdivided into a few stages (Chapskii, 1967). The first stage is the very dark “wingless” type: the dark spots are so densely scattered that they cover the main background almost wholly so that the “wing pattern” is not visible; such animals look like a black, silhouette from a distance.?! Animals with a faintly identifiable “wing pattern,” lusterless like a dim shadow, can be seen in this group. The ages of the males sporting both color variations are wholly identical: roughly 40-43% each of four- and five-year-olds and 15-20% of six-year-olds. The second stage, forms with a “semiwinged pattern,” is distin- guished by fully developed contours of the lower edge of the “wing pattern” and considerably lighter but densely spotted main background (except for a dorsal, very dark clearance between the “wing patterns”) and even more vividly colored snout (Fig. 161). The age of such males is predominantly six years but even five-year-olds (possibly even four-year- olds) are encountered in this stage. The third stage of coloration, spotted “wing pattern,” is the final phase with a sharply distinct posterior boundary in the dark coloration of the head but still with quite a large number of spots or dabs which are generally dull and scattered in the bright main background. Such males are close to the forms with a “semiwinged pattern” in age: more than one-half of them are six-year-olds, about one-third seven-year-olds, while the five-year-olds constitute roughly one-seventh. Thus the transition from infantile to final adult coloration among males occurs over three years (from the fourth through the sixth inclu- sive). All the males do not necessarily undergo the intermediate stages of coloration. The majority enter the “winged” phase, either totally bypass- ing the intermediate phases or passing through only one or two of the later stages. Thus the fourth year is the last year of the infantile period in which all the males still sport the spotted gray coloration (or gray animals), and the fifth year is the first year in which one-fifth or one- third of them acquire the final wing coloration. All the males develop the wing pattern by the eighth year (Potelov and Mikhnevich, 1964, 1967; R.Sh. Khuzin; Chapskii, 1967). The maximum range of individual color variation is noticed in the period of growth and formation of final coloration. It is high among ani- mals preserving the infantile type of coloration as also among mature females of a much younger age and partly also among males in the transitional “semiwinged” phase. Among the younger animals in the 91 White Sea hunters call them “ogar,” “salovar,” etc. 376 population bearing a spotted gray skin, it is difficult to find animals in which the disposition, form, density, and even the color of the spots, and partly the main background are totally identical. There are very light- colored animals, sometimes with dense variegated spots and sometimes with extremely few spots; there are several variations between these types. Among the females with a prominent “wing pattern,” their contours, superimposition with stray spots, and pattern of spots vary markedly. Among the males in the transitional phase to the final form of “winged” type of coloration, all possible variants of dark aberrations and spot patterns are encountered. The skull (Fig. 162), seen from above, is somewhat similar to that of the large larga (Phoca vitulina largha) but the zygomatic arches do not protrude markedly into the sides: the width at the zygoma in adults is only slightly more than that of the cranium measured” between the mastoid processes. It constitutes 90-110%* (x = 102%) of the mastoid width We ye uy lg ( Ky". er: My, Wess : Fig. 162. Skull of the Greenland (harp) seal, Phoca (Pagophilus) groenlandica (figure by N.N. Kondakov). % The craniometric data are from the author’s materials with some corrections by R.Sh. Khuzin (1967a*) marked with and asterisk. 283 ЭЙ or about 55.5% of the condylobasal length. The interorbital constriction varies from 9.5- 13% of the mastoid width. The width of the rostrum (at the level of the upper canines) varies from 23-36% (x = 29%)*. The length of the facial portion (up to the uncinate processes of the pterygoid bone) is roughly one-half (x not over 52%) of the condylobasal length. The basic skull features of the species mainly occur in the basal side of the axis of the skull and also in the structure of the lower jaw and the zygomatic bones. The midportions of the posterior edge of the bony palate are shifted relatively far back and are fairly at level with their lateral sections without forming a distinct palatal notch; a reverse situation is also seen quite often when, at the end of the median suture, a perceptible inverse projection is formed. This formation is promoted by the longitudinal bony septum in the choanae reaching the posterior edge of the bony palate and quite often extending slightly beyond. In such cases the rear section of the palate is in the form of two highly flattened fused arches. The zygomatic process of the temporal bone is turned forward and not perceptibly enlarged. The choanae are low (height roughly one- half their width). The tympanic bullae are roomy, with quite complex contours, and rounded-triangular in the horizontal plane. The angularity intensifies somewhat with age. Their width (together with the lobe of the audi- tory meatus) is greater than their length and averages slightly less than 20% of the condylobasal length. The bony lobe of the external audi- tory meatus is massive and genuflexed forward. The jugular processes in most cases project above the skull surface and are bent backward. The uncinate processes are not bent outward. The nasal bones, consti- tuting about one-fifth of the total length of the skull, are wedge-shaped with a highly enlarged anterior tridentate margin. The nasal processes of the maxillary bones extend along the nasal bones usually for not less than one-fifth the length of the latter. The sagittal crest is almost absent while the occipital crest is not strongly pronounced. The sub- condylar process is well developed in the lower jaw and is perceptibly incurved. The molars and three posterior premolars and the molar on the lower jaw bear well-developed accessory cusps spread fanlike on both sides of the main cusp (one each in front and, in most individuals, two each at the back). In the corresponding teeth of the upper jaw, the anterior accessory cusp is poorly developed (altogether absent in most premolars but quite often preserved in the molar). Skull differences between males and females are not striking but become distinct on statistical processing of the data. The upper contour 284 378 of the skull profile in the frontal portion among males is in the form of a perceptibly curved line; this curvature is far less noticeable among females.” А paired fold on the lateral fronto-sincipital surface from where the temporal muscles originates is more intensely manifest among the males. The bony nasal septum in the choanae is more developed among males while it is often incomplete (with a notch on the posterior margin) among females. The anterior margin of the nasal bones in males has very long lateral teeth while the median teeth are more often the longest among females. The width of the rostrum at the level of the canines in males is 26-33.5% (M 30%) of the width at the mastoid and in the females 23-32% (M 27%). The width at the zygomatic arches quite often slightly exceeds that at the mastoids in males but not in females. Statistical processing of the data revealed that these and several Other craniological values are somewhat higher in males than among females. In general, the skull of the male is perceptibly more massive and heavier. Age-related variability of the skull is very high. Along with an increase in its dimensions and weight, an extremely distinct age-related feature is the development of its crests and the general surface relief of the cranium. Among newborns, its dorsal surface has an almost wholly smooth fronto-sincipital obtuse area angularly turned toward the nose bridge. The sides of this “angle” from where the paired temporal muscles emerge are even less prominent. With age, they become increasingly contoured and are displaced toward the midline. As a result, the anterior width of the area steadily decreases and the anterior angle becomes increasingly acute while the marginal folds formed are transformed almost into a crest and come increasingly closer. Concomitantly, the very faint rugosity observed along the lambdoid suture only among the newborn, develops into a sharp crest with an overhung pointed dentate cornice in the adult. The age-related changes of other craniological characteristics are less striking but are readily discernible in statistical processing of the data. The most significant are the following:** (1) the posterior margin of the bony palate in the newborn usually has slightly more concave contours, sometimes with a mark on the notch, the latter filling up dur- ing growth and even transforming into a reverse fold in the midportion; (2) the base alone of the bony nasal septum in the choanae in most young animals reaches (not invariably though) the posterior margin of 33 This is not a characteristic of the race (М. Smirnov, 1929) but is of sex-related sec- ondary importance (Plekhanov, 1932*; Chapskii, 1952; and others). % Data pertain to the White Sea population. 285 379 the dony palate; hence, there is an angular notch along its height. In adult and older animals, however, it is not only filled up but its upper portion even emerges beyond the margin of the bony palate; (3) the subcondylar process of the lower jaw is fully developed (i.e., reaches the vertical of the condylar process), also only in the adult; (4) the tympanic bullae in young animals are relatively more swollen, with a smoother, round surface, and more simply contoured than in adults. Their posi- tion is more firmly fixed with age; in the young they may be disposed anterior to the rear crest of the articular fossa while they are invariably posterior to the latter in the adult; (5) the flexure of the upper pro- file in the zone of the nose bridge becomes apparent with age but is absent in the newborn; (6) the premolars and molars which have already emerged from the alveoli are disposed in young animals almost with- Out gaps, their edges often touching each other and sometimes even slightly extending beyond the crowns one behind the other. With age, gaps arise between them as a result of the elongation of the jaws and become widest by the time total maturity sets in. Tooth wearing becomes perceptible only in very old animals; in males and females with the “wing pattern,” the accessory cusps of the crowns become worn to a variable extent; in much older animals, even the main cusps are highly worn down but the crowns are not usually completely worn down. The cusp and partly the side of the canines are worn but fully worn down canines are almost never found. The incisors in adults are also quite often highly worn down; moreover, with increasing age, some straight- ening of their upper row is noticed (the incisors are set anterior to the margin in an even file while they are arranged in a slightly semicir- cular fashion in the young); and (7) the apex of the nasal bones (the depth of their wedging into the frontals) changes with advancing age in the following manner, as a percentage of the total length of the nasal bones: Mean Wi patozone months ni) yaa beh emia tee tae ee ae 41.5 (n = 28) Fromvone tosthree уваги eo at oor 43 (n = 20) Males with “semiwinged pattern” (five to six years old) 44 (n = 12) Males with “winged pattern” (adults) ................ 44.5 (n = 13) The skull proportions too vary with age, especially the ratio between total skull length and width of the cerebral portion (the skull becomes increasingly longitudinal). The width of the skull in the region of the zygomatic arches increases Steadily (toward the time of attaining maturity, it is comparable to and even Slightly exceeds the mastoid width). The rostral width (level of the 380 canines) expressed as an average percentage of the mastoid width reveals the following course of changes: % Age up to one month ......... 227 One-year-olds ................ 24.5 Two- and three-year-olds ..... 26.2 Mature animals .............. 28.6 In young animals the rostral portion is not only noticeably narrower, but also considerably shorter and lower than in adults. The cranium reveals a reverse tendency (in the young it is relatively more swollen; the length of the tympanic bullae as a percentage of condylobasal length decreases from the first year to old age by 4%), etc. The range of individual variation in craniological features is quite significant even among such animals as reveal systematic changes sexwise. Thus the index of rostral width which points to a harmonious increase from one age level to another in animals of nearly the same age (for example, in males with the “winged pattern’) varies from 26 to 34 while the index of the interorbital constriction varies from 10.5 to 16.5, index of the width at the zygoma from 98 to 109.5, etc. Skull elements such as length of the nasal bones, shape of their anterior notch, contours of the posterior margin of the bony palate, etc. also vary. The body length of the adult, fully grown. White Sea males measured between tip of nose and tip of tail in a straight line (Lev) is 155-188 cm. However, the length measured along the dorsal surface (Lc) is 169-205 cm. In the White Sea adult females, Гсу varies from 153-188 cm and Lc from 167-202 ст? (Khuzin, 1963; M.Ya. Yakovenko). The length of the os penis in the adults averages 160 mm. The total weight of the White Sea well-fed adult can reach a maxi- mum of 164 kg and that of the skin with blubber removed by the com- mercial method up to 69 kg (Yu.I. Nazarenko). Weight of heart 600-960 g; liver 1,500-2,400 g; total length of the intestine at 1,900-2,870 cm exceeds the body length of the adult 13.6 times (Yablokov, 1963). The condylobasal length of the skull of adult males (37 animals from the White Sea) was 199.8 - 235.8 mm (х = 217.0); zygomatic width 107.0- 142.0 mm; mastoid width 113.0- 133.3 mm (x = 122.4); width of 9° 229 cm in an extraordinary case. 286 381 the snout above the canines 30.0 -43.5 mm (х = 36.2); and the smallest interorbital width 8.8-21.0 mm ( = 15.0). The condylobasal length of the skull of adult females (27 animals) from the White Sea was 193.0-223.2 mm (x = 210.6); zygomatic width 105.3 - 130.3 mm; mastoid width 11.4-138.0 mm (х = 119.5); the ros- tral width at the canines 28.6-40.0 mm (х = 32.8); and the smallest interorbital width 9.1- 16.0 mm ( = 12.7) (Khuzin, 1963, 1967). The reliability of the craniological differences between males and females is also confirmed by statistical variance analysis. (¢ more than 3 or 2). The differences are reliable in all the populations: condylobasal length, width at level of canines, and smallest interorbital width (Khuzin, 1963; Yablokov and Sergeant, 1963; Khuzin, 1967). The difference in the index of rostral width (as percentage of the condylobasal length) is reliably or significantly higher in males than in females (Khuzin, 1967). Within the territorial waters of the USSR, the overall dimensions and craniometric values of the harp seal reveal no geographic variation since Our waters are host to animals of one single population that repro- duces in the White Sea. The breeding center of the other, i.e., the Jan Mayen population, nearest to the White Sea population, is separated by over 2,000 km from the breeding site of the latter and the transgressions of Jan Mayen animals into our waters are very rare (see р. 407). The differences between the Jan Mayen population, which is isolated from the White Sea population spatially and in breeding sites, are neverthe- less very minor (see pp. 387-390) although the Jan Mayen population is Classified as a special group. (K.Ch.) Taxonomy Pagophilus Gray may be regarded as a special subgenus within the genus Phoca $5. 1., Ph. groenlandica. The seals of Phoca groenlandica differ more from those of the subgenera Pusa and Phoca s. str. than from species of the subgenus Histriophoca. The harp seal has several characteristics proximating it with Histriophoca, the most important being the common final coloration of the hair coat, i.e., alternation of large sections of very dark and bright coloration. Some craniological features too are similar: (1) relatively poorly developed and shortened lower posterior process of the zygomatic bones, the length of which often does not exceed or only slightly exceeds the length of the upper process; (2) forward projec- tion of the temporal bone without enlargement on the anterior margin of the terminal section of the zygomatic process; and (3) some simi- larity in the shape of the posterior margin of the bony palate and in the development of the compact longitudinal septum in the choanae; etc. There are elements of similarity in ecology also: the seals of both 287 382 these species are confined to drifting ice floes without emerging, as a rule, onto the coasts, lead a pelagic mode of life, and perform fairly significant migrations. Based on these elements of similarity, an attempt was made (how- ever, not supported by later authors) to combine the harp seal with the ribbon seal into one genus, Histriophoca (N. Smirnov, 1929, 1935). Much later, it was proposed that these species be combined (at the level of monotypical genera) into a subtribe, Histriophocina (Chapskii, 1948, 1955). These species figure in different genera in several contemporary. works (Scheffer, 1958; king, 1963*; Chapskii, 1963). The similarity between these genera is also seen in the number of cartilaginous rings of the trachea (average 43), relative length of the intestine (13.6 to 14.4 times the body length), variation coefficient of the weight of some internal organs, and relative similarity of the number of labial whiskers (Shustov and Yablokov, 1967). A greater generic proximity between the species compared, than with any others, is also detected in the response to precipitation (V.I. Borisov). The evolution of the harp seal from a common ancestor with the ribbon seal is hardly debatable, but the divergence of these species should be placed not in the Quaternary period (Davies, 1958), but in a much earlier period, probably the Pliocene. There is a view (Winge, 1924, 1941) that the harp seal is more advanced in some respects than the other species, especially subgenera Phoca s. str. and Pusa. The more advanced evolutionary features of this branch are seen in the elongation of the bony palate (also due to the equalizing of its posterior margin) and in the growth of the longitudinal bony septum in the same direction (up to the posterior margin of the bony palate).” Geographic Distribution The subarctic and arctic expanses of the Atlantic Ocean and parts of the northern Arctic Ocean adjoining the eastern fringes of the Atlantic Ocean. Geographic Range in the USSR Constitutes the easternmost part of the general range (Fig. 163). The range of this species within our territorial regions and the adjoining international waters covers the entire coastal belt of the Barents Sea % Such an interpretation of the structural features the skull is somewhat debatable in the light of some new facts (Chapskii, in litt.). 383 along Murman from the boundary with Norway to the White Sea inlet, including all the bays and straits, even those penetrating deeply inland, such as the Kola and Motovsk. Farther east, the range encompasses the entire Kanin-Kolguev shallow-water zone, Cheshsk and Indigsk bays, the mainland portion of the sea to the north and northeast of Cape Timansk of St. Nos, extending along the Timansk coast toward the Pechora Sea from its northwestern, northern, and northeastern regions right up to Vaigach itself. However, the southern and southeastern continental sec- tions of the mainland sections of the Pechora Sea bound roughly by a line traversing from the Russkii Zavorot to Yugorsk Shar, including the latter, and also the coastal waters of Dolgii Island, and all other expanses south of the above line fall outside the limits of the range. Farther north, the region of the regular habitation of the White Sea harp seal covers the Kara Strait zone, coastal waters of the northern one- third and probably also one-half of the western coast of Vaigach, south- ern extremity and entire western coast of Novaya Zemlya, and almost all the rest of the wide expanses of the Barents Sea except evidently its extreme southwestern pelagic portion falling under the warming influ- ence of the Nordkapp branch of the Gulf Stream and bound by the actual position of the edges of drifting ice floes in the period of their maximum distribution. It is difficult to draw any precise boundary here. 287 Fig. 163. Range of the harp seal, Phoca (Pagophilus) groenlandica in the USSR (K.K. Chapskii). 384 The northern limits of the range in the Barents Sea cover the Franz Josef Land archipelago and roughly the same latitudes between it and ~ Spitsbergen. The Kara Sea does not wholly fall within the range: the extensive mainland expanses which, with some approximation, can be described by a wide arc from Vaigach roughly to Minin skerry fall outside the range. The harp seal is evidently not found southeast of this line. Thus it is wholly absent in Baidaratsk Bay, on the western coast of Yamal, in all the bays and straits fed by the waters of the Ob’, Taz, Yenisey, Pyasina, and other rivers (Chapskii, 1938). The harp seal is extremely rare in the eastern mainland regions of this sea but is more common in the Novaya Zemlya strip from the Kara inlet right up to Cape Zhelaniya. In the northeastern regions of the Kara Sea at 77 to 80° lat., the range extends in the form of a fairly long tongue from the line separat- ing this sea from the Barents Sea, i.e., through a corridor between the northern extremity of Novaya Zemlya and Franz Josef Land. Much of the population entering here is evidently scattered in the western sec- tions while a small proportion reaches Severnaya Zemlya. In very rare cases, under a favorable icy environment, extremely small stray groups are evidently capable of entering even into Vil’kitsk Strait. The range of the harp seal in the White Sea is nowhere restricted and covers its entire area, including the bays. In the last century, seals often penetrated deep into Dvina Bay and were encountered even on Mud’yug Island (Danilevskii, 1862), 1.е., almost up to the estuary of the Northern Dvina. They have been seen around those places and also on the Letnii coast, e.g., in Unsk Bay, even in this century. They transgress into Kandalakshsk Bay although they usually do not traverse far into its deep northern portion; however, under unfavorable conditions of the ice drifts stray young animals may be seen even in its extreme cul-de-sac sections. In Onezhsk Bay adults are not usually seen except for very rare transgressions into its northernmost section not far from the Solovetsk Islands; but young ones are seen there at times (rather rarely) (in the years of broken drifting “young” ice).”” In Mezensk Bay this seal is quite common except perhaps for the extreme coastal waters along the south- eastern fringe. A feature of the range of the harp seal is its sharply manifest dynamic character caused by the migratory nature of the animal. These seals are not encountered simultaneously throughout the entire expanse of the range. They cover parts of the range in a certain sequence of regular 97 The “invasion” of molted juveniles in 1966 following drifting “young” ice pools in a reverse direction against the general flow, was an unusual event (see page 425). 289 385 seasonal migration of the entire population or its individual age and sex groups. From late autumn throughout winter and in early spring all the harp seals are concentrated at the inlet to the White Sea and in the adjoining sections of the Barents Sea, from western Murman to the Pechora Sea. The range undergoes maximum constriction in the winter- spring period when almost all the White Sea population is concentrated in the White Sea and in the very near border sections of the Barents Sea. In this period the range evidently forms a ring of broad ice floe fringes whose rough position can be schematically depicted in the form of an uneven arc, with one end resting on the eastern Murman coast somewhere in the region of Cape St. Nos (and from there extends in a narrow fringe along the Murman coast even farther west and, in some years, up to the Norwegian coastal waters). The other end of the arc runs northeast beyond Kanin Nos toward the northern coasts of Kolguev Island and farther in the direction of Novaya Zemlya somewhat taking advantage of the edges of the ice floe belt. In a prolonged autumn and late winter, this expanse is more enlarged and extends farther northeast toward the Kara inlet. Such was the situation particularly in the first half of the winter of 1966 -67, when the seals were confined in isolated herds throughout such a long broken edge (Beloborodov, 1969). From autumn and very early in winter, the seals are often seen close to the coasts but there is not much information about their encounters due to the darkness and cessation of navigation. The animals remain exclusively in the water and tracing out their distribution even in the White Sea is quite difficult. Only from the beginning of February, when the period of reproduction is quite close and ice rookeries begin to form, is it possible to establish the location of the breeding section of the population. However, even right at the peak of whelping and lactation, when the female population with offspring is localized in certain parts of the White Sea, the location is not clearly known of the immature portion of the population and of those adults which, for some reason, have not participated in reproduction in a given season. In April and up to early May, almost all the White Sea population is even more localized, mainly at the inlet and in the adjoining regions of Mezensk Bay, but quite often in the neck and even in the central basin of the White Sea, and partly right in the nearest regions of the Barents Sea, in the zones of the ice fringe. From mid-May the seals usually desert the White Sea, move rightward on the fringes in the most proximate region of the southern part of the Barents Sea, into the region of Kanin and Kolguev, and farther toward Gusin Land (Novaya Zemlya); later they even scatter on the new ice fringes of the Barents Sea right up to the Medvezhii and Spitsbergen islands, moving from the fringe increasingly 290 386 north and northeastward. At the end of July and in August, they reach the northern limits of the Barents Sea, transgress into the Franz Josef Land archipelago, in the northwestern regions of the Kara Sea, totally disappearing at this time not only from the White Sea, but also from the immense expanse of the Barents Sea. In the high latitudes, along the edges of the arctic packs to the east of Spitsbergen and among the sparse drifting ice north and east of the northern extremity of Novaya Zemlya, the White Sea harp seal is quite scattered in small groups (sometimes adult males even singly”*) right up to early September, and later extends southward into the wintering ground. It is again concentrated at the time of reproduction in the White Sea region and the southeastern parts of the Barents Sea and small numbers are even seen in Cheshsk Bay. This, then, is a general outline of the seasonal dynamics of the range of the easternmost White Sea populations of the harp seal. Geographic Range outside the USSR (Fig. 164) Extends from the central and peripheral regions of the Barents Sea in a broad strip through the entire northern half of the Norwegian Sea and Spitsbergen Strait, northeastern and southwestern regions of the Green- land Sea (to the west and northwest of the Martovsk ice fringe) into Denmark Strait. The range encompasses the coastal waters of northern Iceland and encircles the eastern and western coast of southern Green- land in a relatively narrow strip. To the west of the meridian passing through Cape Farewell, the range includes Davis Strait and Baffin Bay (including the whole coastal periphery on the Greenland side as also on the western Canadian fringes). The seals are encountered in small numbers in Kane Basin and are more abundant in Jones Sound and traverse along Lancaster Sound up to the Wellington, Barrow, and the northern part of Peele straits inclusive. Depending on ice conditions, advance populations enter the Gulf of Boothia through Prince Regent Strait. The seals are common along the entire eastern coast of Baffin Island, the Labrador coast, in Hudson Strait, and in the northern part of Hudson Bay, descending along it almost up to the Belcher Islands. The southernmost part of the range encompasses all the sides of Newfound- land including the Gulf of St. Lawrence, the shelf along Nova Scotia, and probably the region of the Great Newfoundland coast. The branch of the range diverging from the White Sea basin into the northwest along Murman extends rather irregularly around the Varanger 98 Observations in the Kara Sea, southeast of the Blagopoluchiya Strait (К.К. Chapskii). 290 291 и FOR ока Fig. 164. Species range of the harp seal, Phoca (Pagophilus) groenlandica, and the site of its concentration during whelping and molt (dotted spots) (K.K. Chapskii). Peninsula and the more western highly rugged Finmark coasts, descend- ing to the Vesteralen Islands and even to the southern Lofoten Islands sporadically and only in cold winters and in spring. In extremely severe winters, аз in the very early years of this century (especially the win- ter of 1902-1903), the herd of “invading” seals spread out even farther. Somewhat similar but much smaller “invasions” were observed on the Norwegian coasts even later. The presence of stray specimens far beyond the limits of the normal range in the seas of Western Europe is extremely rare, episodic, at times even totally improbable. Such was the case of a gestating animal found floating in the Elba River 500 km upward from the estuary in 1896. Stray animals were sighted time and again on the coasts of Great Britain, from Scotland to the Thames. Stray animals were found on the northern coast of France. There is no reliable information about the sighting of these seals on the coasts of Sweden. In Norway the transgression of a stray juvenile into the extreme south was reported at Oslo Fjord in 1936.” (K.Ch.) Geographic Variation The intraspecific structure of the harp seal was discussed even forty years ago. It has long been known that the entire population of this species is divided into at least three geographically isolated populations (or herds), % Data on the episodic transgressions of seals were taken from Collett (1911-1912) and Moore (1952*). 388 each of which has its own features, extremely restricted areas of repro- ` duction, and different regions of wintering and molt. Based on this, it was quite natural to decipher some morphological features characteristic of each of these populations. Even the earliest attempts to identify specific features of craniological indices (N. Smirnov, 1924, 1927, 1929) recorded some differences in average values between the different populations. These differences were subsequently refined. Only one subspecies is known within the USSR. White Sea harp seal, or “Lysun,” Phoca (P.) groenlandica oceanica Lepechin (1778). This is the largest form of the species. The body length measured along the dorsal surface (Lc), according to the accurate data of Khuzin (1963), for males (100) is 169-204 cm (x = 185), for females (300) 163-229 cm (х = 183). The skull dimen- sions (according to the same source) are: condylobasal length in males (37) 200-234 mm (х = 217), in females (300) 200.5-223 mm ( = 209.5); mastoid width in males (37) 113-127 mm ( = 121), in females (27) 111-124.5 mm ( = 117); rostral width (at level of canines) ш. males (38) 32-46 mm (x = 36.2), in females (27) 27.5-37.0 mm ( = 31.5). White Sea, waters of the USSR in the Barents Sea, and Kara Sea. The White Sea represents the zone of breeding. Outside the USSR, this species is found in the waters of northern Norway and Spitsbergen and western and northern parts of the Barents Sea. * OK * Outside the USSR, usually only one subspecies is recognized, 1.е., the Newfoundland harp seal, Phoca (P.) g. groenlandica Erxl., 1777; dur- ing reproduction and molt, these are concentrated in two sections of the range: slightly northeast of Newfoundland Island and in the Gulf of St. Lawrence. According to the accurate data of Khuzin (1963), the body length of males (83) is 152-195 cm (5 = 176.5), of females (127) 156-201 cm (х = 175.5); condylobasal length in males (39) 200.0-219.5 mm ( = 208.5), in females (41) 191.0-219.0 mm (х = 204.5); mastoid width in males (40) 109.0- 124.0 mm (х = 117.5), in females (41) 109.0- 123.5 mm (< = 116.0 mm); rostral width in males (40) 30.0- 40.5 mm (х = 33.5), in females (41) 27.0-36.0 mm (x = 31.5). Statistically, the most reliable differences between the White Sea and Newfoundland populations are seen in the overall body length (¢ = 8.52) and in the length of the palate (¢ = 4.61) as well as in the condylobasal length and mastoid width of the skull, etc. 292 389 The range includes the northwestern, predominantly subarctic and arctic Atlantic including Davis Strait, Baffin and Hudson bays, and regions of the southwestern and perhaps southeastern coastal waters of Greenland. The real subspecific differences between the Newfoundland and east- ern Atlantic populations are also confirmed by the presence of a char- acteristic genotype in each of these groups (Naevdal, 1965). Insofar as the Jan Mayen (Greenland) population, which is spatially well isolated, is concerned, it reproduces in the region of Jan Mayen Island and is generally distributed in Denmark Strait and the Greenland Sea; it reveals almost no statistically reliable craniological differences from the White Sea subspecies (Khuzin, 1963; Yablokov and Sergeant, 1963) and does not differ from the latter biochemically in the protein polymorphism (Meller, Naevdal and Valen, 1966*).100 Thus this population evidently cannot yet be regarded as an inde- pendent subspecies; it represents a variety, tribe, or herd closely related to the White Sea harp seal; in fact, the two were regarded in the past as essentially similar (N. Smirnov, 1929). (K.Ch.) Biology Population. Although the total world population of the harp seal has decreased very significantly over the long period of its hunting, it was still regarded as considerable in the mid-1950s. It was approximately, but evidently overestimated, put at 5.5 million (Dorofeev, 1956) or 4.5 -7 mil- lion (Scheffer, 1958). Out of the former assessment, 3 to 4.5 million were regarded as the Canadian-Newfoundland herd. This was followed by the White Sea population at 1-1.5 million. The Jan Mayen population at 0.5-1 million came third. A total figure of 3-3.5 million is evidently closer to the real position (Chapskii, 1966). The harp seal is one of the few species for which the population has been determined by different methods, including aerial photographic survey. The latter procedure was used for the first time in the world in 1927 to count our White Sea herd. Part of the concentrated molting rookery was photographed, its area determined, and by extrapolation and assumption the population put at one million (Dorofeev and Freiman, 1928). This operation was repeated in 1928 and the total population of the White Sea herd was placed at 3-3.5 million (Dorofeev, 1939). 100 At the same time, some other morphological features of the rank of the population were also identified; differences in body length, in distance between teats, and in number of asternal ribs and whiskers (Yablokov, 1963). 390 In the second half of the 1930s, new population counts were made based on different methodological procedures, i.e., by counting the mother seals (or the pups). The main stock (М. Smirnov, 1928)101 was initially put at 600,000-700,000 (Р.А. Rudakov and N.V. Provorov). Somewhat later, double this figure or 1,300,000 was cited (N.V. Provorov). The first postwar experiment in studying the White Sea population enabled a rough evaluation of this herd in 1947 at 1,200,000 (K.K. Chap- skii).!°? A quarter of a century after the above surveys and calculations, the same aerial photographic survey was again employed, which gave a figure of 1.2-1.5 million (Surkov, 1957). These figures incontrovertibly confirmed the sharp (by more than one-half) reduction in the popu- lation compared with the situation prevailing some 30 years ago. On the basis of this figure, however, no conclusion was drawn “about the - adverse effect on the status of the reserves” by excessive hunting (Surkov 293 and Khuzin, 1959)103 and the herd continued to degrade for some time thereafter. In the 1960s, extremely disturbing warnings about its distinctly unfavorable position appeared in the press (Nazarenko and Yablokov, 1962; Yablokov, 1962; Yakovenko and Nazarenko, 1962; Yakovenko, Nazarenko, and Timoshenko, 1963; Yakovenko, 1963, 1967, etc.). The total population of the herd in the early mid-1960s was put at differ- ent levels: 400,000 -700,000 (Nazarenko and Yablokov, 1962), 750,000 (Yablokov, 1962), a minimum of 400,000 (Chapskii, 1966), and a min- imum of 225,000 (Yakovenko, 1963). The last figure is regarded as an underestimation (Khuzin, 1972*) although the mother population based on the data of a 1963 aerial photographic survey of rookeries was put at only 65,000. The total population in all the photographed molting rookeries in 1962 and 1963 did not amount even to 200,000. The total of the White Sea herd calculated on this basis did not exceed 300,000 by 1963 (Yakovenko, 1967). The reduction in total population of the White Sea harp seal in the 1950s is also strikingly demonstrated by the dynamics of the shrinking areas of its nurseries and molting rookeries (Table 15). The age-related composition of the productive population (Doro- feev, 1939) also served as an index of the adverse status of the pro- ductive proportion of the herd in which animals older than 10 years became very few in the early 1960s (Khuzin and Potelov, 1963; Khuzin, 101 That is, without pups. 102 Data for the 1930s-1940s have been taken from Khuzin (1972*). 103 Assuming that all the 100,000 animals killed were pups. 293 293 294 Sil Table 15. Variation in the area of concentration (rookeries) of harp seals in the White Sea, km? (Yakovenko, 1967) Year of observation Type of rookery А ео ава oe crt 28 беби в 1953 1954 1955 1956 1957 1958 1959 1960 Rookeries in the lactation period (nurseries) 294 290 200 167 143 126 130 120 Rookeries in the molting period (molting) 851 27564 58) 58 48) 46 48 Table 16. Total average annual kill of the harp seal in the White Sea and in adjoining areas of the Barents Sea at five-year intervals from 1947 - 1964 (in thousands) 1947-1951 1952-1956 1957-1961 1962-1964 190.0 125.2 117.6 89.7 1964). Hunting statistics too provided an equally striking account of the population reduction (Table 16). AS a result, an agreement was reaced with Norway in 1965 to sharply reduce and modify the trend of marine hunting in this region. The mea- sures adopted have already begun to show positive results. A similar process of population reduction of the harp seal was also reported in the western fringe of its range where the Canadian population concentrates in the period of reproduction and molt. According to the calculations based on aerial photographic surveys in 1950 and 1951 of the nurseries in the Gulf of St. Lawrence and in the open Atlantic northwest of Newfoundland Island, the total reserves of these herds were put at roughly 3.3 million at the beginning of the second half of this century; of this, some 650,000 were regarded as pups (Fisher, 1952, 1955; Sergeant, 1959). Survey-based calculations gave somewhat lower figures for 1959 and 1960: the total population of pups in both the Newfoundland regions (in the gulf and in the “front” region) was put at 365,000. The basic stock (1.е., without pups) of both these herds was, however, put at only 1.2 million (Sergeant, 1963, 1965). The 1964 census,confirmed these figures (350,000 pups) (Sergeant, 1965). No census has been done of the Jan Mayen population but it has been roughly put at a maximum of a million (Fisher, 1954; Scheffer, 1958; Dorofeev, 1965; and others). With such a dynamic nature of the range of the harp seal, its local population year round is also highly variable. By changing location from season to season, the population gathers sometimes almost wholly in relatively small sections of the range and at other times is scattered over a wide expanse (see above). 392 Habitat. The harp seal is a distinct inhabitant of cold waters but avoids the arctic pack ice, preferring the marginal zones and regions of stable drifting ice floes prevailing year round (as near Jan Mayen Island) or during low and high tides (as in the neck or inlet zones of the White Sea). It is this type of biotopic conditions that is selected by this seal in the periods of breeding and molt. In the course of these biological cycles, its herds appear deep in the ice massifs quite far from the marginal zones. In the breeding period it selects large stable ice floes and ice fields even with hummocks as a solid substratum. It is less choosy in the molting period and is seen on the fringes of large ice fields as also on groups of small ice floes if they provide a good link with the water body. This seal does not use the stationary shore ice (fast ice), as far as is known, under any circumstances. Like the other pagophilic seals, the harp seal also crawls onto the ice floe but resorts to this mainly when the open water pools between the drifting ice floes on which it whelps are frozen. At the end of the winter-spring period of reproduction and molt, the seals move to the fringes of drifting ice floes (at the beginning of summer) or are confined generally among the fairly sparse ice floes (in summer). At the same time, it is not quite correct to stress that the “harp seal spends its life among the ice floes round the year” (Freiman, 1939). At the end of summer, in autumn, and even in early winter, when these seals begin and continue to migrate southward and even appear in the region of breeding, the environment is no longer icebound (Chapskii, 1961). Pelagic life is highly characteristic of the White Sea harp seal but nevertheless during the autumn migrations and on Murman even in spring, it is quite often seen around the coasts, transgresses into the bays, even those penetrating deep inland, such as the Kola, and may traverse through narrow Straits, at times extremely narrow, such as the Zheleznye Inlet (on Loginov Island south of Novaya Zemlya). In the ice-free period, however, the seals distinctly exhibit a preference for the coastal strip and do not stray far from the coasts. Food. The food of the harp seal, including our herd, in spite of the earlier optimistic assessment (Ognev, 1935) has not been adequately studied to date; the list of food objects is far from complete for much of the annual cycle, especially from June through January and February, when its feeding is very intense. Among the invertebrates found in its food are the crustaceans, mainly euphausids (genus Thysanoessa) and amphipods (genera Anonyx, Gammarus, Themisto, Gammarocanthys, and Others), and also shrimps (Crangon sp., Sclerocrangon boreas, Pandalus sp., etc.); among the. 295 393 plankton, mollusks of the genera Clio and Limacina. There is no accurate list of the species of caphalopods consumed, but these are mainly squids as also cuttlefish. Among the fishes consumed are capelin, polar cod, navaga, cod, coalfish [pollack], herring, plaice (Hippoglossus), sea bass, and even goby. At the end of the last and the beginning of the present century, harp seals fed along the Murman coasts quite regularly in spring as well as in autumn. This was the period of considerable increase in ice coverage of the Barents Sea and hence of a sharply reduced ice-free expanse, which constituted a seasonal contraction of effective area for the White Sea population of this seal. Fish was evidently the mainstay here, as confirmed by actual observations. Earlier investigators may have erred in assuming that the harp seal consumed a sizable quantity of cod and even chased this fish away from the Murman coast (Knipovich, 1895; Knipovich, Yagodovskii, and Zhikharev, 1902; partly N. Smirnov, 1903; Breitfus, 1903) and also evidently in other regions (Allen, 1880; Wolleback, 1907; and others). Direct observations and dissection of the animals contradict these views. In the spring of 1900, from early March through the 20s of April, along the Murman coast, especially in the region of Kil’din Island, seals fed intensively on capelin (Mallotus villosus) and the stomach of 34 of 58 dissected animals contained fairly significant quantities of this fish (Breitfus, 1903). From the end of November, throughout December, 1902, and also in January and February, 1903, herds of seals wandered in the coastal waters of the western Murman and “their stomachs were stuffed with herring” (Breitfus, 1906). From the last 10 days of February through April end, 1905, the arrival of groups of harp seals on the Murman coasts was associated with the abundance of small coalfish (Prigorovskii and Breitfus, 1912). Commencing from the end of lactation and molt, the juvaniles seek food independently, feeding initially on large components of zooplank- ton present right among the ice floes. At the end of March the molted pups almost cease to look for small crustaceans and prefer to rest on the ice floes, resuming an active search for food in April. In this month their stomach or intestines mainly contained remnants of euphausids (Thysa- noessa inermis and Th. газспи) and amphipods (Anonyx nugax, Parath- emisto sp., etc.) (N. Smirnov, 1903, 1927; Dorofeev, 1936; Sivertsen, 1941; Chapskii, 1961, 1964; and others). It is possible that the list includes the pteropod mollusk (sea butterfly) in the more northern regions. Data are inadequate on the subsequent diet changes of pups on emerging (or drifting) into the fringes of the Barents Sea and also on the food of the adults there. At the very beginning of the 1920s, when the ice conditions along the Murman and northern Norwegian coasts 296 394 were unusual, the pattern of gradual intensification of feeding of the underyearlings and the enlarged species range of food items used by them were as follows. Initially feeding on minute plankton, the pups “later take to fish, at first polar cod ... available abundantly among the ice floes ... and later take to the food generally characteristic of this species, leaving no fish that is readily available, be it capelin or cod, surfacing from the bottom after it, and herring, and others” (N.A. Smirnov, 1903). There is some exaggeration here too about the cod. Smirnov also stated that “if herring were plentiful in the Murman, the bulk of ‘skin’ (1.е., harp seal, K.Ch.), mainly the young ones, would remain there” (N. Smirnov, 1903). Later, however, the views on this subject were more reserved and mainly “small pelagic fish or those confined close to the ice fringes” were added to the list of fishes consumed by the White Sea harp seal (N. Smirnov, 1924, 1927, 1935).104 According to some investigators (М. Smirnov, 1903; Ognev, 1935), there are no differences in the nature of feeding between underyearlings and yearlings; according to others, however (Sivertsen, 1941), some dif- ferences do exist. Fish and deep-water and benthic invertebrates occupy a prominent position in the food of yearlings in addition to the pelagic crustaceans On which 1.5-month-old pups feed. Thus, the stomach and intestines of some yearlings caught in the White Sea in the last 10 days of April, 1934, sometimes contained in addition to the euphausid, much larger crustaceans (Crangon crangon and Spirontocaris turgida) and amphipods, capelin being found more often (Sivertsen, 1941). Regard- less of the foregoing situation, there is evidently no significant difference in the food of yearlings and adults. The seals do not feed regularly during reproduction and molt, at least not daily; many generally remain hungry. Their stomach is often empty and highly shrunk. Of the hundreds of adult animals dissected at hunting sites in the 1920s and 1930s (Sivertsen, 1961), food was found in the stomach of two: March 15, 1929—male (the stomach was filled with euphausids) and April 3, 1931—female (the stomach contained shrimps). A similar picture was observed in special investigations conducted at the beginning of the 1960s in the White, Barents and also Greenland seas. Not even one of the hundreds of animals examined contained food remains in the stomach. In most cases the stomach was markedly shrunk but its wall was very thick. In this period of such physical starvation the gastric juices were neutral (Shepeleva, 1963). According to other reports, the lactating females satiate themselves from time to time “if some food 104 The “invasion” in Murman waters of this seal and the disappearance of cod there is a consequence of sharp atmospheric cooling (Linko, 1912; and others). 395 is available in the nursing region”. The food comprises mainly planktonic or nectonic crustaceans (Surkov, 1960). The seals not engaged in lactation or reproduction and living in this season in the more northern regions of the White Sea, along the fringes in the adjoining sections of the Barents Sea, and on the open Murman coasts, evidently feed regularly. Feeding is interrupted only in the period of molt when the animals mainly rest on the ice floes. In the years of deep penetration of these seals westward, into the Norwegian Sea, some instances are known of their being trapped in fishing nets set at a depth of about 200 m. Two animals were trapped even in the Arctic Circle, one of them in the region of Vardo even at a depth of 280 m (Collett, 1911-1912). With inadequate information on the behavior of such deep submergence, Nansen (1924, 1939) himself pointed out that, apart from the common fish food, the White Sea harp seal was capable of surviving on sea bass and flounder as well as on cuttlefish. Thus the “food spectrum” of the White Sea harp seal is evidently quite broad and has not been adequately studied. In the 1920s, when the nurseries were located more northward, on the threshold of the Barents Sea, it was affirmed that the females at the end of the breeding season migrated for sometime, especially for feeding, to the Murman coast (Skvortsov, 1927; Dorofeev and Freiman, 1928). It is possible that such local migrations did in fact occur since the “young” ice floes at the time of the cessation of lactation often drifted away toward the Barents Sea. At the same time, such an interpretation of the move- ments of mother animals can be the result of inadequate reliable data. Regardless, such massive migrations have not been recorded in the past 40 years. In a much earlier period, as already pointed out, not only the adult females, but also other groups of the White Sea population were generally encountered feeding in considerable numbers on the Murman coast commencing from late autumn through the end of spring. Harp seals in particular were noticed here right from early March through the last 10 days of April, i.e., in the period covered by the formation of nurs- eries and molting rookeries. Evidently, the severe weather conditions in winter, as for example in 1901-1902 and some other years, exerted an unusual influence; in such years the edges of ice floes extended almost up to Kil’din Island. The period of intense feeding now commences evidently from May, immediately after the cycle of reproduction, after which the seals desert the White Sea to molt. Breaking up into small groups along the entire edge and among thin ice floes, they feed mainly on macroplankton (the more massive species of crustaceans) and the polar cod scattered over immense expanses of the arctic seas. The summer-autumn range of the 297 396 harp seal is mainly dependent оп the regions of distribution of this fish, which has great food value for the seal. Feeding in the high latitudes continues all through summer. With the onset of autumn, when the polar cod begins to form large schools and moves toward the coastal regions for spawning, harp seals pursue it. This fish is now almost the mainstay of these seals on the coasts of Novaya Zemlya, Kolguev, and Kanin, and especially in the northern regions of the White Sea. In the latter region, apart from polar cod, herring and partly perhaps navaga, represent important food items. Feeding in the autumn-winter period is no less intensive than in summer: everywhere in the coastal waters of the Soviet arctic, concentrations of seals and en masse arrivals of polar cod are noticed simultaneously (Klyuche,* 1936; Manteifel’, 1943; Chapskii, 1938, 1961; and others). Home range. The concept of home range is difficult to apply to the ecological features of this species whose populations not only migrate widely, but vary sharply in concentration in different seasons. The herd instinct in the harp seal is quite sharply manifest; it is evident even in summer when the population is highly dispersed, in small groups of usu- ally not less than some 10 animals. One such “congregation” in which the animals are confined sometimes more closely and at other times less so (although usually not closer than 5-10 m) is separated from another by varying distances that are not conducive to numerical averaging. Evi- dently in regions of highly dispersed food objects (for example, the polar cod), the seals can be found in some cases even singly and far removed from each other. During the autumn migrations, the animals gather into much closer and larger “congregations” but usually do not remain for long at one place. The age-sex structure of the feeding and migrating groups is not yet clearly understood; however, there is considerable justification for assuming that the migrants segregate into young and adults; the latter, even in high latitudes, then separate into sex groups, which is particularly noticeable among animals on the threshold of the winter-spring grounds. Here the concentration coefficient is also highly increased. The maximum density of animals in the White Sea occurs at the time of formation of the rookeries (Fig. 165). In 1963, mother seals were concentrated with their offspring at three sites over a total area of 108.3 km? where 60,000 whelped females were photographed (Yakovenko, 1967). In fact, the females there must have been somewhat more since not all of them were basking on the ice floes at the time of the aerial photographic survey. Assuming that some 10% of them had remained in water and making a correction of at least 5,000 accordingly, the average density of disposition of lactating females in the nurseries works out to roughly 600 298 397 Fig. 165. Part of the harp seal rookery on ice floes in the White Sea. Pho- tographed from an airplane (material of PINRO) [Polar Scientific Research Institute of Sea Fisheries and Oceanography, named after N.M. Knipovich]. animals per km? or one female in an area of roughly 1,660 m2, 1.е., ава distance of roughly 40 m from each other. Actually, however, the mother seals are disposed far more densely and are not evenly dispersed, remaining in groups concentrated mainly along the fringes of large or finely broken ice floes that refreeze due to the formation of young ice floes at the site of open water pools or crevices. On these new ice floes extending in weird, twisted, and inter- laced strips, tongues and much broader sections, the females and their pups gather in far denser groups than in the central parts of the ice 298 299 398 floes. Moreover, drifting ice floes with animals on them generally alter- nate with “vacant” ice floes (see p. 397). Thus the lactating females in fact live within some 5-10 m of, or even closer to each other. In the molting rookeries the density is much higher: 165,000 animals of mixed composition (adult males, immature animals of either sex in the age of one year to five or six years, and adult females) covered a total area of 76 km? in 1963 (Yakovenko, 1967), 1.е., average of 2,200 animals рег km?. In other words, there was one animal in an area of 455 п? (average distance of 20 m from each other). In fact, however, in this case too the molting animals are usually much closer, especially in the initial, exclusive male preserves where they literally rub shoulders, quite often in very dense rows almost fringing the edge of the ice floes (see Fig. 165). The sequence in the selection of biotopes is determined not only by the requirements under the definitive conditions of reproduction and migration, but also by the food factor, in particular the nature of the seasonal distribution of the polar cod, which is almost the mainstay of the White Sea harp seal and hence a factor responsible for the disposition of these seals. Hideouts and shelters. A hard substratum (ice) is used by harp seals almost exclusively in the period of reproduction (for whelping, lacta- tion, and partly mating) and molt, i.e., in the early spring (winter-spring) period. Very rarely, animals are encountered on ice floes even in the midsummer season in high latitudes (evidently the sick or well-satisfied animals induced by the excellent weather). Thus these seals reside in water for an incomparably long part of the year. This seal never attempts to build hideouts in the snow on top of an ice floe, as done by the ringed or the Baikal seal. The only thing the White Sea harp seal is capable of doing is making air holes in the ice floe, a characteristic of most other species of pagophilic seals. The mechanism of formation of the air holes evidently has no dis- tinctive features. The seals resort to making them only in those cases when the open water pools between ice floes with pups on them begin to be covered by thick ice. The animals surfacing for respiration pierce the ice with their head. As long as they do not move to another site, they use the openings already formed, which are kept unfrozen by the animals through frequent surfacing and crawling out onto the ice. In the course of time these air holes assume the form of a low crater initially fringed by broken bits of ice and later by the water splashed during surfacing. Many animals can use the same air hole and hence there are usually fewer air holes in a nursery section compared to the number of lactating females. Since the ice floes in the breeding region are on the move constantly with open water pools usually occurring between them, the air holes do 399 not represent the lone means of contact for the animals between water and air. The seals hardly use air holes in the much later spring molting rook- eries, being accommodated by the natural openings in the ice floes in the form of open water pools, crevices, “gapes,”!° and “partitions,” which never close for long because of high and low tides, drifting ice floes, and floe movements caused by winds. In fact, there have been instances when the open water pools closed in high tide and froze when filled with ice bits. The seals surprised by hunters found themselves separated from the water. If, however, the “gape” is not densely or firmly frozen, the animals can put to advantage their weight and pierce the ice cover to hide below it. Similarly, the heavy animals can break up even thin young ice extending in quiet frosty weather over an open water pool or in cracks and crevices in the ice floes, especially by their combined might. These seals resort to no other shelters. Only the newborn in the very first days of life on the ice floes seek some shelter from foul weather, creeping time and again into niches or crevices formed before they were born during the hummocking of the ice floes, or hiding under an ice floe overhang. Sometimes the pups hide so deeply under the ledges in piled up ice floes that they readily escape detection. Daily activity and behaviour. The activity of the harp seal is not stereo- typical from day to day; it differs in different seasons of the year depend- ing on the characteristics of the successively changing biological cycles to which not all the age and sex groups of the population respond to the same extent. Thus there are periods of high and low activity. Animals participating in reproduction exhibit hyperactivity, which is somewhat more prolonged among females than males. For much of this period the males are even passive (up to the moment when their sexuality peaks, which is of relatively short duration). Females with pups, however, are active for at least 2-2.5 weeks of very intense lactation, though with intervals for whole days, since the pups are suckled not only during the day but at night, and go into the water many times between suck- lings. An extremely high motor activity, increasingly manifest in daylight, is noticed in the short mating period, accompanied by chasing competitors and seizures (see p. 400). 105 Discontinuities formed between the frozen masses of ice floes, sometimes compressed (during compression) and sometimes diverging or enlarging (in low tide) are called “рарез” by the coastal hunters. 300 400 The last period, 1.е., molt, is characterized on the contrary by maxi- mum passivity, especially among males who rest in small groups on the ice for days on end without going into the water. The females, however, and also the young (commencing from yearlings) are still in the water at this time and join the males later. At the end of molt, the animals again enter a period of high motor activity and migrate into the zone of summer residence and intense feeding. Their “activity,” directed toward | procuring food in the zone of ice fringes in this period, can be assumed to be manifested predominantly in daylight hours which rapidly increase. Nevertheless, their activity does not cease at night. The summer behavior has not yet been clearly understood. Evidently, having expended considerable effort on migration into higher latitudes and feeding well on the way, the animals are now widely scattered in the northern limits of the range and do not exhibit much activity. The latter increases somewhat later with the commencement of reverse migrations, formation of herds, and transition to intensive feeding on fish in the autumn. Moving from place to place in autumn and early winter, on their way to the White Sea and in it, the seals avidly hunt for polar cod and other fishes during the day and also at night. Further, they are capable of submerging quite deeply, as demonstrated earlier by the example of some stray animals being trapped in fish nets set at considerable depths (see below). The herd instinct is highly typical of the harp seal and is mani- fest not only in the formation of strictly localized rookeries in which almost the entire population of a region gathers. It is also reflected in all the behavioral features of these animals. The seals remain in herds everywhere, on the migratory courses, in the wintering sites, and in the feeding grounds. Only in the summer period is some deviation sometimes seen. No aggressive tendency whatsoever is exhibited toward man under normal conditions; on the contrary, with the approach of man, the herd tends to leave the ice floe. However, the maternal instinct is so intense among the females that it suppresses the instinct for self-preservation and they bravely guard their pups without regard for themselves. There are no reports of any scuffle whatsoever among the animals except during the mating season. Adults and immature animals are not heard but a hungry pup calls its mother with a loud wail, quite similar to the cry of a child. It is difficult to establish with certainty which of the sense organs are better developed. Reports that these seals are “frightened” by ship smoke have no serious basis as the olfactory faculty is the least devel- oped in all the pinnipeds. Their vision is quite good in water as well as 301 401 on land. Near, rather than distant, sight is probably better on land. The auditory faculty is evidently well developed. The ability to echolocate is beyond doubt among harp seals; otherwise, it is difficult to explain how this animal can orientate itself in water and catch quarry at depth even under conditions of the polar night. This faculty has been demonstrated experimentally by recording underwater sounds in a hydrophone (Mohl, 1968; L. Popov and Pleshakov, 1970). Seasonal migrations and transgressions. In the nature of its migra- tion, the harp seal has almost no peer in the family of true seals (Pho- cidae). Only the hooded seal and partly probably the Caspian seal are comparable, but the latter falls behind sharply in the magnitude of this phenomenon. Although this aspect of the ecology of the harp seal was known from the earliest published works, i.e., at least from the last quar- ter of the eighteenth century, it has not been thoroughly investigated so far and much is yet unknown. In particular, all the details of the migrations of a well-fed juvenile which leaves the breeding site for the first time, are not known for cer- tain. In fact, the initial migrations of the under-yearlings bear a passive character depending on the general drift of the ice floes on which the pup continues to rest for sometimes (up to March end to early April) after the final molt (i.e., transformed into a gray pup). Under normal conditions of the White Sea basin with sharp drifts (i.e., when the ice floes drift northward through the neck into the inlet), the gray pups present partly on the ice floes and partly in the open water pools between them, drift in the same direction as the ice floes. Under the influence of this drifting, the pups ultimately find themselves in the northernmost regions of the White Sea or even beyond, on the fringes of ice floes in the southern sections of the Barents Sea. Here the gray pups feed from time to time and remain for sometime, possibly until they are overtaken by the molted adults, 1.е., firstly the adult males and later | the immature animals of both sexes, as well as gestating and wandering females. All this represents only a working model as accurately as the actual picture can be deciphered. One cannot entirely agree that the gray pups abandon “young” ice floes for arctic latitudes solely guided by some subconscious desire to reach the north (Danilevskii, 1862). The situation that prevailed in 1966 demonstrated that such is not so. The exceptional complexity of the icy environment in that year, caused by steady northeastern stormy winds, disturbed the normal pattern of the ice drifts. The ice floes on which whelping occurred were not transported into the White Sea inlet nor into the Barents Sea, but were pressed to the western and southwestern coasts of the White Sea where they thawed with the onset of warmth. The 402 molted juveniles on these ice floes and in the water around them thus found themselves not in the northern sections of the White Sea, but on its western and southern fringes where they strayed in early summer into totally unexpected places, up to Arkhangel’sk and Kandalaksh inclusive. By the end of April to early May, when the White Sea harp seal is quite prepared to emerge from the region of breeding but not yet wholly joined by the migrating stream, almost all the White Sea population is concentrated in the northern parts of the inlet zone, on the threshold of the Barents Sea, and in its southermost sections. Here the seal pop- ulation is partly confined to large rookeries and partly highly scattered in the form of small herds and groups on the ice floes as well as in the water. Their actual disposition, no doubt influenced in the recent past by hunting, depends mainly on the position of the ice fringes, on the characteristics of the ice regime, the nature of distribution of the ice floes, and on other as yet unidentified factors. As soon as the majority of the seals complete molt and the molting rookeries in the White Sea are disbanded, the movement of the animals along the fringes of the ice floes of the Barents Sea becomes evidently somewhat more active. The herds spread increasingly along the fringes and, as they recede, move farther northward and ultimately reach the summer range. The directions of spring migrations (Fig. 166) do not remain strictly constant but depend on the contour of the ice fringes. However, these migrations can be generalized schematically as follows. During May and June the movement of the seals from the southern parts of the Barents Sea proceeds initially along an arc approaching quite closely to Kanin Nos, Kolguev Island, and Gusin Land. Roughly on the latitude of the latter, in spite of the disposition of dense massive ice floes, the animals sometimes move north, sometimes to the northwest along the ice fringe receding gradually northward, and scatter almost up to Medvezhii Island and to the ice floes surrounding Spitsbergen. When, however, a rather large batch of the population thus moves west of the meridian of the Kola Peninsula, the ice floes even to the east of the Barents Sea rise markedly northward; they open up in July and by August provide access for the seals into the northeastern regions of the sea. Following the receding ice floes and holding on mainly to their fringes, large numbers of the White Sea harp seal reach high latitudes in August, presumably spreading increasingly predominantly in the eastern regions of the summer range. However, on their northward journey in the spring-summer period, they often cannot reach the coasts of Novaya Zemlya directly which are still blocked by ice floes, although the seals are 303 evidently numerous along the outer edges of this icy belt. In July-August, 403 "ош pue чоцопролАэл jo $1018э1—ф ‘зиоцел8ци ишизие—$ pue ‘8и19$—2 ‘э8иел ay] JO зэперипоа—1 :(нуз4ецо yy) вофрив]изо48 (snjiydo3vg) vo0yg Чеэз diey еэб эйцм ey) JO зиопел8ни ay) BuIMoYsS эла8 ‘99т ‘ЗЫ ZOE 71 IWN Е-7 ef") 1b] 404 а considerable number of them are seen in the broad corridor between Novaya Zemlya and Franz Josef Land. Some herds penetrate the straits of this archipelago while a much larger number of them transgress into the northwestern part of the Kara Sea. Small groups go east, sometimes very far, reaching the western coasts of Severnaya Zemlya. Migration in the reverse direction commences in September. Now, abandoning the zone of the ice fringes, which in the autumn holds no special interest in the context of food availability, the seals go south and southwest (and in the western boundaries of the range possibly even southeast), and probably through the open sea. Unlike the spring migra- tions, on their reverse course, which has not been thoroughly studied, the seals determinedly adhere to the Novaya Zemlya coasts. One of the most important stimuli for their approach to land is the commencement of arrival of spawning polar cod along the coasts on which all the arctic seals feed in this period. In the region of Cape Zhelaniya, harp séals are seen even in Septem- ber and later at almost all the other points on the west coast right up to the southern extremity of this vast twin island. More often, they move from one cape to another and more rarely are seen in the deep bays and straits. The White Sea harp seals migrate southward in small numbers along the Kara Sea, 1.е., the eastern side of Novaya Zemlya, emerging into the Barents Sea through the Kara Strait. Further advance to the White Sea has not been well traced but nevertheless it may be assumed that the migratory routes in the Pechora Sea run for the most part away from the mainland coast, which the herds of seals rarely approach and, that too, only on the western side of the Russkii bend. They probably do not transgress deep into the Cheshsk Bay; they are, however, more common on the northeastern coasts of the Kanin and partly Kolguev islands. The seals arrive in the White Sea usually slightly earlier and begin to form their first nurseries there. In one or the other sections of it, predominantly on the right bank, these seals are encountered even from November end to early December, their appearance being also asso- ciated with the arrival of the polar cod. The autumn migrations, like the summer-spring ones, are mainly based on the food factor, especially the schools of spawning polar cod. It is therefore not wholly correct to interpret the autumn course of the harp seal to the White Sea as exclu- sively “migration for reproduction” (Freiman, 1939). This phenomenon is evidently more complex (Chapskii, 1961) since it begins to manifest at least 5.5 months before the forthcoming actual breeding season and the phenomenon covers even immature animals. This, if one may say so, complex stage of stereotyped behavior (countless generations have 304 405 well-worked out the course) in which the feeding seasons and the return of the animals to their original sites, guaranteeing the performance of the concluding stages of the annual cycle, has evolved with an accuracy that is perhaps maximal for biological systems. In the past, in very cold years, when drifting ice floes in the Barents Sea came close to the western Murman and the White Sea harp seal rookeries were disposed in the inlet of the White Sea and the entire seal population in the spring period gathered in the inlet and in the comparatively small space extremely close to the Barents Sea bound by the fringes of drifting ice floes, the seals undertook extensive migrations along the Murman coast to the west. In particularly icy, cold springs, they moved not only up to Finmarken, but even to the coasts of northern Norway; turning round Nordkapp, they approached Lofoten and moved even more southward. Particularly memorable are the invasions of seals in 1901-1902 and 1902-1903 when the winters were unusually severe and ice floes were seen even in May close to the western Murman coasts and the southeastern fringes of the compact floating ice floes “lay” on the Murman coast considerably more westward of the White Sea inlet, almost around Kil’din Island. The large concentration of seals pressed by the ice floes at this time toward the Murman coasts caused the distant movement of some herds into Norwegian waters. In the relatively less snowy years, however, when the waters on the coasts of the western Murman were not so intensely cooled and the winter-spring fringes of the ice floes remained far away in the north and east, transgressions of the seals along the Murman coasts were not so massive. Until the beginning of the 1930s, harp seals were quite common in the winter-spring months all along the Murman and were trapped there at many places in nets. Invariably, the herds initially came from the east, from the side of the White Sea and, after wandering, invariably returned again eastward. Their encounters continued usually throughout the winter-spring period, mainly in the spring, up to May, and sometimes even in June. Now, in the 1960s and the 1970s, the nurseries are usually disposed deep in the White Sea while the spring boundary of the drifting ice floes in the Barents Sea runs far northward of the Murman. For this reason and also because of the sharp population reduction on the Murman coasts, harp seals are encountered comparatively rarely. The Newfoundland group of seals begins to riove northward from the regions of reproduction and molt even from the end of April. They move along the fringes of the ice floes blocking the Labrador coast toward Davis Strait and their courses branch somewhere at the latitude of 60° and along 60° W long. A relatively small portion of the population 305 406 continues, as far as the snowy environment permits, to move northward up to the southeastern region of Baffin Island. Another, much larger portion, as far as possible, moves through Hudson Strait into the north- western and eastern regions of Hudson Bay. A third group, representing the greater bulk of the migrants, deviates into the northeast and reaches Greenland roughly at the latitude of 65° or slightly more southward; later, the majority of them proceed northward and, depending on the situation of the ice floes, diverge radially along the straits of the Cana- dian archipelago, along the eastern coast of Baffin Island, and north into Kane Basin. A small group of the population turns southward up to Cape Farewell and, probably running round it, reaches the eastern side of southern Greenland (Sergeant, 1963; Mansfield, 1963). The reverse migrations, traced slightly better, proceed from Baffin Bay along two main routes: one runs along the coasts of Greenland up to its southern tip and from there traverses through the open sea toward the southern part of Labrador. Another group, gathering from the Canadian Straits, runs along Baffin Island and taking the branch from Hudson Strait continues along the Labrador coast. Evidently, at the southern corner of the Labrador Peninsula, which the seals reach by November, both the routes converge and deviate again along the Strait of Belle Isle. Along one of them following the eastern banks of Newfoundland, the animals turn farther southward. It was formerly assumed (Robinson, 1897; Chafe, 1923; Nansen, 1927*, 1939) that the seals rushed toward the great fish banks in the southwest of Newfoundland. New information (Sergeant, 1963) does not, however, confirm this view. Another path proceeds through the Strait of Belle Isle into the Gulf of St. Lawrence. It is not clear where the seals of this herd remain from the middle of January to February end and which courses they take to gather in the nurseries. Exact information is not available on the migrations of the Jan Mayen seals. At the end of the lactation period, the pups drift with the ice floes and later emerge onto the fringes; the rest of the animals wander in small herds to the northeast and southwest of the region of the rookeries. It is difficult to say whether the seals diverge simultaneously in Opposite directions or these directions alternate in some manner. Judging from tagged pups, in some years (for example, 1953), the animals are mainly carried away in one direction toward Denmark Strait; in other years (especially in 1955), mainly in the same direction, but also in the opposite direction (Rasmussen and Oritsland, 1964). The zone of summer-autumn dispersal of the Jan Mayen herd extends along the fringes of the ice floes blocking the eastern coasts of. Greenland, 407 from Denmark Strait (and probably even from the southernmost part of Greenland) to Spitsbergen. In the latter region these seals probably meet with their White Sea kin and sometimes are drawn by them onto the icy rookeries in the White Sea. Transgressions of stray tagged animals from one region to another are known. Reproduction. Although the harp seal forms no harems whatsoever like the other seals reproducing on ice floes, it should not evidently be regarded as strictly monogamous. This concept is in general not appli- cable to the White Sea harp seal. Even in the days preceding mating, no single male picks up a female for mating as happens in the case of the larga. In the same manner, and immediately after mating, the male does not stay with the female nor does he accompany her in the follow- ing days. The males and females converge only for coitus and the entire mating behavior of any given pair, including “heat” and contentions over a female among the suitors, lasts barely for a day. Not long before the onset of the mating period, which becomes evident from the growing activity of the males, the latter gather near the whelped females in small but fairly close rookeries (“monasteries”) as though in anticipation of mating. When the time comes, they are aroused from their torpor and these heavy and awkward animals exhibit surprising activity on the ice floe. They crawl along it, dive into the water one after the other, surface from the open water pools, noisily chase each other, as if in play exhibit surprising dexterity, and again dive into the water. In earlier years it was not always possible to perceive from afar in this melee, even with binoculars, who was chasing whom—whether the males were chasing the females or the suitors were chasing each other. As a result of organized scientific floating stations on ice floes, it has become possible to study the intimate life of the White Sea harp seals quite closely and quite completely. The period of mating is accompanied by powerful excitation of the productive males and rather serious scuffles occur among competitors which can even draw blood (R.Sh. Khuzin). Sometimes the competitors are severely injured with bleeding wounds. The victor remains alone with the chosen female far from the rest of the animals. Coitus proceeds in water (evidently more often) as well as ice floes.!°° In the latter case, it extends for 20-25 min (Yakovenko and Nazarenko, 1971*). It is highly possible that the scuffles taking place between productive males is a consequence of the acute competition among them arising from the 106 There are some other views, although hardly substantiated, that the animals do not mate in water but only on ice floes (L. Popov, 1966). 306 306 408 quantitative disproportion between males and females due to long-time preferential killing of the latter. The mating period among White Sea seals sets in at the beginning of the first week of March and terminates evidently around the 20th of the same month. Thus the total duration of the mating period is not long—about 1.5-2 weeks for most of the animals. However, there are various views on this subject: not more than 2 weeks (Surkov, 1957), 10-12 days (Rasmussen, 1957), about 2 or 2-3 weeks (Dorofeev, 1960; L. Popov, 1960), and 10-20 days (Freiman, 1939). The views of some about a late mating season (as against the average). from March end to mid-April (G. Nikol’skii, 1933) and more so that mating occurred after the cessation of molt (Sleptsev, 1949) are erroneous. Even in 1902, the first of the mating pairs on ice floes were noticed on March 10 (N. Smirnov, 1903). Most of the whelped females mate even before suckling of the pups has ceased—roughly in the second half of the period of lactation, closer to its end. The earliest recorded date of mating is March 4 (Sivertsen, 1941). Rarely, the last of the winged males was observed hanging around suckling females in 1948 on March 20. Twenty years later, these dates were rendered more accurate by researchers on the floating research stationsin the White Sea (Yu.I. Nazarenko, L.A. Popov, M.Ya. Yakovenko). The first pair in coitus was observed on March 10 and the last on March 24 (Yakovenko and Nazarenko, 1961). At the beginning of April the testes of the White Sea harp seals revealed no mature sperm (Surkov, 1957). Fig. 167. Mother suckling her pup. White Sea, March, 1967 (photograph by M. Ya. Yakovenko). 409 The conclusion of the mating period is marked by the quiescence of the males, cessation of scuffles among them, a distinct coolness toward the females, and their departure with the young ice floes. By this time (in the second half of March), the males begin to form different herds (rookeries) initially consisting exclusively of adult males. These very inti- mate and massive groups, extending for kilometers along the edges of the ice floes, along the open water pools and partitions, mark the com- mencement of a new cycle of life, i.e., the molting period. The period between mating and whelping is about 11.5 months. In fact, however, embryogeny occurs with a fairly prolonged lag in the stage of the blastocyst. The growth of the embryo commences roughly 2.5 -3 months later when implantation sets in. Excluding this duration, the actual development of the fetus extends for slightly over 8.5-9 months. Until June, no distinct signs of gestation are seen in the womb (Nansen, 1924, 1939; G. Nikol’skii, 1933; Sivertsen, 1941; Fisher, 1955). Having attained maturity, the females usually undergo parturition every year although there are no accurate data on barrenness to date. The view was expressed before (N. Smirnov, 1903, 1927, 1935; Kulagin, 1929) that the females of the White Sea herd do not give birth to young ones every year.. However, this view was based on incomplete data and has not been adequately proved. A thorough analysis of the generative organs leads more to a contrary conclusion (С. Nikol’skii, 1983; Chapskii, 1963): most of the whelped females mate again in the same year. In any case, there is no large-scale barrenness among the White Sea females; it does not exceed 10-15% of the mother population in this herd.!°” The proportion of barren females among the Canadian population too is not high though it has been put at 10 to 16% (Sergeant, 1966) and even 20% (Fisher, 1952, 1955) for the population of the Gulf of St. Lawrence. According to the morphological indices of age used at present, males as well as females retain the reproductive capacity for over 15 years after attaining sexual maturity. The oldest of the investigated females that had whelped in a season were aged 20 years while the productive males included even 25-year-olds (Fisher, 1952; Rasmussen, 1957; Sergeant, 1966; Yakovenko, 1967). Thus it may be assumed that males as well as females retain their reproductive capacity up to 25-30 years although under conditions of extremely intense hunting it is indeed a rare animal that attains this age (fraction of a percentage). Unlike the other arctic seals, harp seals bring forth their offspring in strictly localized sections of the winter-spring range, which for the 107 Tn calculating the population dynamics, the barren animals among the females were frequently not taken into consideration (Yakovenko, 1967). 308 410 species as а whole number four: (1) Gulf of St. Lawrence; (2) region to the north-northeast of Newfoundland Island close to the Strait of Belle Isles (seals of the Canadian population); (3) region of Jan Mayen Island; and (4) the White Sea. The ice regime characteristics in a given year or even a very prolonged period of this regime exert an influence on the actual location of the nurseries, which therefore vary from year to year within certain limits. These locations also change in the course of a given season under the influence of drifting ice floes. Seals of the White Sea herd form nurseries depending on the condi- - tions of the ice formation either in the neck or in the central basin (more often on the threshold of the neck and sometimes even in Dvina Bay) or in the inlet of the White Sea. The possibility in some rare years of a small proportion of mothers giving birth in Cheshsk Bay (Danilevskii, 1862; Zhitkov, 1904; Suvorov, 1913*; Vinogradov, 1949; Dorofeev, 1956) has not been confirmed and should be regarded as erroneous. Over the last 30 years the formation centers of White Sea nurseries have undergone significant changes. In the 1920s, rookeries were seen almost exclusively in the inlet comparatively close to the fringes of the ice floes in the Barents Sea. From the mid-1930s (1935 - 1936), however, they began being formed considerably farther away, toward the south- west, i.e., in the neck and the central basin. The desire to provide for the juvenile adequate conditions in which it can grow well until it becomes self-supporting has compelled the females to select regions much before parturition with sufficiently large and stable ice floes capable of with- standing compression and hence able to serve as a reliable substratum for the pup’s residence for many days. Another invariable condition for the formation of the nursery is the presence of a fairly dense network of open water pools, fissures, and gapes, which permits the female to approach the chosen ice floes while maintaining contact with the water (Fig. 165). The sum total of these conditions together with the herding ten- dency (which to a large extent is probably caused by these factors) and also the sharply manifest seasonal migrations due to concentrations of the population in a restricted section of the range, necessitate that ges- tating females gather periodically in large numbers in the regions of reproduction. Immediately before whelping, they crawl onto the ice to form nurseries. A nursery may run into several tens of kilometers and the area several hundreds of square kilometers. Initially, there is only one nursery (rarely two or more) but over time, under the influence of drifts and shuffling of the ice floe, the single massive nursery is fragmented. As a result, at the end of lactation, when the suckling females leave the young ice floes, several individual groups of juveniles are seen in what 411 was once a large single nursery. Usually, by this time the juveniles are transported by ice drifts for long distances from the site of whelping. Under the influence of the prevailing surface flow from the White Sea into the Barents, nurseries formed initially in the central basin or in the neck are gradually transported into the inlet where they cease to exist. There are, of course, exceptions to this rule. An example is the situation that prevailed in 1937 when ice floes with pups in the final stages of molt were transported far westward into Kandalakshsk Bay. The very same phenomenon, far more acutely, was repeated in 1966. This clarifies the view expressed by Acad. I. Lepekhin (1805) about some retreats in the spring migrations of the White Sea harp seal juveniles when they attempted the inner regions of the White Sea. The animals are disposed unevenly in the nurseries. The gestating females usually select large ice floes and are disposed initially along their peripheries. In time, they often crawl toward the center of the ice fields and thus their distribution becomes more uniform. On the whole, however, within the nursery the animals occupy not all the ice floes but only some of them, usually those along the open water pools and fissures. Therefore, the rookery does not appear compact but has alternating occupied and vacant sections on the ice floes. Ice floes with seals densely or sparsely disposed on them alternating with “vacant” ice floes, cover an extensive area away from the coastal regions of the sea. The lesser the density of the rookery, the wider its area, and vice versa. Further, much depends here on the population of the animals, which is affected by the magnitude of hunting. Shortened periods of whelping are a characteristic feature. Among the majority of White Sea mothers, parturition extends for not more than a week, from the last few days of February to the first few days of March. In 1968, by March 4, almost all the gestating females had undergone parturition. П is the rare female that gives birth earlier than this period (but nevertheless not before the 20th of February) or slightly later. A survey of the animals in the month of March revealed that of the total number of white pups (916), the newborn (aged less than a day) on March 5 were only 12, or 1.4%; on March 7, 0.8%; March 9 about 0.4%; and on March 12 about 0.3% (Khuzin, 1970). In fact, some very late dates of the appearance of newborns, for example March 18 and 31 and even mid-April (Sivertsen, 1941) have been recorded but such instances are totally sporadic.1°* Evidently, taking into consideration such anomalous instances, the total duration of the whelping period is sometimes put 108 In 1966, one white female pup with a firm hair coat was detected as late as April 26 (M.Ya. Yakovenko). 309 309 412 at about two months (Smirnov, 1935) and even extended to 2.5 months (Sivertsen, 1941). However, it is not correct to include in this period the extreme, absolutely atypical instances. The exact moment of birth is usually beyond observation although one such instance has been described (Sivertsen, 1941). Some members of our hunting expeditions affirm unanimously that births occur very rapidly and, as a rule, with no particular birth pangs. Nevertheless, an instance is known of a dead, full-term pregnant female on the ice floes in the White Sea in 1959 (M.Ya. Yakovenko). Instances are more common of stillborn pups or those that perished soon after birth. The birth of twins has not been established although, a century ago, it was stated that the females produce one pup, often two or possibly three, based on the fact that hunters sometimes found up to three pups on the stretch of an ice floe around a single female (Brown,!” 1868). Similarly, rare instances of two pups cared for by a female equally well could point to such a possibility but only indirectly (M.Ya. Yakovenko). Growth, development, and той. The newborn is covered with a long dense, silky hair coat that is yellowish-white with a faint green tinge; hence a just-born pup is called a greenling by local hunters. Its length at birth averages 83 cm (in a straight line, Lcv) and along the dorsal surface Fig. 168. White pup of the harp seal. White Sea (photograph by A.V. Yablokov). 109 Significantly, however, this author pointed out that not a single hunter saw more than two fetuses in one womb. 311 413 (Lc) about 90 cm. The subcutaneous fat layer in a newborn is negligible or almost non-existent. The weight is generally about 7 to 8 kg. In the first few days after whelping the females are particularly solicitous of their pups and remain with them almost constantly on the ice floes. The milk composition of the White Sea mothers, according to the latest data (Khuzin, 1970), reveals considerable variation: fat from 13.5-40.2% (average 29.35%), nitrogen 1.21-2.95% (average 1.76%), and protein content from 7.7-18.8% (average 11.17%) [see also Table 17]. Suckling is frequent, not only in the day but even at night. The suckling mother lies on her side and the pup alongside her stomach finds one or the other teat without difficulty. At times, even an approaching ship cannot coerce a suckling mother to abandon her pup. Sensing danger, she will crawl away attempting to carry the pup with her, then abandon it to dive into an open water pool or through an air hole in the ice but immediately emerge on the ice again. Having overcome her fright at the ship, she comes close to the pup and again dives only to reappear on the surface merely a few tens of seconds later. Confronted with imminent danger, she will not leave the pup out of her sight. Even in the early period of its growth, the pup exhibits quite good mobility and traverses considerable distances, evidently not so much in search of its mother, as to find protection from wind, to which, lacking adequate fat reserves, it is initially very sensitive (Dorofeev, 1939). Per- haps, too, there is a natural need for movement. At favored sites, well protected from winds by an icy projection or the cornice of a hummock, the pups remain for long periods. Their prolonged resting at one place, due to the effect of body heat, forms a basket-like oval depression on the ice. Inside the depression the pup is always dry. A few days after birth the green tinge of the pup’s coat disappears and it turns white; hence hunters refer to it as a white pup. It gains weight rapidly and by early in the second week after birth has accumu- lated 9-11 kg of subcutaneous fat; its total body weight now goes up to 17-18 kg. The body length at the end of the white-pup stage aver- ages 95-96 cm (Lcv) ог 102-103 cm (Lc). The firm white hair coat is sported for roughly a week or one to three days longer (Sivertsen, 1941; Chapskii, 1964; M.Ya. Yakovenko, Yu.I. Nazarenko); the assumption of a long duration of this period, from 9 to 14 days (Dorofeev, 1936), is hardly correct. The commencement of a perceptible loosening of the neonatal hair coat in the White Sea seal occurs roughly on March 8-9. Initially, the hair begins to weaken slightly, loses its original brightness and purity of color, and acquires a gray bloom. The gray bloom is often due to the thinning of the white hair coat (as a result of the pup’s growth and 310 414 Fig. 169. Well-fed white pup of the harp seal. White Sea, early middle 10-day period of March, 1965 (photograph by A.V. Yablokov). its dark skin showing through) and the growing new, already pigmented coat. Often the guard hair of the white coat turns gray at the base and the pigment penetrates there as long as the hair bulb is not atrophied (M.Ya. Yakovenko). This state extends for a week, after which the next stage of molt sets in (Fig. 170). By this time the white cover has become even sparser and the growing stubs of new hairs can easily be seen through it. A few days later, molting becomes even more evident: the white hair that has turned gray falls out in large clumps almost simultaneously from the head, hind and fore flippers, exposing a darker, short, and rigid hair coat. Following this and almost simultaneously, rapidly growing bald patches on the back and tail appear. The last hair to molt is that of the ventral side and body flanks. The better fed the pup, the more rapid the molting of the white coat. In underfed pups, who have lost their mothers early, molting is not only impeded or even halted, but acquires a different sequence: the primary coat is retained on the flippers, tail, and head longer than on other parts of the body. Lactation ceases in a normally molting pup at three weeks of age (Fig. 171). The normally fed molting pup at this time is, on average, 106-108 cm (Lev) or 114-115 cm (Lc). Its total body weight averages 32-33 kg, and some even 38-40 kg or more; the subcutaneous fat with the skin (blubber) weighs 22-23 kg. The fully molted pup is called a 311 415 Fig. 170. Normally fed molting pup and a pygmy (sick pup of the harp seal). White Sea (photograph by M.Ya. Yakovenko). 312 gray pup. The hair coat is extremely similar in color in yearlings and two- or three-year-olds but differs in greater density and softness; the older juveniles also differ in body proportions: relatively larger head and longer flippers. The main gray background of the skin among gray pups is darker on the dorsum with brown angular, predominantly small spots scattered here and there. On attaining roughly one month of age, the pup under normal conditions enters the water for the first time and commences an independent life. The whelped females no longer spend as much time on the ice flow as necessitated before. Their residence outside water follows a definite pattern ordered by periodic lactation, depending on the time elapsed from the moment of parturition, and the weather factors.!!? On pleas- ant, quiet, and sunny days, 85, 90, and even 94% of the total mother population can be seen on the ice surface in some sections of the rook- ery. '!! On the contrary, in stormy weather, this index drops to 36-45% 110 Detailed information on the periodicity of lactation and the general regime in nurseries was provided by the floating ice stations organized from the end of 1966 directly in the rookeries. Groups of investigators landed by helicopter on the White Sea ice with tents and the requisite equipment and carried out extremely interesting studies (L. Popov, 1966, 1967; Khuzin, 1970; Yakovenko, 1970; Yakovenko and Nazarenko, 1971”). 111 According to others (L. Popov, 1966), however, on such fine days, 45-55% of the whelped females were seen on the ice during the day and up to 70-80% in the evening. 312. 313 416 Fig. 171. Normally molting pup of the harp seal (photograph by M.Ya. Yakovenko). and on March 7 even further, to 11% (Yakovenko and Nazarenko, 1971*; L. Popov, 1966). Even in the early period of lactation, not all the whelped and lactating females are seen simultaneously in the nursery although they maintain a definite feeding cycle. The maximum number on the ice floe varies at different times: evening, midday, or morning hours, but more often peaks at midday and evening. The maximum percentage rarely approaches as high as 85-90%; it is usually lower, around 70%. It decreases in the course of time and the residence of the suckling mothers on the ice floe dwindles. In the first few days immediately after parturition, they remain on the ice floe for long periods and feed the pups most often at 3-3.5 hr intervals even during the night. At this time of lactation, the pup receives roughly 0.75 liter of milk in one feed. Later, as the pup grows, feeding becomes less frequent (4-5 times) and the pup then suckles a larger amount of milk per feed (about 1.2 liters). At this time the lactation schedule in most cases is as follows: (1) at 6:00-7:00 a.m., (2) 10:00-12:00 Noon, (3) 4:00-6:00 p.m., and (4) 9:00 - 11:00 p.m. Further, suckling continues even after midnight until the morning feed, as evidenced by the cries of pups and their quiescence after sometime, evidently after satiation of hunger (Yakovenko and Nazarenko, 1971*; Popov, 1966). The daily requirement for milk in 314 313 417 the first few days is about 4.5 liters, which later goes up to 5.8 liters (Yakovenko and Nazarenko, 1971*; Popov, 1966). The milk of the harp seal contains 42 -44.5% fat and 8.4- 12% pro- tein (Sivertsen, 1941; Dorofeev, 1960). Table 17 shows the composition of milk (10 samples) according to the more accurate data of Khuzin (1970). With such a high level of well-being, the daily growth of the pup in a short period of time—from greenling to white pup beginning to molt intensely—averages 1.4-2.3 kg (Yakovenko and Nazarenko, 1971*). Pups which have completed normal lactation and molt grow heavy and round like dumplings; they remain for one more week and sometimes longer on the ice floe until the white wool is completely shed and the gray coat begins to grow. During this period of starvation, extending for about one-and-a-half weeks, the normally fed pups lose, on average, about 5 kg of body weight, roughly averaging 0.5 kg (Chapskii, 1964) or 0.6-0.7 kg (Yakovenko and Nazarenko, 1971*) per day. Such a weight loss in no way affects the future well-being of the young one on its taking to independent living in water. However, not all pups receive normal feeding; pups are encountered which have been abandoned early by the suckling mothers or have lost their mother for other reasons (Fig. 172). If at the age of one week, the fat deposition in the pup does not exceed 6-7 kg (and their total weight not over 10-13 kg or slightly more), cessation of lactation inevitably leads to emaciation and later to the pup’s death. The subsequent growth of the animals proceeds more slowly but con- tinues not only in the first few years, but right up to the time of attaining sexual maturity though at a very slow tempo (Chapskii, 1952; Yakovenko, Nazarenko and Timoshenko, 1964*; Yakovenko and Nazarenko, 1967; Khuzin, 1967; and others). Earlier there were no reliable morphological age criteria and the period of attaining sexual maturity was reported variously as: not before two years of age (Nansen, 1924, 1939), at three years (N.A. Smirnov, 1927), not before four years (Bartlett, 1927*; Nikol’skii, 1933), and even five years (Degerbol and Freuchen, 1935). At present, when the age of Table 17. Composition of the milk of the harp seal, % (Khuzin, 1970) Fat Nitrogen Protein Ash Min Max Mean Min Max Mean Min Max Mean Min Max Mean 13:53) 13922400 2935 [22:95 176 ty Le Ol 1117: 01147 O82 314 418 Fig. 172. A typical underfed pup (dwarf) of the White Sea harp seal with remnants of embryonal (white) hair coat on the head and flippers. White Sea (photograph by M.Ya. Yakovenko). a given animal can be accurately established within a year, the age at which reproduction commences has been well substantiated. The period of maturity among the female harp seals extends for several years. In the White Sea population some females ovulate for the first time even at the age of three years, with the percentage of such early maturing females reported variously as 5-12 to 15.5 (Yakovenko and Nazarenko, 1967) or 30% (Chapskii, 1963). According to these same data, about 50% mature by the age of four years but slightly less (36%) according to other authors (Yu.K. Timoshenko). The rest of the females mature at five or even six years (Khuzin and Timoshenko, 1968* ) but some, however, at seven years (Fisher, 1954; Rasmussen, 1957; Sergeant and Fisher, 1960; Sergeant, 1966; Yakovenko and Nazarenko, 1967; R.Sh. Khuzin). The average age of the onset of sexual maturity among the female White Sea population in the second half of the 1960s was established roughly at 4.5 years (R.Sh. Khuzin). Evidently, how- ever, the average age of females that have attained maturity varies within certain limits, depending on the intensity of hunting, especially on the replenishment of the mother population by those of the younger gener- ation that have escaped the hunters. [t is significant that over the decade from 1953 through 1962, the average age of the Newfoundland females that had attained sexual maturity fell from 5.5 to four years (Sergeant, 315 419 1966). Males develop somewhat more slowly and their maturity sets in, in most cases, at five years of age (Chapskii, 1963). According to some other data (Yakovenko and Nazarenko, 1967), even four-year-olds are mature. On the other hand, Newfoundland males become capable of mating generally at seven to eight years of age (Fisher, 1954). The uterus (without vagina) of the lactating females two months after parturition weighed about 200 g. The weight of the paired ovaries (their average) in such females varied markedly, from 6 to 10 g; the testes of adult males in the breeding season without the spermatic cord weighed 135-300 g and 200-370 g with it; their length varied from 10- 14.5 cm. The growth intensity of the White Sea harp seal (males) is shown in Table 18. It can be seen from Table 18 that the increment in the first five years is quite marked (sharp variations are undoubtedly due to the small number of animals measured). Later, however, it is very small and can be perceived only in a series of animals. Similar age variations are seen among females but their growth tapers off slightly sooner (roughly by a year), after which their length increase is very small. Molting in animals of all later generations, commencing from year- lings, is an extremely perceptible periodic phenomenon in the life of the harp seal; at the same time, it has played an extremely important role in hunting since it serves as a no less powerful impulse than reproduction for the concentration of large numbers of animals in certain periods in relatively small sections of drifting ice floes. Hunters attempt to reach such dense molting rookeries as well as the nurseries. Chronologically, molting follows the periods of parturition, lactation, and mating, extend- ing among the White Sea seals from the last 10 days of March up to mid-May. This period does not cover the duration of individual shed- ding of the hair coat (which in some animals is much shorter) but the duration of the overall period of molt among animals in a population. The animals setting out to molt initially concentrate in nearly the same areas (or in their proximity) where the nurseries existed quite recently. In the 1960s, the molting nurseries were quite often encountered in the neck region of the central basin of the White Sea. Quite often, espe- cially when access to the basin was delayed, the animals migrated to the northeastern regions of the sea into Mezensk Bay and into the inlet (or collected there). By mid-May the animals departed (or drifted away) from there to the ice floe fringes in the Barents Sea. Sometimes the rookeries were formed opposite the western sections of Tersk coast. In 1964, a large rookery was detected even close to the Karelia coast, in the region of Gridin (M.Ya. Yakovenko). 420 315 Table 18. Increase т body length (Lc) of the White Sea harp seal (males) in relation to age (Yakovenko ef al., 1964*) Age, years Number Range, cm Average, Annual measured cm increment, cm 1 8 130-150 138.1 2 4 152-169 161.0 22.9 3 5) 156-172 165.0 4.0 4 6 166-182 174.7 Od 5 16 170-191 180.9 6.2 6 57 162-196 184.7 3.8 7 70 170-207 186.5 1.8 8 42 162 - 201 188.6 2.1 9 45 161-205 189.3 0.7 10 14 181 - 203 192.9 3.6 т 11 185 - 198 193.4 0.5 12 9 178-200 189.6 3.2 13 3 196-202 199.0 9.4 14 2 200 - 203 201.5 25 15 2 185 - 190 187.6 —4 The seals select various types of ice floes for molting rookeries with 316 the only proviso that they should not drift rapidly toward the open fringes and at the same time provide access to the water. The animals lie along the edges of large- and medium-sized ice floes forming a live border with their bodies lying close together. Such ice floes are separated from each other by narrow or sometimes broad open water pools or partitions or gapes; the animals lie on either side of these, presenting a picture of com- plex branched strips or bands. Such a deposition is highly characteristic of the early male rookeries. The animals are disposed more haphazardly on the so-called conglomerate ice, i.e., predominantly on small broken ice pressed together by winds or currents and held together by frost. The density of disposition, the size of the rookeries, their number, and composition vary widely. The animals are sometimes concentrated in a single huge rookery or sometimes divided into several rookeries of much smaller size. Under favorable conditions, a rookery can exist continuously for quite a long time. An extremely characteristic feature of the molting rookeries is the fluid state of their age-sex composition. The first (excluding pups) to molt are the mature males and the first molting rookeries consist almost exclu- sively of animals with a wing pattern. At the commencement of April immature animals of both sexes except under-yearlings (gray animals) join them. After shedding the embryonic white coat, the under-yearlings 317 421 molt no more in the first year. However, at this time, young animals (aged one to four-five years still preserving infantile coloration) constitute no more than 15%. The proportion of mature females in the rookeries at this time runs into a few percentages but in any case does not exceed 10%. In the last 10 days of April, the immature animals increase in num- bers up to equal those of animals with a wing pattern and the percentage of the adult females slightly rises. According to the situation prevailing in the 1920s and the 1930s and later (Dorofeev and Freiman, 1928; Surkov, 1957), at the end of April and in the first few days of May, the percentage ratios existing in the molting rookeries between the main constituent animals somewhat reflect the potential ratio characteristic of the herd: adult males and adult females 25% each and immature animals 50%. These figures can hardly be regarded as factual since females predominate among the animals caught. There is reason to believe that the actual position is considerably more complex. The above age-sex ratios in the molting rookeries at April end to early May represent a regular process observable even at present. It is graphically depicted as the crossing of curves showing the different peri- ods of molt among the different constituent animals of the herd. This is strikingly demonstrated, especially by the data for the early 1960s, when no one doubted the distinct disproportion between the number of productive males and mother animals (Yakovenko and Nazarenko, 1962; Yakovenko, Nazarenko and Timoshenko, 1963; Yablokov, 1962; and others). Consequently, from the equilibrium (or similarity) of the percentage ratios between males and females in the late molting rook- eries, no conclusion whatsoever can be drawn about the actual balance between the different groups in a herd. The final period of the molting rookeries of the White Sea herd was studied less fully since the seals desert the White Sea and hunting ceases. It is usually assumed that the very late rookeries consist mainly of immature animals of both sexes and adult females. However, even in the first of the molting animal concentrations along the fringes of the western part of the White Sea inlet, young ones were in a majority from April 5 (Khuzin, 1970) and observations of the migrating animals point out no perceptible division among them into age groups as the animals advance toward the White Sea (Beloborodov, 1969). Some direct observations (Sivertsen, 1941) also point to a large variation of the composition of the rookeries even in the last 10 days of May. The process of molting is quite prolonged. The old hair coat con- tinues to be sported until the new hairs grow to half the length of those 422 shed. However, the implantation strength of the older hairs gradually decreases. During molt, whiskers too are shed along with the hairs; evi- dently claws commence growth at this time. The horny layer of the epi- dermis is also shed in small and large strips simultaneously. Some new data available only recently suggest that molting among the various groups of seals entering the White Sea occurs at different times in different regions. It has been found that among an overwhelming majority of the animals residing in the open sea, especially the adult males, molting was at its peak on April 27 while molting of animals arriving later in the inlet region had only just commenced (Khuzin, 1970). Enemies, diseases, parasites, mortality, and competitors. From among the vertebrates, the potential enemies of the harp seal in water are the Greenland shark (Somniosus microcephalus) and the killer whale (Orci- nus orca) and on ice floes, the polar bear. Raven and arctic fox can also be listed among them. The destructive role of all these animals is extremely variable. The Greenland shark and the killer whale pose the greatest danger to the White Sea herd. However, the shark is confined mainly to the deep sections although it is capable of surfacing and, what is more important, it is generally rare in the White Sea. Seal remnants were detected in the stomach of Greenland sharks caught on Murman and on Kanin Island (Breitfus, 1906; Smirnov, 1935; Andriyashev, 1954). The role of the killer whale has not been supported by actual data since it is not actually caught in the Barents Sea; yet it cannot be ruled out as a potential enemy. In fact, in the eastern part of the sea, especially along the coasts, this whale is encountered extremely rarely during the migration of the White Sea harp seal. The polar bear, by and large, poses no real danger to the seals of our herd for several reasons. It has not been sighted, as far as people can remember, in the zone of reproduction and is extremely rare in the high latitudes of our western Arctic. Further, the bear has access only to seals on ice floes where the harp seal is almost nowhere seen in the summer. Foxes can probably attack the newborn but they are hardly even seen in the nursery zones; by the time the rookeries drift toward Kanin Nos, the young one has grown considerably and molted and the fox is no longer capable of attacking it.!!? The fox, if it strays onto an ice floe, has to satisfy itself under the best of circumstances with only the frozen placenta. In this respect, ravens cause serious damage and are often encoun- tered in the nurseries. They feed mainly on the placenta or the dead 112 See: V.G. Heptner et al. Mammals of the Soviet Union, vol. 2, pt. 1. 318 423 pups but, in rare cases, may even attack live pups, primarily undergrown, sick, underfed, and hence incapacitated ones. Regardless, the skeletons of pups with a pecked head have been encountered from time to time in the White Sea rookeries. Sometimes, large gulls also indulge in this activity. In some years the number of dead pups with pecked heads has reached 25% of the total dead pups recorded in a given section of the nursery (Rudakov, 1936). In other years, however, as in 1947, no such instances were reported (K.K. Chapskii). An instance of attack even by ermines has been recorded (L. Popov, 1955). Among the other more common factors for pup mortality are: (1) defective births (birth with the amniotic sac and placenta) and other instances of still births; (2) freezing and death due to emaciation; (3) crushing by hummocking ice floes; and (4) washing away by waves. Pups in the White Sea perish relatively more often due to the second and third factors though sometimes the first factor as well as the second play a predominant role (Yakovenko and Nazarenko, 1962). The emaciation of a pup transformed into a starveling is caused by the cessation of lactation, due to the loss of its suckling mother. If the pup ceases to receive milk at a weight of less than 15 kg, it cannot survive (Yakovenko and Nazarenko, 1962). Pups whose lactation has been interrupted even somewhat later do not attain normal growth. The destructive effect of the hummocking of ice floes, which crushes the pups, is very difficult to estimate since the dead ones buried under the vast ice escape attention. Such causes of death can be judged only from instances when the crushed pup is partly visible among the broken ice (or the tracks of the animal are lost under it). Such finds are very rare and do not reflect the true scale of mortality, which is perhaps relatively high since hummocking in the White Sea is a fairly frequent phenomenon. The washing away by waves of pups which have yet to complete lac- tation (or their falling into the water while negotiating on the ice floes), causing excessive cooling of the body, is no mean factor in their mor- tality. However, at present, when the nurseries are formed in the open waters of the White Sea and not close to the fringes (as, for example, in the 1920s), very few pups perish for this reason. In the other locations of the White Sea rookeries, however, such deaths of pups represent one of the most significant factors (N. Smirnov, 1927). Yet situations that prove fatal to the pups do arise from time to time in the White Sea nurseries. One such is when the young ice breaks up intensely during compres- sion and later greatly expands, as happened in 1967 (Г. Popov, 1971*). This factor is of equally great relevance to the infant mortality of the 319 424 5 Jan Mayen and eastern Newfoundland (in the “front” zone) populations (Nansen, 1924, and others). There is yet one more, albeit episodic, very intensely manifest natural factor of infant mortality. In years of anomalous wind directions in the White Sea, when the persistent northeastern and even northern winds prevent the drifting of ice floes into the inlet zone in spring, the well-fed young find themselves carried away by the young ice floes in the opposite direction, i.e., into the western part of the White Sea (Fig. 173). As soon as the ice floes break up and thaw, pups scatter in small groups, at times all along the periphery of the sea, right from the uppermost part of Kandalakshsk Bay to the Onega River and Arkhangel’sk. The weakened and emaciated pups, not finding their natural food, become incapable of independent survival under the unusual conditions and often even crawl to the coasts, thus becoming easy prey to any land predator, even man. ZONE OF RECOVERY ® ARKHANGELSK Fig. 173. Some sites of tagged White Sea harp seal pups in 1966 scattered along the periphery of the White Sea (data of the Polar Research Institute of Sea Fisheries and Oceanography, PINRO). 319 425 Such a situation prevailed in particular in 1966 and probably led to the large-scale death of pups (Tambovtsev, 1966; Bianki and Kar- povich, 1968; Beloborodov and Potelov, 1968). Such instances undoubt- edly existed even in earlier times although not very often. The overall magnitude of the natural mortality of White Sea pups during their residence in the nurseries is not amenable to precise compu- tation since the newborn can be washed off, crushed during hummocking, fall into water, and the dead buried under snow. The figures cited below should therefore be regarded as extremely approximate. For the 1930s, the figures cited at 10 to 11% are slightly exaggerated (Rudakov, 1936) and for the 1940s, on the contrary, the figure of less than 5% is an underestimate (K.K. Chapskii). The actual figure is obviously 5 to 7.5%. The overall mortality of the pups in their first year for the White Sea herd was taken at roughly 20% in the calculations of population dynam- ics (М. Smirnov, 1928; Р.А. Rudakov, К.К. Chapskil). The mortality in the subsequent age groups has been roughly assumed at 9% in the sec- ond year, 6% in the third year, and 5% each in the fourth and fifth years; the average is 7.5% for the mother population and the produc- tive males (K.K. Chapskii). These same indices were used in the latest calculations of the population dynamics of the White Sea harp seal for 1970 (Yakovenko, 1967). For seals of the western Newfoundland herds, the Canadian investigators put the approximate extent of natural mor- tality in the first year at 50% of the generation (Rasmussen, 1957). The average index of natural mortality for the entire population reproducing in the northeast of the Strait of Belle Isle has been assumed at roughly 15% (Fisher, 1952). There are no clear references to the diseases they cause but seals do represent a source of disease for the people around them (called “chingi”). When skinning the killed animal to remove blubber, infec- tion of any wound produces an extremely serious inflammatory purulent process [seal finger] leading ultimately to abnormal bone deformation (or damage to the other parts of the limbs). The helminth fauna of the harp seal has not been studied equally well in all parts of its range. Evidently the helminth fauna was better studied for the harp seal inhabiting the White Sea and in the region of Jan Mayen (Vagin, 1933*; Skryabin, 1948*; Mozgovoi, 1953; Delya- mure, 1955; Delyamure, A. Skryabin and Alekseev, 1964*; Delyamure, A. Skryabin and Treshchev, 1965*; Khest, 1932*; Zhuar, 1935*; Stankard and Shoenborn, 1936*). Ten species of helminths and two larval forms are known among harp seals. The trematodes Orthosplanchnus arcticus and Pseudamphistomum truncatum infect the gall bladder and the bile 320 426 ducts of the liver. The cestodes Diphyllobothrium cordatum, D. schis- tochilus, Diplogonoporos tetrapterus, Diphyllobothriidae g. sp., and others are sometimes parasites in the intestine. The nematodes Contracaecum osculatum, Phocascaris phocae, Ph. cystophorae, Anisakidae g. sp., and others infect the stomach and intestine, with as many as a thousand present at one time; Terranova decipiens is encountered only in the intes- tine. The acanthocephalan Corynosoma strumosum infects the intestine (V.V. Treshchev). A comparison of the helminth fauna of the White Sea and Jan Mayen harp seals showed that acute differences along with features of similarity exist (Table 19). Of the 121 Jan Mayen animals studied (V.V. Treshchev), 113 (84.8%) were infected. Newborns (seven) were free from helminths; however, the yearlings as also almost all animals of other age groups were 100% infected. The most infected body parts were the stomach (in 91.7% of the animals) and the small intestine (87.6%), and less frequently the duodenum (29.7%). It was established that five- to eight-year-old White Sea harp seals are infected more severely than other age groups. An average of 131 helminths were found in the infected animals. According to the data of investigations covering 169 White Sea harp seals (V.V. Treshchev), 95 (56.2%) were infected. Newborns with white hair coat and normally molting pups were not dissected for helmintholog- ical studies but two of the 72 moited pups (gray ones) aged 1.5 -2 months were found to be infected with immature nematodes. All animals older than one year (97), except for two, were infected (97.7%). The seals of the White Sea herd were more severely infected with helminths at the age of 13-16 years. Often the stomach (in 56.2% of the animals) and the small intestine (52.0%), and less frequently the duodenum (44.4%) were infected. An average of 341 helminths was found, indicating the severity of the invasion, which led to the formation of innumerable unhealed sores up to 20 mm in diameter on the stomach walls.'!° There are no serious competitors of the harp seal in the White, Bar- ents, and Kara seas with regard to food. In the pelagic regions in sum- mer, stray ringed seals are sometimes found along with this species on the polar fringes. In autumn, in the coastal regions, especially on Novaya Zemlya where large masses of polar cod gather for spawning, ringed seal and bearded seal appear along with the harp seal. However, they rep- resent no competition since fish food is abundantly available there. A similar situation prevails later on the northern coasts of the White Sea. In the pelagic sections, however, in winter and early spring, other seals ПЗ A short review of the helminths of harp seals was made by the staff of the helmintho- logical laboratory of Crimea State University under the guidance of Prof. S.L. Delyamure. 427 320 Table 19. Comparison of the helminth fauna of the harp seals of Jan Mayen and White 321 Sea herds Helminth From Jan Mayen From White Sea region region Orthosplanchnus arcticus ar Pseudamphistomum truncatum Diphyllobothrium cordatum Diphyllobothrium schistochilus Diplogonoporus tetrapterus Diphyllobothriidae g. sp. ++++ Contracaecum osculatum Phocascaris phocae +++ Phocascaris cystophorae Terranova decipiens Anisakidae g. sp. Corynosoma strumosum — —> + + are not found near the nurseries and molting rookeries. Only the white whale can be seen rather frequently in the large open water pools but predominantly in the central basin; these whales are not, however, seen right within the rookeries. Population dynamics. Man’s hunting activity is almost the lone factor responsible for the population dynamics of this seal. Until the beginning of this century, hunting in the White Sea was relatively low, the herd reduction was much less than the births, and the growth of the herd was therefore good. In the first 15 years of this century, hunting intensified noticeably. and the growth of the herd slowed down as a consequence. From the 1920s, the population began declining sharply and continued to fall almost up to the early 1940s (see pp. 391, 430). During the Great Patriotic War (especially in 1942, 1943, and 1944), further decline in the population was arrested and the herd slightly improved but the 3- or 4-year break was inadequate. The subsequent fresh hunting spree, although not so intense as before 1940, caused further population reduction. From 1965, when the White Sea herds quite clearly became very lean, the use of ships for hunting in the White Sea was banned for five years. An exception was permitted only in favor of the coastal collectives who were assigned the right to hunt for 20,000 pups a year. Additionally, nearly one-half this number was killed outside the White Sea boundaries by the Norwegian hunting fleets. 322 428 The sharp reduction in hunting activity stopped the degradation of the herd but the measures did not produce the expected results; the population rose at a slow tempo. By the end of five years’ restriction, however, the White Sea population had improved considerably and began to rise; nonetheless, by 1968 the mother seals only slightly exceeded the 1963 strength (M.Ya. Yakovenko). However, the accumulated reserves should promote the growth of the herd. The seal population is regularly controlled by the scientific institutions of the USSR and especially the Soviet-Norwegian Commission for Seal Research in the Northeastern Subarctic and Arctic Atlantic. The status of the other populations of harp seals, the Jan Mayen and Newfoundland seals, is not so fortunate. The population dynamics are adverse in the case of the Jan Mayen herd, geographically closest to the territorial waters of the USSR and even representing a hunting base for our fleets (see pp. 389-391). The steadily declining numbers of animals caught per ship reflect the significant reduction of reserves. A century before (1866-1870), the average catch per ship was 4,341; at the beginning of this century (1901-1905) 1,400; in 1921-1925, 1,046; and in 1931-1935, fell to 862. Hunting practically ceased in 1941 - 1945. In the postwar years, in spite of the ban and improved fleet, the catch tended to decline.!4 Judging from the results of aerial photographic surveys of the harp seal nurseries in the Newfoundland region, even the Canadian population underwent sharp reduction, with the mother seals in it decreasing by nearly one-half (see p. 391). Field characteristics. The adult animals in the final phase of the hair coat coloration are easily recognized from the intensely dark wing-shaped patterns on the body standing out in contrast to the light-colored back- ground and also from the head which is as dark as the pattern. Among the animals of transitional type coloration, these dark patches are quite distinct along the body flanks. They are diffuse toward the outer fringe although the wing-shaped patches are of smaller dimensions. The imma- ture animals of both sexes and the very young (infantile type of col- oration) gestating or whelping females have fairly distinct contoured, small, dark patches that are sparsely scattered on a gray background. These seals usually live in herds on the coasts and do not emerge onto stationary coastal ice (fast ice). (K.Ch.) 14 The numerical data were borrowed from R.Sh. Khuzin who processed the hunt- ing statistics published by Sivertsen (1941), Iversen (1927*), and the Norwegian Fishing Directorate. 323 429 Economic Importance The harp seal is of primary importance to hunting. Its potential resources оп restoration and subsequent rational utilization would ensure a catch in future (for the population as a whole) of at least 400,000 animals per year for a very long time. Unfortunately, due to mindless plundering of reserves even in the relatively recent past under pressure of international competition and markets, the reserves of this species of seals had been rather depleted by the middle of the present century. This situation was largely due to the long absence of a reliable census. The numerical data given in Table 20 show the volume of hunting in the past and at present. The advance of the harp seal to first place among the seals hunted was promoted not so much by the very high potential of its population increase (the total actual reserves of the ringed seal are slightly more than the present-day strength of the harp seal), as by the favorable conditions for its hunting. These conditions are: the high degree of concentration in localized sections of the range where these seals form massive and fairly dense groups in rookeries on the ice and are quite accessible even to poorly equipped hunters. With the present organization and hunting techniques, however (powerful icebreakers, better hunting ships, trans- port and survey helicopters and planes, radio communications, etc.), the high concentration of these seals makes for highly efficient hunting. The harp seal is caught at present mainly for the fur of the juveniles. An additional raw material is the subcutaneous fat used as medicinal, edible, and commercial fat. The meat portion is used as feed in the farms of fur animals. The harp seal has long been hunted in the White Sea. The origin of such hunting runs into the prehistoric period. There was regular seal hunting in the Neolithic period, mainly (judging from kitchen remnants) of the harp seal, even in the basin of the contemporary Baltic Sea. In the remote past the seals played a very significant role in the life of prehistoric man at many places by providing him not only with hides and fat, but also edible meat. Until recently, the coastal Eskimos and Chukchis depended almost wholly for their existence on the hunting of the walrus and various seals. Even at present for people from the south who settled along the White Sea coasts and took to raising cattle and at places cultivation and fishing, sealing (in which the hunting of the harp seal was of exclusive importance) is not the least of their activities. Right up to the beginning of this century (especially in 1901), earnings from hunting the White Sea harp seal in Mezensk district were put at 27,000 rubles and exceeded 322 430 Table 20. Average number of White Sea harp seal* killed per annum at five-year intervals from 1875 through 1945 (in thousands) Years Killed by Russian Killed by Norwegian Total hunters hunters average ее ВРК pos ee ted res kel tee for the Min. Max. Average Min. Max. Average period 1875 - 1880 19.3 48.3 30.7 5.6 9.1 73 38.1 1881 - 1885 8.9 58.3 26.6 gril 19.8 9.6 36.2 1886 - 1890 12.9 33.0 19.1 15.8 22.7 19.6 34.8 1891 - 1895 13.3 40.7 23.6 24.1 33.0 28.5 S251 1896 - 1900 37.0 74.5 57.1 27.1 38.0 34.7 91.7 1901 - 1905 19.3 64.6 33.9 32.5 79.8 57.9. 92.0 1906 - 1910 19.2 25.6 21.8 42.6 107.0 76.8 98.7 19111915 27.7 50.2 46.3 84.5 118.4 99.2 125.6 1916 - 1920 28.0 49.6 37.8 74.9 154.2 106.6 144.4 1921 - 1925 36.4 124.6 70.4 69.0 343.0 188.8 259.3 1926 -1930 92.5 182.1 140.5 90.7 РЗ 167.2 316.7 1931-1935 102.1 197.8 141.1 47.1 150.5 108.1 245.1 1936 - 1940 10.1 168.9 95.3 34.1 42.7 37.1 124.9 1941-1945 Poot 131.8 55.0 — — = 55.0 *Calculation based on material compiled by R.Sh. Khuzin, who used such primary sources as Iort and Knipovich (1907) and Siverstsen (1941*), whose data included other species as well (their quantum does not exceed 1%); data for the postwar Norwegian statistics (Fish Catch, 1949 to 1959) and Russian statistics. earnings from salmon fishing (23,000 rubles). A higher income came only from forestry (wood processing and sawing) at 100,000 rubles, agriculture 77,000 rubles, cattle breeding 47,000 rubles, and navaga fishing 34,000 rubles. The income was less from deer farming, river and lake fisheries, hunting in forests, and transport. The primitive seal hunting on the ice floes in the White Sea was cum- bersome and risky in the past and was organized by groups of hunters. The people on the coasts and in Mezensk district called such groups “bursas”. For the most part, a group consisted of five to seven hunters in a spacious boat. On the Tersk and Karela coasts, these groups (called “romshas”) comprised only a few people. When the rookeries began forming, the “bursas” set out with their paraphernalia for the sea ice. Dragging heavy boats, experiencing quite some hardships, and exposed to danger, the hunters wandered in search of the rookeries for many days. It was most risky to land on drifting ice floes singly or in twos or even threes (without boats) while chasing the seals sighted from the coasts. It was necessary to hurry back, towing the hides and the blubber, to negotiate the island before the ice broke apart excessively. The first Russian ships began being detailed for seal hunting in the White Sea right at the beginning of this century but group 324 431 hunting continued until the Revolution and after. From the 1920s to the 1930s, most of the hunters from the coastal collective farms engaged icebreakers which, in the 1920s - 1930s as well as in the postwar years, carried out state hunting operations in the White Sea. Apart from the icebreakers, special hunting ships were recruited. On the whole, hunting, regularly assisted by reconnaisance planes, was more intense. The White Sea harp seal was killed in the largest numbers (exceeding 300,000 annually) for five years from 1924 through 1928. In the next five- year period (from 1929 through 1933), the average annual kill dropped to 250,000, and still later from 1934 through 1938, fell further to 188,800 on average. There is no doubt that hunting in the 1920s was extremely intense. The average annual kill in the postwar years steadily fell as follows: О. 166,300 Spy О оное 151,700 OSD Об: а, 115,000 О A), Su res aa) ot 81,900 Norwegian hunting ships also cruised along with our hunting opera- tions in the White Sea region and in the adjoining southeastern sections of the Barents Sea. Before the end of the 1930s, the Norwegian hunters were given a concessional right to hunt for the harp seal in the north- ern part of the White Sea (north of the line joining Capes Orlov and Konushin). At the end of the Great Patriotic War, the agreement was not renewed and at present the Norwegian hunters hunt outside the White Sea limits and in the adjoining areas in a few dozen small hunting boats. Hunting is regulated by agreements. From the second half of the 1960s, the organization of our hunting activity in the White Sea underwent radical change. In the five years ending with 1969, state hunting ceased because of the depleted reserves. Neither icebreakers nor special hunting ships carried out hunting in the White Sea. An exception was made only for the local collective farms, which were given the right to hunt 20,000 pups a year. The collectives rented helicopters to reach the hunters on the ice and transport the killed animals to the coastal base. No more than 10 days were spent in the entire hunting operations under favorable conditions. Until recently, well-fed normally molting young animals were brought onto the coasts where they were allowed to complete molt in special enclosures on land. The Norwegian hunters in the immediate proximity of the White Sea inlet area caught no more than 14,500 animals per annum in the 1960s (Table 21), mainly the older juveniles and adults. It was thought prudent to maintain the hunting level as at the end of the 1960s in 325 432 the next decade to promote a more rapid population recovery. After a short period of relative restriction, the population had not reverted to the level of the 1920s and 1930s (the year 1925 was a record for the White Sea region; Soviet hunters caught 124,600 animals and the Norwegians 343,000 animals, 1.е., slightly less than half a million seals). The ban imposed in 1963 on killing adult females in the nurseries had an extremely salutary effect in restoring the White Sea population. The technique of hunting in the nurseries was extremely simple in the past. The hapless white pups could not escape from the hunter and were killed directly using clubs. Since suckling mothers usually do not leave their pups, “hunting” them also posed no problem. Nevertheless, rifles were used to kill them. In the past, hunting in the molting rookeries was more complicated. Hunters with harpoons [rifles] approached quietly in white masks and aimed from behind cover. The shots frightened the sleeping animals and soon only dead animals remained on the ice floe. In some cases, when the ice floes became compact (usually in high tide), open water pools disappeared and the ice lumps filling them froze and the animals were thus cut off from the water. Hunters could then kill them even with clubs. Usually, however, even in compact frozen areas, the animals could ultimately get into the water by pushing aside the ice lump or pressing it with their bodies. Table 21. Harp seals (White Sea population) killed in the postwar years (in thousands) (after R.Sh. Khuzin) Year Killed by Killed by Total Soviet hunters Norwegian hunters killed 1946 79.1 8.5 87.6 1947 161.1 6.6 167.7 1948 146.3 8.9 155.2 1949 183.4 25.1 208.5 1950 194.7 LT, 212.4 OSH 192:2 33.8 226.0 1952 131.4 19.1 150.5 1953 88.3 12.4 100.7 1954 152.8 11.6 164.4 1955 97.8 191 116.9 1956 68.0 25:1 93.1 1957 107.9 22,3 130.2 1958 119.6 15.1 134.7 1959 101.4 8.5 109.9 1960 95.9 10.7 106.6 1961 93.8 1:2 105.0 1962 106.9 8.3 15.2. 1963 69.5 13.3 82.8 1964 62.7 14.6 77.3 1965 20.1 6.4 26.5 1966 20.0 12.2 32.2 326 433 The first of the denser rookeries of males with a “winged” pattern and disposed in rows along the edges of open water pools and fissures (Fig. 174) are the least accessible to hunters. It is difficult to approach the animals within the required distance as they abandon the ice floe at the first shot and dive pell-mell into the water. The killed animals are processed very quickly: first a cut is made along the abdomen from head to tail, next circular incisions made through the skin with the fat layer around the base of the flippers, and then the subcutaneous fat separated from the muscles. The skins with the fat upward are spread out on the ice and stacked for loading in a vessel that comes later. Nowadays, the extensive used of airplanes and killing pups exclusively has made the job quite simple and fast. At the coastal base the fat is separated from the skin, the skins degreased to the max- imum possible extent, and salted. In the natural or even dyed form, the white embryonic pelage and now, more so, the skins of the molted gray pups are in great demand in the internal and well as international markets. Drifting ice floes in the region of Jan Mayen Island represent another region of hunting. Exploitation of this international hunting zone began over 200 years ago. The maximum intensity of hunting in Jan Mayen was recorded in the 1870s (200,000 seals were killed there in 1874). The decline in killing began subsequently, touching the lowest level at the end of the first 10 years of this century. World War I interrupted the Fig. 174. Early spring rookery of adult males. White Sea, April 18, 1966 (photo- graph by Уи.Г. Nazarenko). 326 327 326 434 hunting activity and this helped the herd to restore itself to some extent. After the war, hunting resumed and reached a fairly high level. World War II again interrupted hunting activity but it was rapidly resumed thereafter and, in the first half of the 1950s, exceeded the prewar level. On average, the kill in the first five years of the 1950s was 39,300 seals with a very large number of modernized vessels operating. However, the success was short-lived and unreliable. The pressure on the animal population was excessive and it rapidly declined, an index of this decline was the reduction in the number of animals killed per ship (see Table 22). The lowest annual yield for the entire hunting duration in this century occurred in the first half of the 1960s (average kill per ship fell to 467 seals in one expedition although the vessels had been equipped with new diesel motors) (R.Sh. Khuzin). Soviet ships began hunting in the ice floes of Jan Mayen Island region in 1955 and continued hunting there until 1965 (see Table 23). Hunting was discontinued as it was unprofitable. The commercial exploitation of the Newfoundland populations by the local people, in which the USSR became somewhat interested in the 1960s, began almost concurrently with hunting of the White Sea resources. Hunting of harp seals using ships had begun there at the end of the eighteenth century and reached an immense scale of 500,000 a year in the 1820s. Hunting continued at this level for forty years (Allen, 1880). Table 22. Norwegian hunting of the harp seal in the Jan Mayen Island region from the 18805 (after R.Sh. Khuzin) Years Average annual Average number of Average kill per kill, thousands expeditions per ship expedition, ship per annum numbers 1881 - 1885 82.8 17.5 5,241 1886 - 1890 33.3 22.8 1,469 1891 - 1895 45.8 19.6 2,338 1896 - 1900 24.3 12.4 1,960 1901 - 1905 15.1 10.8 1,400 1906 - 1910 14.0 21.6 648 1911-1915 17.4 29:2 596 1916-1920 32.3 59.7 542 1921 - 1925 19.5 18.6 1,046 1926 - 1930 32.6 33.5 969 1931 - 1935 32.3 37.4 862 1936 - 1940 36.7 45.6 803 1941 -1945 — = — 1946 - 1950 36.1 37.0 976 1951-1955 39.2 55.6 859 1956-1960 25.0 39.8 585 1961 - 1965 18.6 39.8 467 435 Table 23. Jan Mayen harp seals killed by Soviet hunting ships in the 1960s (after R.Sh. Khuzin) Year Number of Total killed by all ships, hunting ships number Total Pups 1960 8 3,356 420 1961 8 6,043 755 1962 8 2,423 303 1963 i, LOG, 222 1964 8 4,483 560 1965 8 6,273 896 One can well conjecture the original population level of the local harp seals to be able to sustain such a prolonged and intense rate of hunting. By the end of the nineteenth century, the number of animals killed had fallen to half in spite of maintaining the same intensity of hunting; in the first few decades of this century, the number decreased even further, to 160,000 a year. World War II served as a fortuitous interlude that extended for sometime after the war. The continuous interruption of hunting using ships undoubtedly promoted a partial recovery of the animal population. But hunting resumed in the 1950s and exceeded the prewar level, with the annual kill reaching 262,000 (Fisher and Sergeant, 1960*). At the present level of the animal population, given such large kills the animal population can hardly restore itself to the original level. At the beginning of the 1960s, hunting was carried out in the New- foundland region twice on a trial basis by Soviet hunters. As it proved unprofitable and Canada extended its coastal waters to 12 miles, the Soviet Union ceased hunting in the Newfoundland region. At the end of the last century and in the first thirty years of this century, this seal was also caught in nets. This was mainly practiced by the locals off Murman but here and there the White Sea harp seal was caught even in the White Sea before total icing set it. Catching in nets was quite profitable off Murman. For example, at Kharlovka about 1,000 seals were thus caught from the end of winter to early spring in 1898 - 1899.1 This hunting practice flourished even in the 1920s (Skvortsov, 1927). Nets were no longer used in subsequent years off Murman mainly because the seals changed their approach to the Murman coast. Sighting the harp seal on the Murman coasts, people unjustifiably assumed its adverse effect on the catch of cod there. Even very old reports 15 See: Proceedings of the Murman Scientific-Hunting Expedition, St. Petersburg, 1912. 328 436 contain frequent references to the fatal consequences of the presence of seals on the Murman fishing industry. Thus, in 1801, the seals report- edly “chased away the entire fish from the Murman coast” right up to Rybach’ii Peninsula. In 1803, the “catch was good from the spring but kozha!!© arrived insummer and chased away the entire cod”. In 1807, “tight from the spring, kozha spread all along the coast like tree stumps in a forest, as a result of which the fishermen could not catch even a single fish”. Similar complaints were registered in subsequent years (Reineke, 1830*). In fact, the White Sea harp seal continued to be accused for quite some years, often with even greater exaggeration. The accusations that the harp seal consumed large quantities of com- mercial fish, especially cod, or chased it away were never confirmed. On the contrary, the arrival of cod was usually not accompanied by the arrival of seals; instead the absence of cod coincided with the arrival of the White Sea harp seal. Thus the cod per se holds no attraction for the seal (N. Smirnov, 1924); this is explained by hydrological conditions. The harp seal is a distinct element of the arctic fauna; it is sensitive to the thermal regime of the water (even early in this century it served as an indicator of the cooling of the water or of the invasions of arctic waters (Linko, 1912)). Thus, instances of the temporary disappearance of the relatively thermophilic cod on the Murman coast and the simultaneous appearance there of the White Sea harp seal were actually impelled by cooling or ingress of arctic waters. The harp seal of the White Sea herd is one of the most harmless species of seals in relation to the fishing industry, coming in third place after the bearded seal and the walrus among the Pinnipedia of the North Atlantic. (K.Ch.) Subgenus of Ribbon Seals Subgenus Histriophoca RIBBON (BANDED) SEAL!” Phoca (Histriophoca) fasciata Zimmermann, 1783 1783. Phoca fasciata. Zimmermann. Geographische gesch. des Men- schen u. der algemein. verbeit vierfussigen Thiere, 3, p. 277. Kuril Islands. 116 «“Kozha” is a local term used by the coastal people for the harp seal. 17 Krylatka—adult animal; belek—newborn pup; and serka—under-yearling after casting the juvenile hair coat. 437 1831. Phoca equestris. Pallas. Zoogr. Rosso-Asiat., I, р. Ш. Kuril Islands, Sea of Okhotsk (“Rarissime in mari Ochotensi, frequentius circa Curilorum insulas. . .”). (V.H.) Diagnosis Four light-colored transverse bands run along the generally dark back- ground of the trunk (Fig. 175). The labial whiskers number 41-42 and orbital whiskers 6-8. The whiskers at the tip of the snout have a wavy surface (Fig. 176). The first digit on the fore flippers is longer than the rest. The paired air sac joined with the trachea is well developed among males (Fig. 177). (V.A.) Description These are seals of moderate dimensions [up to nearly 2 m in length]; the males are usually larger than the females. Body long, streamlined. The color of the adults is typical: four light- colored bands run along the dark background (black or brown among males; brown or brownish-gray among females). One band encircles the neck like a collar, covering also the rear portion of the head, and another ==, 92 Ра > Sa к. < 2 — = о ems A = ее —= a) О i 329 Fig. 175. Color of the adult ribbon seal (banded seal), Phoca (Histriophoca) fasciata (figure by N.N. Kondakov). 329 Fig. 176. Whiskers of the ribbon seal (banded seal), Phoca (Histriophoca) fasciata (figure by N.N. Kondakov). 438 329 Fig. 177. Air sac of the ribbon seal, Phoca (Histriophoca) fasciata (figure by N.N. Kondakov). A—general view of air sac filled with air (seen from the right); B—sketch of respiratory tract in the ribbon seal. 1—cervical section of air sac; 2—trunk section; 3—trachea; 4—enlarged section of lower trachea; 5—lungs; 6—thoracic cage (after M.M. Sleptsov). encircles the body at the base of the hind flippers; two symmetrical bands mark the outline of the base of the fore flippers. From a distance, these 330 flippers look like the attached wings of a bird. The width of each band is 5.5-15 cm and its color varies from pure white to yellowish. Fine mottling is sometimes seen on the light-colored bands of young animals. There is no seasonal variation in coloration. (Age-related color variations are described under “Growth, Development, and Molt”.) The skull is shortened and broad, the cranial portion and zygoma broad, the facial portion short and narrow (Fig. 178). The nasals are short, on average 36.6 mm. The bony nasal septum does not reach the posterior margin of the bony palate. The length of the palatine bones is usually less than their maximum width. The tympanic bullae are large (about 33.5% of the condylobasal length), their width considerably less than their length, and set sharply inclined toward the longitudinal axis of the skull. The alveolar margin of the upper jaw is curved; the lower jaw is curved with a downward convexity corresponding to the curvature of the upper tooth row (Fig. 179). The teeth are small and number 32-36. The canines are small and obtuse, the incisors weak, slightly curved, almost vertical. The molars and premolars are small, set wide apart, their roots tending to fuse. The 330 331 439 Ри iS) <0 cree \ = Si a My у md у % Ip AD \ : \ LX i WR Sy чи y PS Е y a 7 wo Г) 5 . я in, \F2 ( ib inane SY, : oct iat Ve Reo Fig. 178. Skull of the ribbon seal (banded seal) Phoca (Histriophoca) fasciata (figure by N.N. Kondakov). only accessory cusp, located posterior to the base of the last tooth, is most often in-the form of a faintly recognizable denticle. The average body length varies from 155 to 165 cm depending on the age composition of the sample). The maximum length of the male (Bering Sea) is 192 cm, of the female 198 cm (males, however, are gener- ally larger than females). The length of the os penis may reach 150 mm. The condylobasal length of the skull in males averages 191.7 mm, in females 190.7 mm; mastoid width in males 125.1 mm, in females 121.4 mm; length of the upper tooth row in males 57.5 mm, in females 59.0 mm (Ogney, 1935; $. Naumov and М. Smirnov, 1936; Chapskii, 1955, 1963; Shustov and Yablokov, 1967). The maximum weight of these seals is 150 kg, the average being 55 kg. The average weight of the blubber (fat with hide) is 20 kg and the subcutaneous fat layer is 2- 4.5 cm thick. The weight of the os penis is about 20 g. The weight of the internal organs of mature animals is as follows (g): heart 499-513, lungs 978 - 1,030, liver 1,378 - 1,403, stomach 401-441, intestines 1,351 - 1,433, and kidneys 123-144 (A.S. Sokolov et al., 1969). The mode of life of the ribbon seal outside the icy season is not known and hence there is no information on the seasonal changes in weight. (V.A.) 330 332 440 Fig. 179. Bony palate and teeth of the ribbon seal, Phoca (Histriophoca) fasciata (figure by N.N. Kondakov). Taxonomy The ribbon seal is distinguished in the family by its specialized pelagic mode of life, which has led to some characteristic structural features. Some authors (Ognev, 1935; Simpson, 1945; Chapskii, 1963) regard the ribbon seal as a monotypical genus, while others (S. Naumov and N. Smirnov, 1936) combine it in the same genus as the harp seal, with which the ribbon seal shares several common morphological and ecological features and evidently has a common ancestor, although the genesis of the latter is not clearly known. (V.A.) Geographic Distribution Seas of the northern Pacific Ocean—Sea of Japan, Sea of Okhotsk, Bering Sea and Chukchi Sea. Endemic to this section of the world oceans. Geographic Range in the USSR Divided into two isolated portions, 1.е., Sea of Okhotsk and Bering Sea (Fig. 180). In the Sea of Okhotsk this seal is encountered on the spring ice floes from La Perouse Strait to the Shantarsk archipelago and She- likhov Bay. the southern boundary runs along the northern part of Tatar Strait (Dorofeev, 1936) and into the coastal waters of the northeastern 331 441 250 500 750 1000 km — Fig. 180. Distribution of the ribbon seal (banded seal), Phoca (Histriophoca) fasciata in the USSR (V.A. Arsen’ev). coast of Hokkaido Island (Nishiwaki and Nagasaki, 1960; Nishiwaki, 1966). Distribution in the Bering Sea covers the coastal waters of Koryak Land from Olyutor Gulf in the north, Gulf of Anadyr, Bering Strait, and the Chukchi Sea to Kolyuchin Bay in the west (Shustov, 1965). Geographic Range outside the USSR Eastern part of the Bering Sea from Bristol Bay to Bering Strait and waters of the Chukchi Sea to Cape Barrow in the east (Scheffer, 1958; Shustov, 1965). 332 442 The southern boundary in the Bering Sea runs from Olyutor Gulf northeast roughly up to 60°N lat., from there eastward and then south- east up to Bristol Bay (Fig. 181). The southern limit does not cross the islands of the Aleutian range (Allen, 1880). The northern boundary runs at places between the Chukchi Peninsula, Alaska, and Wrangel Island from Kolyuchin Bay to Cape Barrow. (V.A.) Geographic Variation The southwestern boundary of distribution of the ribbon seal in the Bering Sea lies more than a thousand miles away from the northeastern boundary in the Sea of Okhotsk. The existence of two local populations is thus highly probable. All the same, geographic variation has not been established. (V.A.) Biology The mode of life of the ribbon seal is known for a comparatively small duration of the year, 1.е., at the time of spring-summer rookeries on the Fig. 181. Species distribution of the ribbon seal (banded seal), Phoca (Histriophoca) fasciata. Dots represent sites of records of the ribbon seal (V.A. Arsen’ev). 333 443 ice floes. From this viewpoint, И is one of the poorly studied species of Pinnipedia of the Northern hemisphere. Population. An idea of the population of the ribbon seal in the Sea of Okhotsk was provided by the data of the annual catch, aerial surveys, and observations from ships used for hunting. The total population was thus put at a few tens of thousands. No special census was undertaken. The abundant population of the Bering and Chukchi seas began attracting hunters only in 1961. Calculations based on the areas of ice floes, density of disposition of seals on various types of ice floes, and aerial and visual observations put the total strength of this population at 80,000 to 90,000 (Shustov, 1969). Habitat. These seals generally select firm, clean, white hummocky ice blocks invariably with an even surface for their spring-summer rookeries. Instances of finding these seals on dirty ice blocks are very rare and are possible at the end of the icy period when the area of the ice floes has greatly diminished. The height of the ice floes is not of much consequence as the seal can jump onto an ice block 1 m or more in height. The main rookeries are disposed on drifting ice floes far away from the coasts but usually overlying depths not exceeding 200 т. The seals are also sighted in coastal waters but only in the case of early thawing of ice floes. Outside the icy period, the seals evidently lead a pelagic mode of life since they never emerge onto the coasts (barring extremely rare cases). The present sites of distribution of the seals have not been established. Food. Data on the food of the ribbon seal have been collected exclu- sively for the period of their residence in the rookeries on the ice floes with no information at all available outside this period. Most of the dissected stomachs in all the regions of study were found to be empty. Thus, 91.6% of 443 stomachs dissected in the Sea of Okhotsk were empty (Pikharev, 1941); in another case, 82% of 398 stomachs were devoid of food (Arsen’ev, 1941); in the Bering Sea, 97.4% of 1,175 stomachs dis- sected were empty (Shustov, 1965). Based on these data, it was earlier thought that the ribbon seal did not feed in the icy period but thorough investigations established that the intestine of many seals contained food remnants while the rectum was packed with fecal matter. Very rapid food digestion is evidently characteristic of this seal. In addition to the items listed in Table 24, the stomach of seals in the Sea of Okhotsk at one time contained the bone remains of navaga (Freiman, 1936b) and mysids (Nikulin, 1937b). In the Sea of Okhotsk as well as in the Bering Sea, some stomachs revealed the feathers of sea birds, sand, and stones. The geographic variation of the food of ribbon seals was quite signif- icant but data for the Sea of Okhotsk (V.A. Arsen’ev, 1941) are clearly 334 44+ inadequate. The age-related food variations are: young animals (primar- ily the under-yearlings) feed mainly on pelagic invertebrates (amphipods, mysids and isopods) while adults feed on benthic and pelagic forms in spite of the fact that the depths in the regions of their habitat reach 50 to 100 m or more. Home range. The ribbon seal does not form large rookeries on ice floes but lives singly or sometimes in twos or threes. There are no distinct male or female rookeries. These seals are seen only on ice floes with a density of predominantly four to six points. Hideouts and shelters. These seals do not make air holes or snow shelters. Table 24. Food items of ribbon seals (Shustov, 1965a) Food item Number of cases Percentage of number of stomachs with food remains Seaof Bering Seaof Bering Okhotsk Sea Okhotsk Sea CRUSTACEANS Crangon dalli 1 — 2.0 — Nectocrangon lar = 3 = 9.4 Pandolopsis sp. = 6 = 18.7 Pandalus borealis — 8 = 25.0 Pandalus goniurus 5 1 9.8 3:1 Eualus ватагай т 6 = 18.7 Spirintocaris murdochi os 1 = 3.1 Lebbeus sp. = 3 — 9.4 Temisto sp. (2 species) = 6 = 18.7 Stilomysis grandis = 2 = 6.2 Amphipods =a 4 — 12.5 Cephalopod mollusks 24 5 47.0 15.6 FISHES Polar cod, Boreogadus saida = 4 — — Atlantic herring, Clupea harengus = 2 — 6.2 Pacific navaga, Eleginus gracilis — 2 — 6.2 European smelt, Osmerus eperlanus — 1 — 3.1 Stout eelblenny, Lumpenus medius = 4 = 12.5 Alaska pollock, Theragra chalcogramma 47 = 92.1 — Capelin, Mallotus villosus 1 2 2.0 6.2 Common sand eel, Ammodytes hexapterus = 1 = 3:1 Smooth lumpsucker, Aptocyclus ventricosus 1 = 2.0 — Pacific cod, Gadus morrhua macrocephalus 4 — 7.8 = 334 335 445 Daily activity and behavior. The number of stomachs with food rem- nants was 20% more among seals caught during the night, suggesting a more active feeding at night (Shustov, 1965). No other data are available on daily activity. In cloudy weather, many seals are seen resting on ice floes in the morning and evening hours while most of them remain in water at noon. In rainy weather most seals prefer to be in water and rarely venture onto the ice. On quiet, sunny days the seals almost do not enter the water in the day but gather there in the evening hours and at night. A large number of seals remain in water at night, roughly between 8:00 p.m. and 4:00 a.m., regardless of weather conditions (Pikharev, 1941; Shustov, 1965b). The ribbon seal very easily negotiates ice floes with long leaps, without touching the edge of the floe. It lies quite close to the edge, quite often on a level surface at the foot of a hummock with its head invariably toward the water. It is extremely sensitive and approaching it without detection is very difficult. If, however, a boat with the motor switched off runs straight toward the animal and the people in the boat remain quiet, the seal carefully surveys the boat and permits its approach within rifle shot without attempting to escape into the water. Probably its auditory faculty is poorly developed compared to vision. An injured seal attempting to reach the water quickly, will invariably stop and howl sharply, which the hunter promptly takes advantage of. The animal moves rapidly on the ice floe, contorting its body like a snake, dives almost noiselessly, without a splash, moves far away in the water, and never surfaces immediately. A badly wounded animal attempts to return to a block of ice while a slightly injured one can fiercely attack the hunter (Pikharev, 1939, 1941). Sometimes, for no reason at all, the seal begins to move rapidly on the ice floe from one side to the other, rolling and turn- ing from one side to the other, then rushes suddenly into the sea, only to return immediately to the ice floe with a leap. The motive for these actions is not clearly understood; they resemble playful acts although done singly, without a companion. Seasonal migrations and transgressions. Information on migrations is practically not available. In the rookeries on the ice floes, the seals are mainly passive and the movements they do make can hardly be regarded as migratory. On the spring ice floes in the Sea of Okhotsk, the largest concentrations are noticed south of Tauisk Bay, in the region of lony Island, north and east of Sakhalin. At the end of the icy season, herds of these seals migrate into the region of Shantarsk Islands and into the Gulf of Sakhalin. They are distributed on the ice floes, in patches. In relatively nearby regions on the ice floes, appearing almost identical externally, some will be occupied by the animals and others vacant, the 446 occupied and vacant floes alternating. Possibly, this is associated with the depths of the region and hence the chances of finding food (Shustov, 1965b). With the thawing of ice floes in the Sea of Okhotsk and the Bering Sea, the ribbon seal disappears. Its summer residence has not been established. Only some stray observations have been reported on the migrations of some seals through the Bering Strait in spring to the north and in autumn to the south (Shustov, 1965b; Tikhomirov, 1966). Some distant transgressions of the ribbon seal are known. Early September, 1927, a seal was killed in Vladimir Bay in the Sea of Japan (about 44°N lat.) (Pikharev, 1941). In another case, an adult ribbon seal was noticed on June 17, 1944, in Tsushima Strait (35°10’N lat. and 130°34’ E long.) 45 miles from the coast (Vedenskii, 1950). Transgres- sion of the seal was noticed into the western part of the East Siberian Sea (Ognev, 1935) and toward Wrangel Island. A male ribbon seal, 131 cm long, was caught on November 16, 1962, close to Morro Bay in Cali- fornia. [ts body was devoid of a hair coat except for the vertical surface of the fore and hind flippers, head, and upper portion of the neck. The animal was kept in an aquarium where it died a month later (Aryan [Roset], 1964). 335 Fig. 182. Female ribbon seal with a newborn. Bering Sea (photograph by А.Р. Shustov). 336 447 Reproduction. The period of whelping extends from March end to early May. In 1962, mating was noticed for the first time in the Bering Sea on March 29 while newborns were encountered on Мау 2-3, 1963. Mating takes place soon after parturition and thus gestation extends for about a year. Most of the females are capable of producing an offspring every year. Females attain sexual maturity at two to three years of age and males a year later at three to four years. Females older than five years are the most productive. The upper age limit for productivity has not been established nor has any climacteric period been detected among females. The intensity of reproduction is quite high and the number of gestating females in some years varies from 30 to 60% or more of the total number of females in the population (Shustov, 1965b). Growth, development, and molt. The ribbon seal is characterized by a latent period of embryonic growth [delayed implantation], roughly cal- culated as 2.5 months. The newborn averages 85 cm in length, i.e., about one-half the length of the mother. It reaches this length roughly after nine months of uterine development. The average weight of the pups is 8.6 kg. The pups are covered with a long, soft, silvery-white embryonic coat with smoky spots; this coat is sported for about two weeks. In this period the pups feed on the mother’s milk. At 25 to 30 days of age, the molted pups (normally molting pups and gray pups) measure an average length of 112 cm and weigh 28.3 kg. The hair coat is short and coarse, slaty on the back, gray on the flanks, and light gray on the belly. After the first molt a dark band with sharp outlines appears on the generally monochromatic dorsal background. At two years of age the white bands characteristic of adults are seen for the first time (Fig. 183). At this age the female acquires the adult coloration while the male of two (or some- times three) years of age most often is very dull colored like the adult female. Males acquire the adult vividness at 3-4 years of age. The intense growth of seals in the first few years slows down after maturity. Growth cessation among females occurs evidently at seven to eight years of age and among males at 7-9 years. By this time the seals are fully grown. Females lag behind the males in growth though not very significantly (Table 25). The longevity of the ribbon seal has not been established. The oldest male studied was 27 years while two females were 26 years old. These animals did not give the impression of senility (Shustov, 1965b; Shustov and Yablokov, 1967; Tikhomirov, 1968). Shedding of the embryonic hair coat constitutes the first molt, the signs of which are noticed 7-10 days after birth. The hairs are initially shed from the head and the hind flippers and later from the other sections 336 337 Fig. 183. Young ribbon seal. Bering Sea (photograph by А.Р. Shustov). Table 25. Variation in body length of seals with age (measured along the dorsal surface) (Tikhomirov, 1968) Age, years Males No. of animals measured 1 16 2 20 3 29 4 33 5) 1 6 20 7. 17 8 19 9 16 10 21 ее 13 12 14 13-14 22 15-17 26 18 апа абоуе 15 Average length (cm) 132.0 145.0 155.0 156.6 165.0 158.9 160.1 161.0 168.1 163.8 165.0 167.7 165.2 168.7 163.1 Females No. of animals measured Average length (cm) 129.0 145.0 152.0 153.9 160.2 161.5 168.0 169.0 163.9 163.0 163.9 168.8 168.1 of the body. The juvenile coat is preserved for quite sometime in the armpits of the fore flippers. Molting ceases roughly at two weeks of age. The molting of adult seals extends for quite a long period. The sec- ond half of April can be regarded as the beginning of the molting period but the cessation of this period has not been established. A ribbon seal caught in the Sea of Okhotsk on May 16, 1939, was at the peak of 338 449 molt while another one caught on May 20, 1938, was found to be fully molted. In mid-July, molting animals were seen among those that had already completed molt. The duration of molting can be roughly put at three months. During molt, not only a change of the hair coat occurs, but often a simultaneous and intense peeling of the epidermal layer (Pikharev, 1939, 1941). Enemies, diseases, parasites, mortality, and competitors. The ribbon seal has practically no enemies in the icy period. Only on occasion does it fall prey to the killer whale, polar bear, or even a large Greenland shark. No information is available on the subject outside this period although there is undoubtedly greater possibility of death being inflicted by the killer whale or shark. Diseases suffered by ribbon seals have not been studied but sometimes animals with skin diseases are encountered. Such animals are partly or even wholly devoid of hair coat, their epidermis is peeled, and bleeding cracks are seen in the affected sections of the skin (a seal in this state was caught in California). The mobility of such animals is greatly reduced and they do not fight man. A blood analysis of three sick seals gave an erythrocyte sedimentation rate of 18 to 25 mm/hr versus 3-4 mm/hr for healthy animals. Such seals, though few, are nevertheless seen every year. The following ten species of helminths have been detected among the ribbon seals of the Bering Sea. Cestodes Diphyllobothrium sp. and Diplo- gonoporus sp. have been found in the intestine. Nematodes of Anisakidae gen. sp. have been found in the intestine and stomach; Contracaecum osculatum Rud., detected in many species of Pinnipedia in the Northern and Southern hemispheres, localizes in the intestine (not detected among ribbon seals of the Sea of Okhotsk). Phocascaris phocae Host, known only among harp seals as well as ribbon seals, parasitizes predominantly the stomach. Terranova decipiens Krabbe was detected only once in the stomach of ribbon seals; it is known among walruses, sea lions, fur seals, elephant seals, and many species of true seals. Another species of this genus, Ter- ranova azarasi Yamaguti and Arima, was found in the intestine of ribbon seals from the Sea of Okhotsk but not the seals of the Bering Sea. From among the acanthocephalans, Bolbosoma nipponicum Yamaguti was also found among many marine mammals of the Pacific Ocean basin. It local- izes in the intestine, as does Corynosoma strumosum Rudolphi, known among a large number of species of Pinnipedia and also among some land mammals and birds. Corynosoma semerme Forssel, widely distributed among Pinnipedia and cetaceans of the Northern hemisphere, is also found among ribbon seals (Delyamure, 1955; Shustov, 1965c). 450 Natural mortality of this seal has not been studied. As in other species, mortality is maximum in the younger age groups. Death caused - by enemies is evidently insignificant but parasites can cause death, albeit to a small extent. Maximum mortality arises from unfavorable ice con- ditions and other abiotic factors. The scattered disposition on drifting ice floes suggests that there is по competition with other species for a place on them. Only at the end of the icy period, in the years of early thawing, when many seals of four species gather on the rapidly diminishing ice massifs, is competition possible between the bearded seal, common seal, ringed seal, and ribbon seal. There is hardly any competition in relation to food since the food zones of all these species are distinctly “delineated”. Population dynamics. The natural population dynamics of this seal have not been studied but its fluctuations cannot be significant. In recent decades this situation has undergone significant changes as a result ot hunting. Before hunting with the help of ships was organized (1932), the ribbon seal was caught by the local people in very limited numbers since it essentially stayed far away from the coasts. This level of hunting had hardly any impact on the seal reserves and the population maintained a natural equilibrium. In the first two decades, hunting was carried out only in the Sea of Okhotsk where the comparatively low level of killing did not alter the natural state of the population. As the hunting fleet enlarged in the early 1950s, and the catch of all species of seals increased, some 20,000 ribbon seals were caught annually. An analysis of the age composition of the seals caught revealed a gradual “rejuvenation” of the population but pointed to a disturbance of the natural ratio of the age groups and a reduction in the reproductive capacity of the popula- tion. The results of these investigations formed the basis for suggestions regarding the need to restrict the killing of the ribbon seal in the Sea of Okhotsk. Hunting of the Bering Sea population commenced only in 1961; until then the population had remained undisturbed. An analysis of the hunting activity of ships and the age composition of the animals caught in 1961-1963 showed that the ribbon seals killed per day decreased con- siderably irrespective of hunting conditions. The specific proportion of the larga, which is the most difficult seal to kill, increased in the catch of the ships; larga becomes the target only when seals of other species are not available. The age composition of the catch also changed. In 1961, the average age of the ribbon seals caught was 9.8 years; it fell to 6.9 years in 1962, and further dropped to 4.9 years in 1963. This points to a reduction of the total population and its reproductive capacity since females older than five years are the most productive (Shustov, 1965). 339 339 451 With expanded hunting in subsequent years for newborn pups because of their valuable fur, the age composition of the animals killed somewhat balanced but the state of the population still causes anxiety. Field characteristics. Characteristic coloration is an unfailing feature for recognizing the ribbon seal (Fig. 184). The seals are found only on white, clean, and stable ice floes drifting far away from the coasts; they do not form groups and are usually seen singly or in twos and threes. In the summer months these seals adopt a pelagic mode of life; sites of summer gatherings have not been established. (V.A.) Economic Importance The ribbon seal is of little importance to the local people who hunt in the coastal zone where its population is very small. The proportion of the ribbon seal in the ship catches in the Far East is not less than 30% of the total catch (over 80% in the Bering Sea) and thus its importance here is quite substantial. It should be borne in mind that the ringed seal, which represents the largest number of seals caught, is a much smaller animal and provides a smaller quantity of useful products compared to the ribbon seal. Fig. 184. Adult male ribbon seal on an ice floe. Bering Sea (photograph by A.P. Shustov). 340 452 Seals of all the species are killed in the Far East using firearms from motor boats dropped in the water from a schooner in the hunting region. Seals resting on ice floes represent the predominant target since killing the animals in the water inevitably results in great losses as most of these killed in the icy period drown. The products of hunting are the hide, fat recovered from the subcu- taneous blubber, and the carcass. The blubber is removed at the site of the kill (on the ice floe). Its further processing is carried out either on some Suitably equipped ship or at the coastal base where the ships deliver the raw products for processing. Salt is used as a temporary preservative of raw skins on the ships. The hides of adult seals are mostly used as raw leather while those of white pups and the better quality skins of adult animals serve as valuable raw material for the fur industry. The fat recovered is used for commercial purposes in tanning, metal, and other industries. The carcass is a valuable product for the animal breed- ing farms extensively organized in the Far East; the meat and bones are used as feed for fur animals. The local population fully utilizes the animals killed. Both the populations of the ribbon seal are exposed to heavy hunting (see “Population Dynamics’’). The results of investigations indicate the need for an extremely careful approach to utilizing these populations. Hunting norms should be fixed annually for each population on a ratio- nal basis depending on its condition and strict adherence to the norms enforced. (V.A.) Genus of Gray Seals Genus Halichoerus Nilsson, 1820 1820. Halichoerus. Nilsson. Skand. Fauna. Dagg. Djur., I, p. 376. Hali- choerus griseus Nilsson = Phoca grypus Fabricius. These are relatively large seals, the adults measuring about 2 m or more (Lc). The facial portion of the head is noticeably elongated (more than twice the distance from the eyes to the ear opening) and elevated. Its height in front is almost the same as at any other point while the upper contour of the profile draws an even or slightly convex line at the bridge of the nose. The whiskers are flattened, with wavy edges. The first and the second digits on the fore flippers are longer than the others. The profile of the skull is seen as an almost perfect straight upper contour from the occipital crests to the anterior margin of the nasal bones; the height of the facial portion on the anterior margin of the nasal bones is no less than the height of the cranium. 341 453 The anterior nasal opening is extremely elongated and opens upward; the maximum width occurs in its posterior one-fourth. The nasal bones are relatively short (not more than one-fifth the condylobasal length) and broad; their total width at the anterior margin is about one-half or more their length. On the posterior (orbital) side of the zygomatic processes of the maxillary bones lies a horizontal carinate crest which conceals the suborbital aperture when viewed from above. The length of the zygomatic bones without process is more than double their smallest width; the lower posterior process of these bones is considerably longer than the upper. The features of the tympanic bullae, viewed from below, appear as a triangle with rounded apices and resemble those in seals of the subgenus Phoca s. str. The bony lobe of the external auditory meatus is simple in design, flattened and broad. The premolars and molars, except for the first molar, are massive; the second and third premolars and molars of the upper jaw and also the first three premolars of the lower jaw have conical crowns and lack accessory Cusps; accessory cusps are seen on the fourth premolar of the upper and lower jaws and on the lower molars; highly reduced acces- ОГУ cusps are sometimes seen even on the initial premolars. A second molar is sometimes seen in the upper jaw. At least the three initial premolars of the upper and lower jaws among adults have a single root. True molars, however, invariably have two isolated roots set in separate alveoli. Incisors have laterally flattened roots. The dental formula is: SA ЩАС. 1 2 5, Ст, Ру, M; = 34 of М = 36. The neonatal pelage is white with a creamy tone; after the first molt the hair coat is spotted. The females are somewhat smaller than the males. There is one pair of teats. Ecologically, the genus is represented by two forms. One form inhab- its the Baltic Sea and also partly the Canadian-Newfoundland waters and is biologically associated with ice floes. These seals reproduce, suckle the pups, and molt on ice floes. The herd instinct and polygamy are less characteristic of this form. The population of the second form, inhabiting unfrozen waters, exhibits no affinity for ice floes, spends part of the time on the coasts, and forms rookeries of a distinct harem type. Whelping, lactation, and partly even molt occur on the coast. The distribution of the genus is restricted to the coastal waters of the boreal belt of the North Atlantic and is now mainly confined to three isolated portions of this belt: western Atlantic (American), east- ern Atlantic, and Baltic. The American section is bound by waters from the southern boundary of the Gulf of Maine almost up to northern 454 Labrador, including the Gulf of St. Lawrence and Newfoundland coasts. The European section comprises the waters surrounding Iceland, the Faeroe Islands, Ireland, and Great Britain, and also the coastal waters from Bretagne and La Mancha to eastern Murman, including the North and Baltic seas, and also the inlet zone of the White Sea. The boundaries of distribution are mainly determined by the lines of the polar front and heavy pack ice, continental shelf zone, and boundaries of the temperate boreal waters. The distribution as a whole is characterized by interrup- tions, being broken into fairly distinct isolated sections. Thus one of these, the Baltic, is isolated from the eastern Atlantic which, in turn, is wholly isolated from the western Atlantic. The origin of the genus has been traced to the Pliocene as genus Gryphoca van Bened. Deeper roots of the genus have not been detected. It is possible that the Miocene seal, Miophoca vetusa Zapfe, represents only one of the much older ancestral branches of this phylogenetic group. These seals exhibit the most proximate contemporary genetic asso- ciations with seals of the genus Phoca (especially of subgenera Phoca s. Str. and Pusa) together with which they form the subtribe Phocina (Chapskii, 1955). The genus consists of a single species, the gray seal or “tevyak,” Най- choerus grypus Fabricius, 1791, which constitutes 5.5% of the number of species in the family. There is no basis for assuming that the genus con- sisted of a larger number of species in the geological past. The economic importance of gray seals is not significant. The USSR is host to this single species, or 7.7% of the number of pinnipeds in our fauna and 0.3% of the number of species in our mammalian fauna. Its distribution is restricted to the Baltic Sea and southwestern part of the Barents Sea. The economic importance of the genus in our waters is altogether negligible. At places, some probably damage the fish reserves by consuming particularly valuable fishes, i.e., salmon and eels. (K.Ch.) GRAY SEAL!'® Halichoerus grypus (Fabricius, 1791) 1791. Phoca grypus. Fabricius. Scrivter of Naturhist.-Selskabet, Kjoben- havn 1, p. 167, pl. 13. Greenland. 118 “ТеууаК” is a name used by the coastal people but is not less common. Known as “zhirovets” at places in Murman. Other names—“long-snouted,” “gray,” “hump-nosed,” “pig,” and other artificial, bookish, often translated names—are almost out of use since the coastal inhabitants are little acquainted with these names or this species. 342 343 455 1820. Halichoerus griseus. Nilsson. Scand. Fauna, Dagg. Djur., I, р. 377. Greenland. 1851. Halichoerus macrorhynchus. Hornschuh et Schilling. Arch. Natur- gesch., 17, p. 28: Baltic Sea. 1851. Halichoerus pachyrhynchus. Hornschuh et Schilling. [bid. Baltic Sea. 1886. Halichoerus grypus var. atlantica. Nehring. Sitz.-Ber. Ges. Natur- forsch. Freunde, Berlin, p. 122. Western coast of Norway. 1886. Halichoerus grypus vat. baltica. Nehring. [bid. Baltic Sea. (V.H.) Diagnosis Only species of the genus. Description General build of body massive, without distinctive features, while pre- serving all the typical features of the subfamily of true seals (Fig. 185). However, the snout is elongated, high; the profile at the eye level shows no curvature. Some skin #014511? are seen posterior to the ear opening. The whiskers are set in six rows on the upper lip; the first to the third rows from below have 7-9 each. On the fore flippers, the first digit is the longest (with the claw), the second slightly shorter, while the length of the others decreases markedly toward the last. The color of the hair coat reveals considerable individual variation depending on age and sex and, further, is subject to some geographic variation. The coloration essentially consists of two elements: the gen- eral background and the fairly dark spots dispersed on it. The background and partly the spots exhibit a wide range of gray tones and shades, from extremely light silvery-white to, at places, almost white to very dark, and even totally black. The main background in most cases is considerably lighter than the spots and is usually different on the dorsal and ventral sides, the latter almost invariably being perceptibly lighter. The spots are of diverse sizes and shapes; the sharpness of their contours, number, and disposition also vary. From very early times, many authors (Millais, 1904; Collett, 1911-1912; Ognev, 1935; and others) attempted to group all the 19 This formation was described by Pocock (1933) who regarded it as a rudiment of the outer ear shell. However, these skin prominences, measuring 3 to 15 mm (Mohr, 1952), cannot be regarded as typical of the species. There is even less justification to view them as analogous to the rudiments of outer ear shells. These skin folds are altogether lacking in the Baltic gray seals. 342 456 Fig. 185. Gray seal (‘ЧеууаК”), Halichoerus grypus (figure Бу М.М. Kondakov). diverse colors into two main types. One type covered animals with a rel- atively lighter, contrasting spotted coloration and the other type animals with a much darker, dull-spotted coloration. According to this scheme, the light gray main background on the dorsal side of animals with light coloration is somewhat darker and covered with innumerable, predomi- nantly dark gray, blackish spots, sometimes scattered singly though quite densely and sometimes gathered in clusters which are often fused. Some spots have distinct contours while others are quite diffuse. The angular, sometimes complexly contoured spots on the sides of the neck, on the chest, and on the shoulder blades exhibit the utmost contrast. The spots at these sites are also the most vividly pigmented, at places almost pure black. On the dorsal side, the spots are duller and generally lack sharp outlines. The dark-colored type animals are distinguished by a very dark gen- eral shade that depends not only on the darkening of the main back- ground, but also on the intensity of pigmentation of the spots covering it and the increased number of spots or their sizes. Concurrently, the contrast of the pattern either disappears or weakens due to the indis- tinct, diffuse contours of the spots. When the background is particularly 344 457 darkened and the spots are fused, the animals appear from a distance to be monochromatically dark-colored, almost black. On closer exam- ination, the spots are definitely apparent; furthermore, small, sparsely scattered, light-colored specks are discernible on the dark skin, either in streaks or blotches. Various transitional coloration forms are noticed between these two extreme types. The following features are characteristic of males as well as females among all the diverse types. The dorsal side is gray or bluish- gray, with fairly innumerable almost black spots; the underside of the body is almost white with highly contrasting black spots that are partic- ularly prominent in the anterior portion of the body. In spite of the extensive studies on the color variation of this species, its earlier descriptions (Millais, 1904, and authors citing Millais; Ognev, 1935; and others) are no longer adequate, 1.е., attempts to differenti- ate two main color types—light- and dark-colored animals, or at least the greater number of them. The information collected in the last two decades has provided data pertinent to the prevailing situation. There is no doubt about the existence of predominant color types but these can now be more conclusively interpreted as individual, age-related, sex- related, and evidently geographic variations. The magnitude of this vari- ation differs in different population groups. Evidently, its range is most extensive among the land-loving (pagophobic) gray seals inhabiting the European and American Atlantic. The Pagophilic Baltic seals are evi- dently characterized by a relatively greater color stability. Only the Murman gray seal can be regarded as a representative of the Atlantic population in our waters. Its color is very similar to that of the British-Irish populations and is perhaps indistinguishable from the color of the latter. It is characterized by as much range in individual varia- tion as in the first year after the initial neonatal molt (Fig. 186). Among such molted pups, the color intensity of the main background as well as the nature of spots (size, shape, and sharpness) and the intensity of their color vary; the color may be very light or sometimes very dark “on any general background”. Wholly monochromatic black-colored animals are also encountered (Karpovich, Kokhanov and Tatarinkova, 1967). A similar color variation is seen among the juveniles of British-Irish and Norwegian populations. In these regions pups are encountered with and without spots, with a bluish-gray main background, even greenish, some- times creamy-white or dark-colored or pure black; some animals are brownish or even reddish-brown (Millais, 1904; Collett, 1911-1912). Color variation among the adults is even more diverse. At times, it is difficult to find two animals with totally identical coloration. Never- theless, the color of Murman gray seals does not go beyond the various ааа Tee ~ съ SRG SR Ne RA SS 344 Fig. 186. Individual color variation of molted juvenile gray seal, Halichoerus grypus. Great Ainov Island, Murman coast (after figures by V.D. Kokhanov). shades of gray and varies from almost white to almost or even totally black. This is true of the main background on which the spots are dis- persed and of the spots themselves. The main background is more often of light gray shades, sometimes whitish and at places almost totally white; specimens with a very dark, almost black background are not very rare. 345 On the underside of the body, the background color is somewhat lighter 459 than on the upper side, but this difference can sometimes be evened out by the significantly greater spottiness on the throat, chest, and belly, compared to the dorsal side on which the spots are frequently duller. The spots vary widely in shape, size, sharpness, color intensity, and density. In most cases the spots are diverse, often of queer shapes and extremely dissimilar in size even in the same animal. The larger spots have sharp as well as highly diffuse contours. The spots on the flanks and underside of the body are the most intensely pigmented and contrasting; they are often totally black. The spots are usually 10-15 cm in size; innu- merable much smaller spots, down to mere specks or dabs, are scattered between very large ones as well as separately from them, descending onto the throat, occiput, and the base of the fore flippers. Extremely matte and comparatively small spots sharply predominate on the dorsal side. Thus the entire dorsal surface very often appears from a distance light- colored and monochromatic. At the same time, animals are encountered with totally dark or black coloration, this being characteristic of males. The main background as well as the spots never contain admixtures of brown, cinnamon, rust, or yellow shades but these are encountered, albeit very rarely, among the animals in British rookeries.!”° A very similar sharply manifest sexual dimorphism points to the genetic kinship of the Murman gray seals with the other gray seals of the open Atlantic waters breeding on the land. Adult males are distinguished from females by a very dark and fairly monochromatic color. Males are not characterized by contrasting spots and their body underside is only slightly in color than the upper side. On the contrary, brighter coloration is characteristic of the females, among which the main background is much lighter, especially on the flanks and ventral side, where it is very light. The spots on this background are highly contrasting but vary in degree of sharpness of contour, which is sometimes well defined and sometimes highly diffuse. These spots are concentrated predominantly on the front portion of the body where even contrasting specks stand out.'?! The age-related color variation among the pagophobic and pagophilic forms has not been thoroughly studied. Evidently, in the early stages of postnatal growth, their color differs only as much as the groups under comparison differ in the range of individual variation. Pups of 120 The above color characteristics of the Murman gray seal are largely based on the reports of V.D. Kokhanov, a zoologist in the Kandalaksh sanctuary. 121 The contrasting fanciful patterns of bright black coloration of the Murman gray seals renders them visible from a distance even in twilight hours. Thus, several tens of animals lolling on the coast appear “surprisingly similar to a herd of spotted cows at rest. Their white skins appear variegated with large and small black spots” (Belopol’skii, 1951). 346 460 different ecological groups form herds of unmolted white pups. In most of the pups of the Atlantic population, the short hair coat after the long embryonal hair has been shed is almost the same as among the pups of the Baltic seals. Its typical features are as follows: The main background is pale gray, usually somewhat darker on the dorsal surface than on the ventral. Comparatively small but numerous dark gray or even Olive-colored spots, often fused, are scattered fairly uniformly on the upper as well as the underside of the body. The sharpness of the spot edges varies in different sections of the body of even the same animal as also in different animals; in the anterior portion of the body, however, and in particular on the flanks, chest, and neck, the spots mainly have sharp outlines and hence stand out contrastingly. Such a coat among the juveniles usually appears brighter and somewhat “fresher” than among the older seals. The molted pups of Murman reveal up to 12 distinct color varia- tions which can be grouped into the following six classes: (1) almost black, monochromatic; (2) dark gray, with very dark spots only on the trunk; (3) similarly dark gray but with spots on the head as well as on the flippers; (4) close to black, with light-colored spots; (5) with light gray main background and dark, almost black snout and extremities of fore and hind flippers; and (6) with the same light-colored background and slightly darker colored head, flippers, and spots of the same shade scat- tered on the trunk (V.N. Karpovich, V.D. Kokhanov, I.P. Tatarinkova). The pups from the Baltic Sea after shedding the embryonic pelage appear more monotypical. They are then gray with innumerable blackish, partly fusing small spots. Instances of black or almost black coloration have not been seen among these pups. The spottiness covers the entire head and extends to the occipital, temporal, and throat regions (Lillje- borg, 1874; Allen, 1880; Schubart, 1929; Lonnberg, 1929; Freund, 1933; Mohr, 1955; and others). The subsequent color changes of the hair coat until the animal attains maturity have not been adequately understood. Some authors (Collett, 1881; Millais, 1904; Mohr, 1955; and others) believe that the original vivid and contrasting spots (with sharp contours) are gradually lost as the animals grow up, giving place ultimately to a uniformly gray or very dark coloration. But this is not actually so. There is no single pattern of age-related color variation that can be regarded as common to all the animals or to different populations. This is indirectly supported by the considerable diversity of the original coloration of the pups, espe- cially the specimens that are found to be almost black from the moment of casting the embryonic pelage. Although there is no possibility of their turning blacker later, even these pups change color. Thus, one gray seal 347 461 that lived for several years in the basin of the Bergen Biological Station was almost black with a brown tinge at the age of one month; it began to lighten in color after two years (especially the main background between the spots) and roughly ten years later again turned very dark (Collett, 1911-1912). Observations of two juveniles sporting different colors in the Berlin Zoological Garden showed that the hair coat of the light- colored animal became even lighter and spottier after each molt, while that of the dark-colored animal, on the contrary, became darker so that the spots became increasingly less distinguishable (Nehring, 1887). Whatever the pattern, the pup at the transitional age is characterized mainly by a transitional color with larger, more contrasting spots than pups that have not yet attained the final pattern (or color type) char- acteristic of the adults of the corresponding sex. Evidently the young animals do possess some features reflecting their affinity to one or the other ecological groups: Atlantic or Baltic. The growing juvenile from the Baltic Sea is apparently characterized by a gradual loss of the con- trast of spots from year to year; the spots either become less distinguish- able on the generally darkened background (Ropelevskii, 1952), or they disappear altogether (Mohr, 1965), or become diffuse or sparser. Among the males in general, the spots are lost sometimes in a wholly dark back- ground (Aul, Ling and Paaver, 1957). The spots are less sharp among young animals of transitional age and among the Canadian population (Mansfield, 1963). On the other hand, the view has been expressed that the color and spottiness of at least part of the British-Irish gray seals are as diverse among young animals as among adults (Millais, 1904). On the whole, a clear picture of the age-related succession of color changes among the various populations is not yet possible. The process of color variation in the much later stages of age pro- ceeds divergently, i.e., begins to show sex-related differences. Concomi- tantly, changes in the configuration of the spots characteristic of the males and females continue and many spots enlarge and lose their sharp- ness of contour. The color of the adult males of the Baltic seals is generally quite uniform and falls in the category of spotted coloration described above, being only somewhat duller. The main background of the dorsal side of the body is Smoky-ash with dove-olive tones. It is noticeably darker on the flanks and on the ventral side and the boundary of the back- ground color is often quite distinct. Innumerable dark-colored spots are scattered all over this general background. On the dorsal surface, they are mostly small with diffuse outlines that are generally regular (less often, highly sinuous) and not very contrasting. On the flanks and ven- tral side the spots are denser, appear very dark (black), often fuse with 462 each other, acquire very complex outlines, and stand in sharp contrast against the light-colored background. The spots are noticeably larger in the anterior half of the body than in the posterior half; the spottiness on the belly is paler and quite often looks like marble streaks. The spots become smaller around the head and on the throat and look like dabs. Fine specks are sometimes seen even on the occiput, sometimes extend- ing also onto the sinciput; on the sides of the head, near the eyes and ears, however, the spot-dabs are as numerous as оп the throat. The snout and the upper portion of the head are also quite often monochromatic, grayish dove-colored. Quite often, too, fine specks descend even onto the fore flippers. On the lower side of the neck and on the chest, the spots form a typical pattern, more often a longitudinally elongated garland of fused dark gray spots with diffuse contours standing out sharply on a very light-colored background. On the belly the spots are not only paler, but their outlines more diffuse. Adult female Baltic seals, compared with males, are generally lighter in color and less spotted (Aul, Ling and Paaver, 1957). It is possible that the spot pattern on the neck and chest is better developed among females than males (brighter and more contrasting). The hair coat fades somewhat in the course of the year. Among the British-Irish and Norwegian populations, the skin turns very pale by spring and the spots lighten; they turn a vague brown on the dorsal sur- face while the main background between them fades to a monochromatic straw color (Millais, 1904; Collett, 1911-1912). Sex-related color differences are quite sharp among the Murman gray seals. Adult males are usually dark, almost black, with spots that appear indistinct from a distance. Only the underside of the body is lighter with dark spots being fairly distinct. The females are lighter and more spotted than the males although dark females do occuroccasionally (V.D. Kokhanov). Seven basic color variations have been recognized among the females. The males and females exhibit deviations (roughly 12%) wherein the males exhibit the color features characteristic of females and vice versa. Sex-related color differences are seen among Murman gray seals right from the earliest age, 1.е., as soon as the pup begins to shed the original (embryonic) pelage. The main body background in most young females is light gray with scattered darker gray spots. Less frequently, the spots are black on a light gray or even darker background. A very dark, almost black main background (lighter on the underside) predominates among the males; in the first case the scattered spots are black and in the second light colored (lighter than the main background) (V.D. Kokhanov, V.N. Karpovich, 1.Р. Tatarinkova). 348 463 Sex-related color changes are also similar among the Norwegian (Collett, 1911-1912) and British-Irish (Hewer and Backhouse, 1959) populations of gray seals. This confirms the greater affinity of the Mur- man gray seals with the European-Atlantic population than with the Baltic. The adult males of the Norwegian population are dark-colored, almost totally monochromatic blackish-brown; the lighter-colored ani- mals show a dark gray main background alternating with large brownish- black spots. The females are lighter in color: the main background is either brownish-gray or silvery-white as in the juveniles and the spots sometimes more, sometimes less vivid; sometimes spots occur only on the shoulders; some animals are totally devoid of spots and appear fairly monochromatically white from afar; the spots vary from light brown to very dark (Collett, 1911-1912). The sex-related color differences of the hair coat are similar among the British-Irish populations. The main background in adult males is a uniform dark color interrupted quite often by not very contrasting and rather small clearances in the form of angular white spots and motleys, sometimes with complicated patterns. In females, innumerable dark spots are scattered on a very light-colored main background (usually differing in vividness on the dorsal and ventral sides). These spots are invari- ably darker than the main background and the contrast is quite good in most females. Among the dark-colored males, the main background is close to black with indistinct small and narrow white streaks show- ing through. Among the dark-colored females, the main background is dark gray and the spots are numerous, quite large, and deeply pig- mented, almost black. Light-colored males are distinguished by a very pale, brownish-gray main background covered with very bright, some- times almost white streaks. Light-colored females have a light gray main background covered with comparatively sparse and small spots (Hewer and Backhouse, 1959). Coloration is also similar among the Canadian gray seals. The males sport a dark gray, almost black, background color patterned with tiny dark gray spots on the flanks which may fuse into large groups. The overall impression is a very dark coloration. The color of the females is smoky-gray dorsally, turning silvery-gray or even white along the flanks and on the belly. In much older animals, the dark gray or black spots may have fused. The female in general is lighter in color and more spotted than the male (Mansfield, 1963). Among the females in the British section of the range, stray speci- mens are encountered, albeit very rarely, with an orange-rust-red head, neck, and lower surfaces of the flippers, and sometimes rust-colored flanks; elsewhere the body color remains normal (Backhouse and Hewer, 349 464 1957, 1960). Animals with this type of coloration have not been reported in our waters and it has been emphasized that even brown shades are not found among the Murman gray seals (V.D. Kokhanov). The skull is massive, the facial portion appears elongated, and the rostral portion broad (Fig. 187). Among adults the width between the zygomatic bones noticeably exceeds that of the skull between the mastoid processes; the maximum lateral bulge of the zygomas falls close to the posterior margin of the orbit, in the posterior half of the skull. The interorbital space is moderately compressed (its width is not less than 15 mm among the adults). The general appearance of the skull resembles more that of a large larga or Atlantic common seal but differs in the structure of the anterior nasal opening, nasal bones, massive rostral portion, nature of crests, form of the lower jaw, and especially the sharp structure of the crowns of the premolars. The broad and short nasal bones have a shortened apex, forming more than half a right angle, with rectilinear sides. The anterior margin of the nasal bones has three deeply incised denticles. The nasal processes of the pre-maxillary bones reach the nasal bones but only adjoin them (not invariably though) and do not wedge deeply between them and the upper maxillary bones. The longitudinal bony septum in the choanae falls far short of the posterior edge of the bony palate incised in the form of a deep arc. The anterior palatine openings are large, with a deep troughlike pit anterior to each. The width of the tympanic bullae together with the bony lobe of the external auditory meatus exceeds their length. The sagittal crest is straight, narrow, and relatively low. Most of the premolars and the upper molars have a high conical crown, without accessory cusps. The latter are developed, though not intensely, only on four premolars and lower molars (and often also on the lower initial premolars). These are in the form of small denticles disposed one each anterior and posterior to the main cusp at its base. Intensely reduced and barely visible denticles are seen even on some other pre- molars at the base of the crowns. The canines are relatively large (the longitudinal diameter of the alveolus is more than 10 mm); the extreme lateral incisors in the upper jaw are somewhat larger than the rest. Individual skull variations are significant in terms of absolute units as well as ratios. The structure of the crown of premolars is variable. Sex-related differences in the skull ar manifest only in sizes and some ratios; the skull of males is larger. Age-related skull variations are seen primarily in the elongation of the skull due to the predominant growth of the facial portion which becomes more massive and increases in height. Among young animals, the facial portion falls much below the occipital and the line of the upper 349 350 465 Fig. 187. Skull of the gray seal, Halichoerus grypus (figure by М.М. Kondakov). profile of the skull is intensely inclined toward the anterior margin of the nasal bones. The width at the zygomatic arches is less developed among young animals than adults; it does not exceed the width at the mastoids in the young; the width of the rostrum at the level of the upper canines too is relatively smaller than in adults. The contour elements of the cranium (crests) are almost undeveloped among young animals. The sagittal crest is not manifest: instead, only folds are seen to the right and left of the median. These folds slightly diverge sideways and form a small elongated area; occipital crests, seen as cornices in adults, are 466 practically absent in the young. As in other seals, the roots of the teeth are initially hollow and thin-walled but intensely thicken over time and | the pulp cavity fills in. Among many old seals, the roots are unusually hypertrophied due to the cement layers while the crowns are worn down almost to the base. Over time, all the skull bones thicken, as a result of which the overall weight of the skull also increases. The individual and partly the geographic variation of body size is significant; females are generally smaller than males. The body length of the adults measured from tip of nose to tip of tail along the dorsal surface (Lc) varies from 170-250 cm. This length in the largest animals can reach 3 m. However, various authors cite different values: 2-2.5 m (Freund, 1933), 2.5-3 m and at places where these seals are killed most intensely (southern Baltic), mostly 2-2.5 m (Mohr, 1952). According to the old data (Lilljeborg, 1874), the body length of the Baltic male gray seals is 195 cm and that of females 175 cm. Data on the body length of male Murman gray seals are extremely limited and not wholly reliable. According to some authors, the average size of an adult animal is 2.5-3 m (Skvortsov, 1928*) and according others (Belopol’skii, 1951), the males are 2.5 т long. A large male caught on Franz Josef Land measured 263 cm along the dorsal surface (Lc) (Tsalkin, 1936). The body length of gray seals of the British and Irish populations (from which the size of Murman gray seals can be judged to some extent) varies from 245 to 260 cm in males (largest male measured 292.5 cm) and 168 to 198 cm in females (Millais, 1904). The average length of adult Canadian male gray seals is 235 cm, of females 200 cm (Mansfield, 1963). The available information on weight is also in general terms. It ranges from 130 to 320 kg for seals of both sexes. During lactation, the females lose over 30 kg of weight and males in the mating period even more than 50 kg (Collett, 1911-1912). The condylobasal length of the skull is 240-270 mm and its maxi- mum length [?] 330 mm (Collett, 1911 - 1912). This length among females is considerably less than among males: 264 mm in the Baltic male seals and 240 mm in females (Lilljeborg, 1874). In the largest males of the Atlantic region, it reaches 320 mm (Allen, 1880) and even 330 mm (Col- lett, 1911-1912) white it hardly exceeds 260 mm in the largest of the females. The width of the skull at the zygomatic arches reaches 200 mm. (K. Ch.) Taxonomy Only species of the genus. 351 351 467 Geographic Distribution Temperate latitudes, partly the cold belt of the North Atlantic from Canada and the adjoining regions of the USA to the coasts of central and Northern Europe and the Baltic Sea. The range is broken into fairly isOlated sections. Geographic Range in the USSR (Fig. 188) This covers the eastermost part of the world distribution and includes two distinct regions: coastal waters of Murman and the Baltic Sea. The gray seal is distributed all along the coastal belt of Murman from Varanger Fjord to St. Nos Strait and possibly farther in the southeast up to the western threshold of the neck of the White Sea. However, the find of the gray seal on the eastern extremity of the range is evidently sporadic and it is only very rarely that this seal is found in the neck region itself (Schrenk, 1955*; N.A. Smirnov, 1929; Kirpichnikov, 1932). Details of the seal distribution and the disposition of all of its coastal Fig. 188. Distribution of gray seal, Halichoerus grypus in the USSR (K.K. Chapskii). 352 468 rookeries have not yet been established. The gray seal reproduces for certain or resides regularly on the Bolshoi (Large) and Malyi (Little) Litskie, Veshnyak, Kuvshin, Malyi Zelenets, Kharlovskie (“baklyshes”), Semiostrovnye (“ludy”), Ainov and other islands, and in the region of Dal’nii Zelenets, lokan’gi, Teriberki, and Pala Bay. The gray seal whelps presumably in the bays of Rakushnaya, Voyak, Fedotovsk, and Kora- bel’naya (Karpovich, Kokhanov and Tatarinkova, 1967).122 The view that the eastern boundary of distribution extends up to Novaya Zemlya, based on the record of this seal by K. Ber on the west coast of Novaya Zemlya, is erroneous. This is an exceptionally rare event of the straying of an animal far beyond the range of its distribution. Very similarly, a lone gray seal was also caught in Franz Josef Land archipelago (Tsalkin, 1956*). The same is the situation with regard to the White Sea, which was included in the distribution of this species (Schrenk, 1855*), although episodic finds of lone animals have been reported from eastern Murman in the inlet region and the adjoining neck regions (N. Smirnov, 1929; Kirpichnikov, 1932; A.G. Beloborodov; and others). It is quite likely that lone animals even more rarely can penetrate into the northern part of the neck region too. In our Baltic waters the gray seal is encountered from time to time in the northern coastal belt of the Gulf of Finland from the boundary with Finland to Nevsk Bay, while it is quite common in the southern coastal zone commencing from Kopor Bay and farther west. It is encountered on the Estonian coasts all along the Gulf of Finland but is mainly confined to regions far removed from the mainland coast, with a preference for the tiny islets scattered there (Aul, Ling and Paaver, 1957). It inhabits Irben Strait and the Gulf of Riga (its Estonian as well as Latvian sec- tions) including Rukhnu Island. Evidently, from time to time, especially in water, it is seen in the Latvian marine coastal waters. Rare encounters in the coastal belt of the sea within Lithuania and Kaliningrad districts are possible since gray seals were found in the past on the coasts of the former eastern Prussia, Pomerania and Ryugen Island, as also on the coasts of the former Polish corridor. 122 The first reference to the distribution of the gray seal beyond the Polar Circle in our territory was made by I. Lepekhin (1805). Concrete evidence of the catch of gray seal in Varanger Fjord (Pleske, 1887), on the coasts of eastern Murman (N. Smirnov, 1903), and on Litskie Island (Telegraph of Murman Research and Hunting Expedition, 1906) became available much later. It took an even longer time for more detailed information to become available (Skvortsov, 1928*; Sdobnikov, 1933*; Belopol’skii, 1941, 1951; У. Uspen- skii, 1941*). New data have been collected in recent years by the workers of the Kandalaksh Sanctuary. 353 469 Geographic Range outside ше USSR (Fig. 189) In the Baltic Sea, the range covers the Finland portion of the Gulf of Fin- land, Gulf of Bothnia, northern and partly central regions of the Baltic Sea itself (from the west, its Swedish belt). To the south of Gotland Island, it is encountered more rarely, in the south up to Eresunn Strait; it is rare in the southern and southwestern parts of the sea, including the coastal waters of the Polish People’s Republic, the German Democratic Republic, on the adjoining coasts of Denmark, and the Federal Repub- lic, on the adjoining coasts of Denmark, and the Federal Republic of Germany where it is encountered sporadically. The gray seal is evidently even more rare now in Denmark Strait although the breeding of this seal in the past on Anholt Island in Kattegat has been recorded. Outside the Baltic Sea, the population in the eastern Atlantic section is concentrated mainly in three regions where the seals breed and reside permanently: 1) region of Great Britain and Ireland including the Hebrides, Orkney, Shetland, and Faeroe islands; 2) coastal waters of Iceland (excluding evidently the northeastern section); and 3) Norwegian coast, predominantly its middle portion where the most important (though not many) breeding sites are concentrated. In general, however, the boundaries of distribution in the European part include the coastal mainland from Bretagne (sporadic), La Mancha, Pa-de-Kal along the southern coast of the North Sea up to the Jutland Peninsula, joining there with the Baltic and Norwegian sections of distribution extending up to Nordkapp. The presence of this species on the coasts of Spitsbergen is rather doubtful in spite of positive references to it (Saemundsson, 1939). There are no settled populations on the coasts of Greenland but stray animals and probably even small groups wander there from time to time (Brown, 1868; Allen, 1880; Winge, 1902; and others). The American part of the distribution includes the Gulf of St. Lawrence and the adjoining regions of open coastal waters in the north from the Strait of Belle Isle along Labrador roughly up to the latitude of Hebron (about 68.5° N lat.) and in the south partly along the west coast of Newfoundland, to the south and southwest of this island and Cabot Strait, including Sable Island, coastal waters of Nova Scotia, Gulf of Maine, and up to Cape Cod inclusive. (K.Ch.) Geographic Variation The geographic variation of the gray seal has not been adequately studied. Nonetheless, the species distribution is quite fragmented into distinct 352 470 Fig. 189. Species distribution of the gray seal, Halichoerus grypus (К.К. Chapskii). sections with extremely incomplete contacts between them. First of all, the western Atlantic (American) zone of habitat of the species is practi- cally isolated from the eastern Atlantic (European) zone since gray seals do not reside regularly in the waters of southern Greenland and are sighted rarely and irregularly. The Baltic zone of distribution is largely isolated from the Atlantic Ocean zone. In the remote past, gray seals resided undoubtedly even in the Denmark Strait and a contact between the populations of the Baltic and North seas did exist though not wholly constant. A real interruption of distribution evidently prevailed for some centuries (for more details, see “Geographic Distribution”). Two subspecies of gray seal can be recognized in our waters at present. 1. Baltic gray seal or “tevyak,” H. g. macrorhynchus Hornschuh et Schilling (syns. pachyrhynchus, baltica) (Fig. 190). The body and skull dimensions are comparatively smaller than those of the nominal form. Sexual dimorphism in dimensions is not very sig- nificant. The color is generally very light with fairly bright, though often diffuse, contrasting spots on the chest and neck with relatively less vari- ation. There are practically no animals in the population with intensely dark, almost black coloration. Sex-related color differences are relatively insignificant. It occupies Gulf of Finland and the waters of Estonia, Latvia, Lithua- nia, and Kaliningrad district. 353 354 471 Fig. 190. Baltic gray seal. Leningrad Zoological Garden (photograph by К.К. Chapskii). Outside the USSR, the Baltic gray seal is found in the rest of the Baltic Sea regions. The form differs not only in morphological, but also in its biological relation with ice floes on which it breeds (pagophilic form). 2. Atlantic gray seal or “tevyak,” НЯ. в. grypus Erxleben, 1777 (syns. gris- seus, gryphus, halichoerus, gris, thienemanni, atlantica) (Fig. 191). The body and skull dimensions on average are slightly larger than in the preceding form and sex-related size differences are more significant. The color on average is darker with somewhat less sharp and con- trasting but extremely variable spottiness. Animals are encountered with very dark, almost black coloration. Sex-related color differences are more intense than in the preceding form. It occupies the Murman coast. Outside the USSR, this subspecies is found in the rest of the area of distribution of the species, at least in the European waters in addition to the Baltic Sea. Seals of this form exhibit no affinity for ice floes during the period of breeding (littoral form). Assigning the nominal name of the subspecies to Murman and gener- ally to the European seals of the oceanic waters is presently still tentative since the waters of southern Greenland represent the terra typica of this form. It may be assumed that the animals existing there are identical with the American form. However, a comparative systematic analysis of the European and American gray seals has not been attempted to date. Yet, while taking into consideration the quite complete (though not absolute) 354 472 Fig. 191. Murman gray seal in water. Great Ainov Island, Murman coast, Мау, 1964 (photograph by V.D. Kokhanov). isolation of these two large populations, it must be considered that they are not identical at the subspecies level. If this is so demonstrated, the name grypus should be given to the American gray seals while their Euro- pean counterpart should be designated by the closest recent synonym. Nevertheless, the systematic homogeneity of the American form may give rise to some doubts since its composition includes pagophilic as well as aegialoid [littoral] populations. It is possible that some differences do exist between the various populations of seals in European waters.!79 Outside our waters, these animals are not classified into particular subspecies. Biology Population. Even an approximate census of the population has not been possibie in our Baltic Sea waters. It was difficult to establish even the relative sizes of the local populations. Evidently, these seals are least numerous in the easternmost portion of the Gulf of Finland in the region of Vyborg (Freund, 1933). In Nevsk Bay only rarely are stray young 123 According to the latest data (К.К. Chapskii), fairly stable craniological differences do exist between the Atlantic and Baltic subspecies. 355 473 seals encountered and that, too, only in the summer-autumn season; the population increases westward. This seal is regarded as common in Estonian waters. In the Gulf of Riga it is relatively more numerous around autumn, especially during the formation of ice floes, in search of which the seals traverse Irben Strait (Aul, Ling, and Paaver, 1957). South of the inlet into the gulf their number decreases sharply down to rare single ani- mals for many kilometers of the coastal zone. The nature of the coastal zone, especially the ruggedness of the coastline, and also the extent of human habitation and hunting activity—all these greatly influence the population of this seal. The numerical strength of the local seal population changes sharply in different seasons of the year; thus a greater concentration of the seals occurs in the period of breeding and a sparse population in the feeding season. Factors such as the formation of ice floes, their distribution and suitability for whelping, as also the availability of food play a decisive role. While conducting a census of the world reserves of the gray seal, the strength of the Baltic Sea population was also evaluated. This evaluation was mainly based on information regarding the number of animals killed. The statistics were incomplete because of the nonavailability of data for our waters. In the first evaluation of the world reserves of gray seals (Lockley, 1954), a figure of 5,000 was cited for the Baltic population, exclusively for the Gulf of Bothnia. The view was later expressed that the total reserves in the Baltic Sea should be in the tens of thousands (Davies, 1957). Practically, however, in subsequent evaluations the total Baltic population was roughly put at 10,000 (Haglund, 1961) ог 5,000 - 10,000 (Hook, 1964*; Curry-Lindahl, 1965). A more realistic figure lies between 7,000 - 10,000. With this background, it is as yet difficult to reckon the population in the waters of the USSR. According to preliminary data (1972), the mother population of our gray seal in the main region of its winter-spring concentration, i.e., in Irben Strait with adjoining sections of the Gulf of Riga, was roughly put at 350-400. In our zone of the Gulf of Finland, however, the strength is a mere 50 (V.A. Zheglov). The total strength of our Murman population was put at 300 in foreign compilations (Smith, 1966*). A direct count of the newborn on all the islands of the archipelago of Semi Islands and on the Ainov Islands in the early 1960s identified a total of 200 seals. By applying the method of extrapolation to other regions in which breeding is wholly likely, the total population of annually reproducing females of gray seals has been put at 500-800 (Kandalaksh Sanctuary; Karpovich, Kokhanov 356 357 474 and Tatarinkova, 1967). The main population (without pups) of Murman seals constitutes roughly 1,500. The population of the gray seal in other sections of its range is as follows: 34,000 in the waters of Great Britain of which 5,000 to 8,500 seals are concentrated on the Orkney Islands, Northern Rona, Outer Hebrides, and Farne Island; 2,000 in Irish waters; and 2,000 on the Norwegian coasts. The population of Iceland has been put at the same number as on the Norwegian coasts and on the Faeroe Islands at 3,000. On the whole, for the eastern Atlantic part of the area of distribution there are 42,500 seals (not counting the Baltic Sea and Murman population). In the western Atlantic (American) part of the distribution, the total population runs roughly into 5,000, of which 3,000 are in the Gulf of St. Lawrence and 2,000 east of Nova Scotia (Smith, 1966*). The total world reserves Of gray seals thus come to 55,000 - 60,000. Habitat. The habitat is closely associated with ecology, which for this species is of a twofold nature, as in the case of seals of the subgenus Phoca s. str. (common seal and larga). In our, as well as in international waters, there are two ecological forms of the gray seal. One of these is biologically associated with the ice floes on which they breed and molt. These are typical pagophilic (ice-loving) animals. This form includes seals of the Baltic Sea and the Gulf of St. Lawrence. The seals in the rest of the European region, including our Murman population as also some populations of the Nova Scotia Islands, reproduce on the coasts. In this season they form fairly cohesive rookeries at definite places on the coast year after year and can be regarded as wholly pagophobic. It is usually assumed, although not always justified, that the pagophobic populations of gray seals are more stationary and reside longer on the same sec- tions of the coastal zone, usually not straying far into the open sea. For rookeries, they select uninhabited, predominantly small islets, inaccessi- ble to man, and made up of smoothly rolled rocky or more rarely pebbly coasts with a flat and fairly level surface. They are disposed right on the coast, slightly above the littoral zone as well as away from it in the more elevated sections, including those overgrown with grasses. Usually, the flat rocky coasts of islands slightly above the line of high tide serve as the site of whelping on Murman but at times the semisub- merged littoral sections are also used (Fig. 192); a distinct preference is shown for smoothly- rolled boulders. Sometimes the ascent of the coast is quite steep (up to 30° or more) but matters not if it is smooth. It is not important how much of the slope is covered with a thick layer of snow. Beaches with boulders or pebbles are rarely selected for whelping (Uspenskii, 1941; Karpovich, Kokhanov and Tatarinkova, 1967). On the 356 475 Great Ainov Island (in the region of Varanger Fjord), the sites of whelp- ing fall predominantly in the littoral zone toward the open sea but are almost wholly absent on the southwestern side turned to the mainland coast. Often, they are disposed there in sections with peat clumps among tall grasses (lyme grass) (V.D. Kokhanov). Our Baltic pagophilic populations during the breeding season are disposed close to the outer rim of drifting ice floes, abounding in fissures, Cape Polenyi ©) Lake Severnoe Lake Zapadnoe O Lake Nedupoynoe AONIV. LVayy Sy 1S) (oop О — о en = ise} — Go} > axv Isl О Lake Srednee Lake Maloe Cape Gagachiya Littoral Disposition of Gray Seal pups Cape Robinson Hut Fig. 192. Figure showing the disposition of newborn gray seal pups on Great Ainov Island, Murman coast, November - December, 1963 (after V.D. Kokhanov). 476 open water pools, and partings, which are essential for keeping contact with the water. Only in very rare cases, when the winter is extremely mild, do some animals use the coastal rocks for whelping (Curry-Lindahl, 1965). Under the influence of ice floe movements, the fissures develop- ing among them, and the expansions and contractions of the open water pools, considerable changes occur in the original distribution of the new- borns over the course of time: some pups and their suckling mothers drift with the ice floes to the very edge or sometimes deep into the ice massif, thereby being islanded in the dense ice “conglomerate”. The Baltic seals are usually regarded as more pelagic (Curry-Lindahl, 1965) than the British animals whose coastal rookeries in summer are very specifically fixed. Food. In our waters this aspect has been thoroughly studied only recently. There are practically no definitive data on the food of the Mur- man gray seal. Judging from observations, however, lumpfish and cod assume a prominent position in its food (V.D. Kokhanov). Only gen- eral information is available on the food of the Baltic gray seal. On the coasts of Estonia, it feeds on Baltic herring, cod, viviparous blenny, eel, salmon, and bream (Aul, Ling, and Paaver, 1957). The animals held in the Leningrad Zoological Garden (caught in the Gulf of Riga) consume diverse fresh fishes from the Gulf of Finland and Lake Ladoga including such freshwater fishes as roach, ide, dace, rudd, bream, etc. In nature, at February end to early March, 1972, only the remnants of benthic fish, 1.е., viviparous blenny and flounder, were found in the stomach of whelped females of the Baltic population (V.A. Zheglov). The gray seal of the west European and Canadian populations is usually regarded as a consumer of predominantly a wide variety of fishes. The instance of catching a seal on a baited hook dropped to a depth of 145 m and also the repeatedly reported facts of the emergence of these seals with deep-water fish between their teeth (Collett, 1881), point to the possibility of these seals “fishing” at considerable depths. In the European seas the gray seals consume, in addition to the above-listed fishes, halibut, other flounders, herring, ling, whiting, and other cods and pike. Apart from fish, they catch cephalopod mollusks (squids—Loligo forbesi and Eledone cinerosa were noticed in the Irish Sea) and at times consume even deep-sea crustaceans (Collett, 1911-1912; Freund, 1933; Duncan, 1956; Lockie, 1962). In the stomach of some tens of gray seals caught in the Gulf of St. Lawrence, the following were identified: among fish—mackerel (?), cod, herring, salmon, hake, small winter flounders, unidentified species of flounders, skate, and shark; among invertebrates—squids, crabs, and shrimps (Fisher and Mackenzie, 1955; Mansfield, 1966). 358 477 The diet of the gray seals varies considerably depending on age, season, and local conditions. These factors are also responsible for the magnitude of losses suffered by the fishing industry (see p. 495). Accord- ing to some sources, the gray seal is such a voracious eater that it can consume its own weight in fish in a single day in captivity (Legendre, 1947). The daily ration of caged gray seals abroad is 6.8 kg (Steven, 1934, 1936). They receive a slightly larger ration in the zoological gardens of Leningrad and Kaliningrad where the seals enjoy good health. The stomach of one animal shot contained 27 completely whole eels that had evidently been swallowed in a very short period of time (Curry-Lindahl, 1965). In another animal 54 common perch and one pike, in addition to several herrings, were found in the stomach; it was said that an adult seal can consume 60 to 80 herrings daily (Friedel, 1882). However, .it should be noted that, in nature, it feeds at inter- vals during the periods of reproduction and molt. Evidently lactating females feed sporadically, with intervals of starvation occurring during the summer-autumn season. When the fish is large (not in the case of an eel), the gray seal tears it apart with the claws of its fore flippers and consumes it in bits. Smaller fish, such as the herring as well as the eel are swallowed whole (Nehring, 1887). These habits have been recorded during the food intake of seals at the Leningrad and Kaliningrad Zoological Gardens. Home range. Historically, each local population has maintained a fairly stable home range. This is primarily and especially true of pago- phobic populations. Observations of the British-Irish populations have established that the bulk of the mother population whelps year after year on the same coastal section. Nevertheless, the animals are not confined to the same site year round. Even the most settled among them abandon the rookery at the end of reproduction, lactation, and molt, wandering from place to place along the coastal belt. Therefore, establishing the dimensions of water bodies covered by a given group or a Single animal is a difficult task. The Murman seals lead a more stationary life. They are distributed in different regions of a comparatively narrow coastal strip as fairly distinct isolated local groups living in herds. Such conditions make it wholly possible to determine the home range areas of the seals in future. Stray animals are, however, encountered close to Mezensk Bay. The Baltic gray seals, on the other hand, reveal a varying affinity for given sections of their habitat. In the winter-spring period, when they are confined to ice floes, mainly close to the edges, the area occupied by them often varies in accordance with ice floe drift. Thus the area of occupancy is sometimes more, sometimes less. Under these conditions, 359 478 the density of disposition of the seals is highly variable not only from year to year, but also in the relatively short icy season. During the periods of reproduction or molt the seals on ice floes generally do not gather into groups; instead they scatter, sometimes at considerable distances from each other, although not as far apart as in the case of the larga at this time in the Bering Sea (see p. 353-354). But sometimes the animals do gather in a small section of the ice floe or even in groups forming small rookeries. At this time the concentration of gray seals is maximum. According to recent observations, gestating females are disposed quite often in small groups of two-five or large groups of 20-40 animals; sometimes they group in even larger numbers or remain singly (V.A. Zheglov). At the end of the reproduction and lactation period, the adult seals that have mated do not evidently go into the water nor do they abandon the region of whelping; the seals begin to molt. The well-fed pups also remain on the ice floe for some time more until the embryonic hair coat has been completely shed. After the disappearance of ice floes in the summer-autumn season, the seals scatter in a wide expanse and keep moving in search of food for much of the time. Hideouts and shelters. The gray seal organizes no special protection for the pups born even on ice floes or on snowbound land. The newborns lie in the open and only sometimes, as for example on the islands along the Murman coast, especially on Great Ainov, enjoy the protection of a projecting stone or a peat clump (V.D. Kokhanov). Information on the disposition of pups on ice floes in the Baltic Sea has only begun to be collected (V.A. Zheglov). For a long time there was no unanimity of opinion regarding the ability of the pagophilic gray seals to make air holes in ice floes for communicating with the water. According to some (Holm, 1921, Aul, Ling, and Paaver, 1957), the seals do make air holes and keep them open. One such hole is sometimes used by many animals. Curry-Lindahl’s (1965) reports do not confirm this view. There is no need for the gray seal to make air holes as the whelping site in the fringe zone of drifting ice floes already has innumerable open water pools. Evidently, they set out to make air holes only when needed, i.e., when the open water pools or an enlarged fissure is covered by a recently formed ice crust. Daily activity and behavior. The daily cycle of activity and the behavior of the Baltic gray seals have not yet been studied thoroughly. In general, the gray seal is a fairly gregarious animal but its herding instinct is not equally well developed among the ecologically different populations; among the pagophilics, especially the Baltic seals, this ten- dency is weaker than among the land seals. The herding reflex evidently 360 479 peaks by the time of reproduction. However, some British rookeries exist long before or after the breeding period or there are those in which the animals do not reproduce at all. The same is noticed among Murman gray seals, which quite often are bound in herds on their favorite sites on islands after the breeding season. The regime of the coastal rookeries also varies. Some exist for a long time and even almost uninterruptedly round the year, as for example on the small islets of St. Kilda to the extreme west of the Hebrides (Waters, 1965). Others, however, function for a much shorter period or are characterized by almost uninterrupted traffic for the reason that all the females do not arrive at the same time and remain on land for 2-3 weeks after whelping, their pups for an average of 4-5 weeks (Fitter, 1961), and later leave the rookery altogether. The coastal collections of animals also vary widely in size: in the British Isles the sizes vary from small groups of a few tens to huge herds of 2,000 or even up to 5,000 seals. During the periods of reproduction and feeding, the animals of different ecological groups are active during the day as well as at night except for hours of sleep, for which there is probably no fixed time schedule. These seals can sleep in water as well. Activity and ability of orientation and finding food at night are characteristic of the Murman gray seals who have lived long under conditions of the polar night, coinciding with their period of reproduction. Activity of the British gray seals right up to their successfully catching salmon in the dark has been confirmed by observations on the east coast of England in the estuary of Tweed River (Rae and Shearer, 1965). During their residence on the coastal rookeries for reproduction in our Murman region as well as in British waters, the seals do not generally feed. During lactation the whelped females of the Murman population remain silent in the water not far from the pups lying on the coast, always ready to respond to the pup’s call. Less often, they spend the intervals between sucklings on land near the pups. The nurseries there, as at other places, break up as the well-fed pups begin to enter the water. A part of the population abandons the reproductive section while others remain for a longer period, until spring, and a very small number even up to summer; at this time, the seals remain in groups in the water. In spring, coastal rookeries are often formed there again. A characteristic features of these rookeries is that the seals are disposed in clusters, quite close to each other (Karpovich, Kokhanov, and Tatarinkova, 1967). Seasonal migrations and transgressions. The seasonal migrations of the gray seals do not exhibit similarity in magnitude or regularity with those of the harp seal, hooded seal, or fur seal. Some remote similarity of these migrations is seen only with the Baltic gray seal. 480 The magnitude of migrations and the duration of residence in the sea and the density of animals of different age groups and in different sections of distribution (with varying extent of food availability) differ among the pagophilics and land seals. It is known that the young of the British population remain continuously at sea for the first two years, often quite far from the site of their birth (Davies, 1956*). In the winter-spring period, like the harp seal, but on an extremely small scale, the bulk of the gray seals concentrate in relatively limited sections of distribution where the ice conditions are most favorable for reproduction and later even for molt; at the end of this season, however, they return to the extensive feeding expanses. Thus, on our Baltic coasts, in the winter-spring season, the gray seal is usually confined mainly to Irben Strait and the western regions of the Gulf of Riga. In the ice-free period, however, these seals are more widely scattered. The main impetus for the spring-winter migrations is the need for finding suitable ice floes that ensure dependable conditions for raising the offspring. The seals seek out stable, white, coarsely broken ice floes that are not intensely hummocky and with a fair amount of open water pools. The selection of Irben Strait for this purpose is wholly justified since suitable conditions required for whelping and lactation are gener- ally available there. The seals find similar conditions also in the central eastern section of the southern strip of the Gulf of Finland. Regions of their breeding are also known outside our territorial waters. The seals begin to move to the breeding zones by late autumn, the younger animals migrating somewhat earlier (Holm, 1921). In the wintering site, they are seen in clusters by December, sometimes even in November. By March end to early April, at the end of the breeding season, the adult animals leave the suckled pups on the ice to complete molt and begin to return to their summer habitat. Initially, this process is passive with the drift of ice floes but later becomes more active. The Murman gray seal, having lost its biological link with ice floes, has become a more settled animal. For three-fourths of the year, from August end, the whole of autumn, winter, and spring months, its popula- tion, with some minor changes, is confined to the Semi and Ainov islands. In the former region the seals may be seen even in spring (before June) in herds of 20-70 animals or more on Litskie, Veshnyak, and Kuvshin islands and also in some sections of the mainland coast where they form fairly dense rookeries in spring. Such groups of gray seals are seen at this time on the Ainov Islands also. But most of the animals depart in June and only rare strays continue to remain until the end of August. The summer habitat of the bulk of the seals is not known for certain. They 481 361 begin to return to the sites on the Semi and Ainov islands by autumn, from the end of August (Karpovich, Kokhanov, and Tatarinkova, 1967). In the context of seasonal migrations of Murman gray seals, the corresponding process among their kin, especially the British popula- tions, is of interest. There are no essential differences in this respect between many British and our Murman populations. The seals remain throughout the year in the waters of western Wales, migrations here being only of local interest (Lockley, 1954). The British populations are sometimes classified as nonmigratory (Davies, 1956*). However, massive tagging of pups, commencing in 1954, has shown their great capacity to migrate and for distant transgressions. Although 85% of a batch of pups marked on Farne Islands did not leave the area of their birth, some were caught on the coasts of Holland, the Federal Republic of Germany, and Denmark, while some reached Norway and the Faeroe Islands (Hickling and Ennion, 1959*). New data (tagging at the com- mencement of the 1960s on the Orkney Islands) confirmed the extensive scattering of juveniles, including even Norway where some twenty tagged animals were recovered from 1960 through 1962 (Hickling, Rasmussen, and Smith, 1962). While these do not represent systematic distant migra- tions, they confirm the great mobility of the juveniles and their ability to negotiate long distances. This has been convincingly demonstrated by the fairly extensive actual exchange even between isolated populations, which appear extremely remote at first glance (see p. 488). Reproduction. The period of mating among this species extends on the whole for almost three-fourths of the year, from July end to March end or early April while some animals mate even somewhat later. Such a wide mating season is explained partly by the fact that the animals belonging to the ecologically different groups have their own periods of mating, partly due to extensive population variations at these periods among the seals in the British part of the distribution. The Baltic seals reproducing on ice floes mate in early spring, mostly in March (predominantly at the end of this month) or in April; it is pos- sible that mating occurs at a much earlier or even at a later period, however, in February (Lilljeborg, 1874; Freund, 1933), May (Nehring, 1886), and even in June (Neif, 1757*). It has further been pointed out that in the more northern Baltic regions, especially in the Gulf of Both- nia, the seals mate later (in June) than in the Gulf of Finland. According to the latest data (V.A. Zheglov), the period of mating in Irben Strait extends from around March 10 to mid-April. In the Baltic Sea the seals concentrate for reproduction in a fairly restricted strip of coarsely broken fringes of ice floes, usually tending in March to south of the Aland Islands, mainly in a latitudinal direction, 362 482 forming а deep loop in the Gulf of Finland. In the southern part. of the Gulf of Bothnia extensive air holes become available by this time but whether the gray seals remain in the gulf or reproduce along its periphery is not known. In general, however, the disposition of the mas- sive drifting ice floes suitable for whelping is highly variable not only in a given season, but also in different years. In such situations there is little justification for assuming fixed dispositions of productive males and the mother population with pups or the existence of sharply iso- lated populations. The affirmation that there are no definite places in the Baltic Sea where these seals can be found for certain in the season of reproduction or molt is not mere speculation (Smith, 1966*). Never- theless, judging from the data of recent years (V.A. Zheglov), the stable regions of reproduction in our waters are Irben Strait as well as the predominantly western part of the Gulf of Riga. A very small portion of the population breeds in the central and eastern regions of the southern part of the Gulf of Finland. Not much is known about the nature of the mating period. Never- theless, in spite of statements to the contrary in the literature (Curry- Lindahl, 1965), harems are not a characteristic of the Baltic seal. The seals mate two weeks after whelping. Copulation occurs in water as well as оп ice floes but more often evidently in water. Among the British gray seals, there is a definite relation between the nature of coitus and the extent of manifestation of the harem regime in the rookery. On land, these seals mate at places where the males hold firmly to their territorial sections, as for example in the largest rookeries on the northern Rona and Farne Islands. But even under these conditions, when fairly deep erosion “tanks” or “channels” are available, the animals eagerly take advantage of them for coitus. When, however, the males are confined preferentially to the sea in front of the whelping site, mating proceeds in water (Hewer, 1960). Under the prevailing harem regime of the rookery, the males holding onto a definite territory fertilize many females. Evi- dently, in water too, the more powerful males stand in an advantageous position by chasing away the much younger and weaker animals. This aspect of the life of Murman gray seals has not been clearly understood. In the Norwegian waters in the remote past, when the seal population was much larger, there were 2 to 4 or 5 females to every bull (Collett, 1881). Murman gray seals mate late in the autumn and early winter, 2 to 3 weeks after whelping or slightly later. This period essen- tially falls at the end of November to the end of December. Any overture to mating before this period (long before completion of lactation of the female) or male proximity to her pup is decisively resisted by the suck- ling female, even to the extent of biting the neck of the intruder. Mating 363 483 is completed mainly on land. In any case, mating in water could not be observed. Coitus is preceded by conjugal play in water or on the coast when the animals gently bite each other. Signs of distinct harem forma- tion were not seen among the Murman population nor were there more females than males among the adult animals. It is nevertheless possible that both monogamy and polygamy prevail on Murman. Polygamy was suggested by the disposition of pups in groups of 2-4 together (Kar- povich, Kokhanov, and Tatarinkova, 1967). Conforming to the extended period for mating, the whelping season too reveals wide variations geographically as well as among the various seal groups in the same region. Thus, in the Great Britain-Ireland region, some main groups in the population have a much extended whelping period while others have a short whelping period. In Wales, the Irish Sea, and on the western coast of Ireland, whelping extends from July through October; among Scottish-Hebrides populations, pups are encountered in September-October and on the Farne Islands (northeastern coast of England) from the second half of October to the middle of December (Coulson and Hickling, 1964). However, the bulk of the females whelp within a very short period of about a month. The view that the eastern Atlantic seals whelp increasingly later toward the south (Mohr, 1952) has not been fully substantiated. The Murman seals are distinguished by a somewhat greater stabil- ity in this respect: the period of reproduction among them extends for not more than 1.5 months in the late autumn. Even according to older information (Prigorovskii and Breitfus, 1912, 1915), newborn pups of gray seals were caught on Murman throughout December but, according to much later information, they were seen on the Semi Islands in early November (S.M. Uspenskii, 1941), from the end of November to the end of December (Sdobnikov, 1933*); they were caught on November 16 and 17, December 6 (Belopol’skii, 1941) or in November and Decem- ber (Belopol’skii, 1951). The above whelping periods in eastern as well as western Murman have been confirmed by the latest observations; they are identical and the total duration of whelping extends for 30 to 40 days. The earliest pups are born in the first ten days of November and the last ones toward mid-December. In 1963, on Great Ainov Island, two peaks were noticed in the birth of pups, one from November 10 to 15 and the second, a sharper peak, at November end to early December (Karpovich, Kokhanov and Tatarinkova, 1967). Though neither the whelping nor mating season of the Baltic seal as reported in the literature is very accurate, there are no particular dispar- ities regarding these aspects. In most cases the pups appear by February to March, the main whelping season falling in March (Lilljeborg, 1874; 484 Nehring, 1886; Aul, Ling, and Раауег, 1957) ог February end to early March (Holm, 1921) or between February end and mid-April (Smith, 1966*). Recent observations have shown that the total whelping period among the Baltic gray seals extends for one month from around Febru- ary 20; most of the pups are born from February end to March 4-5 (V.A. Zheglov). The gestation period has been put variously at 8.5 to 12 months because of the lack of accurately established mating and whelping peri- ods. In the middle of this century, it was assumed as 11 months, as in the case of other seals (Mohr, 1952). Obviously, however, the entire period from mating to parturition stretches two weeks longer, to 11.5 months. But considering that a prolonged delay of zygote implantation is characteristic of gray seals (Lockley, 1954; Backhouse and Hewer, 1956), the actual embryogeny extends for roughly nine months or so. The birth process is extremely brief, being accomplished in a few seconds; the placenta is discarded in an hour and many animals eat it (Matthews, 1952). Twins are very rare among gray seals (Collett, 1881; Curry-Lindahl, 1965) although according to some authors (Аш, Ling, and Paaver, 1957), Baltic seals do produce twins. Births in the normal course are an annual feature. The overall bar- renness has not been established accurately for any of the populations of these seals though the quantum of females not participating in repro- duction, on analogy with fur seals, was put at 20% among the British population (Hewer, 1964). The British female gray seals become capable of reproducing partly (50%) at five years of age and wholly (100%) at six years. By this time, depending on the state of ovogenesis and ovulation, the length of the ovaries increases on average by 32-35 mm and the weight goes up to 5-8 g. Thus the female births her first pup at six years of age. The males become mature at seven years of age but stake their claim at the site of reproduction only on attaining 10 years; they cease mating at 20 years of age. Among mature males, in the mating season, the weight of the testes together with the spermatic cord varies from 100 to 290 g, on average 166 g. The weight decreases by the end of the mating season roughly to 100 g; the length of the os penis is not less than 10 cm. The females remain productive for a very long time; although rare, gestating females older than 30 years (31-32 years) have been recorded. The oldest female encountered in a breeding site was 33-34 years (Hewer, 1964). Similar data are still not available for Murman gray seals. The oldest suckling Baltic seal seen in our waters was aged 21 years (V.A. Zheglov). 364 365 485 Growth and development'**. There is very little concrete information on the size of newborn pups of Baltic seals in the earlier works. The following figures are cited: 95 cm (Lilljeborg, 1874) and 60 cm (Priemel, 1909). The first figure is wholly reliable (evidently, the pup was measured along the dorsal surface) while the latter figure is clearly an undermea- surement. According to the current data (V. A. Zheglov), the body length in a Straight line (i.e., Lcv) varies from 75 to 85 cm and along the dorsal surface (Lc) from 82 to 92 cm; weight ranges from 6.1 to 9.5 kg, on average 8.15 kg. The length of the suckling pup averages 100-110 cm or 110-120 cm (Lc) and its weight 40-50 kg (the lower limit pertains to females and the upper limit to males). The body length of a newborn Murman female gray seal in a straight line (Lcv) is 107 cm, axillary girth 64 cm, thickness of the skin with the subcutaneous fat layer (51) 0.5 cm; total weight 15.2 kg (Uspenskii, 1941). A slightly older male pup, roughly three days old, was 110 cm long (Lcv) with an axillary girth of 66 cm, thickness of skin (St) 0.9 cm, and weight 20.5 kg (Fig. 193). Pups born with a body length (evidently, Lcv) of 80-107 cm add 15-25 cm in the period of suckling, the most intense growth occurring in the first two weeks. After 5-7 days the canines have cut through and the rest of the permanent teeth are cut immediately afterward (Karpovich, Kokhanov, and Tatarinkova, 1967). The newborn British seals weigh 11.5-17.5 kg but most 14-16 kg, with a body length of 91-108 cm; the body weight increases during the period of lactation to 40-50 kg (Matthews, 1952; Coulson and Hickling; 1964). The duration of lactation has been recorded by various authors as two weeks (Darling, 1948) and four weeks (Lilljeborg, 1874). Evidently suckling most often ceases in the middle of the third week (captive pups were weaned on the 18th day, and on Farne Islands, England on the 16th day) (Coulson and Hickling, 1964). During the first three days of lactation the daily weight increment averages 1.5 to 1.8 kg. This rapid increment in weight is ensured by the high fat (52% or more) and protein (11%) contents of the mother’s milk (Amoroso and Matthews, 1951). The frequency of suckling (Fig. 194) changes with time. The newborn receives its first nourishment 30 min after birth and later at 2-hr inter- vals; in the latter days this interval increases and the number of feeds decreases to a few times a day (Smith, 1962). Among the Baltic seals, the 124 The data available in the literature on the size of the newborn, its growth during the period of lactation, and in the subsequent period of its independent living, at least in the first year, as well as the sizes of adults are difficult to utilize since it is usually not known how the animals were measured: along the dorsal curvature or in a straight line to the tip of the tail, or to the end of the extended hind flippers. 364 486 Fig. 193. Newborn pup of Murman gray seal in the first few days after birth. Great Ainov Island, Murman coast, December, 1963 (photograph by V.D. Kokhanov). pups suckle initially at 3 - 4 hour intervals, receiving 400 - 650 р or more of milk in each feed. The daily weight increment varies from 1.2 to 2.5 kg but most pups double their weight in 8- 10 days and reach 33 - 40 kg or more by the end of the lactation period (V.A. Zheglov). The suckling dura- tion increases as the pup grows and the milk intake likewise increases. According to observations on Great Ainov Island (western Murman), in the second half of the lactation period suckling extends for almost 15 min. The pups suckle the two teats alternately, taking 3-4 mouthfuls at a time from each (Karpovich, Kokhanov, and Tatarinkova, 1967). The behavior of pups of the Murman population in the period of suckling has been described as follows: “The newborn pup moves little and permits fondling with the hand with no resistance. Just a few days later, it becomes active and attempts to shy away from man or protect itself, sometimes by trying to attack”. The pups usually remain within the coastal belt, up to 100 m from the line of high tide. Sometimes, however, they do stray up to 300-400 m beyond the coast. The pups spend much of the first two weeks sleeping, mostly on the right side or on the back. With the commencement of intense molting, they become more active, mainly on the rocks, often descending into the pools in the sublittoral and littoral zones. Some pups still not completely molted go into the sea (Karpovich, Kokhanov, and Tatarinkova, 1967). Completely molted pups, however, continue to remain on land for a few more days, 366 367 365 487 often moving away for long distances from the place of birth, or migrate from one island to another in the vicinity (Karpovich, Kokhanov, and Tatarinkova, 1967). The pup retains its white embryonic coat for the same short duration on the ice in the Baltic Sea as well as on the Murman islands, Norway or Great Britain. This coat loses strength in seven to ten days (Neif, 1957*; Collett, 1881; and others); among Baltic Sea newborn males and females, this happens in 5 -6 and 8 - 10 days respectively. In 17 - 20 days hair begins to appear again on the trunk (V.A. Zheglov). Among Murman gray seal juveniles too molting is over by the end of the first week. The first molt follows a fairly common pattern for all the animals of the family. Weak- ening and shedding of the embryonic hair coat usually occurs first on the head, the hind and fore flippers, and the tail (Fig. 195). On the Sth- 8th day the embryonic coat thins out on the snout and on the flippers; dark spots show through the embryonic coat here and there on the back. On the 9th- 12th day dark spots are seen all over the body, which fuse into a continuous gray field, signifying the commencement of intense molting (Fig. 196). By the 13th - 16th day the head, back, and sections around the flippers have completely molted and the embryonic coat elsewhere on the body scales off intensely. On the 17th-22nd day hair remnants are seen only on the flanks and on the abdomen but these too are weak; on Fig. 194. Posture of gray seal during suckling. Great Ainov Island, Murman coast, December, 1963 (photograph by V.D. Kokhanov). 488 the 23г4 - 24th day the remnants are rare; on the 25th- 27th day there is nothing left of the embryonic coat and the pups appear in the new coat with smooth short hair (Karpovich, Kokhanov, and Tatarinkova, 1967) (Fig. 197). Thus the entire molting process in Murman takes roughly three weeks. Little is known in general about the further growth of the juvenile. There is altogether no information on its life and growth tempo in our territorial waters. On taking to independent living in water, the pups are usually scattered widely, quite often straying very far from the place of their birth (see p. 481). The subsequent growth of the juvenile has been studied only among the British populations; by the end of the first year, the males begin to outstrip the females in growth (Table 26). The yearlings are practically no different from the well-fed pups (at the time of first molt of their embryonic coat) and are even lighter in 366 Fig. 195. Some growing gray seal pups; commencement of molt. Great Ainov Island, Murman coast, December, 1963 (photograph by V.D. Kokhanov). 367 Table 26. Average body weight (after Hewer, 1964) Age, Average body weight (approximate), kg in years Female Male 1 41 42 2 49 59 3 55 74 4 65 90 5) 72 105 366 Fig. 196. Molting gray seal pup. Great Ainov Island, Murman coast, December, 1963 (photograph by V.D. Kokhanov). 367 Fig. 197. Molted gray seal pup. Great Ainov Island, Murman coast, December (photograph by V.D. Kokhanov). weight than the latter. This phenomenon (Coulson, 1959*; Coulson and Hickling, 1964) is explained by the characteristics of the further growth conditions of a pup in the first few months of its independent existence. While still on the ice floes, after lactation ceases, the pup suffers a weight loss which continues for sometime even in the water as the youngster 368 490 has to acclimatize and become accustomed to finding its own food. This transitional period is evidently a vital factor in its life (as among other seals) and the pup is often forced to merely survive by expending the reserves accumulated during lactation. The pup can replenish its body losses only after total adaptation to the water and taking to intensive feeding. The growth of the animals, judging from the growth curve of the British seals (Hewer, 1964), continues with gradual attenuation until roughly 10 years among females and up to 15 years among males, or perhaps somewhat later. The average difference in body length between males and females after total growth cessation is roughly 35 cm. Growth is most intense among females for 6-7 years and among males for 8-9 years. Increase in skull length proceeds roughly in this same range. The known longevity in nature is 35 years for females and 25 years for males (Hewer, 1964). In zoological gardens some females have survived for 28 and males 41 years (Matheson, 1950; Mohr, 1952). Information on the molting of adult Baltic seals is very scant. At the end of the period of reproduction, after some nourishment, the Baltic seals return to ice floes and form fairly large molting rookeries on them (Holm, 1921). According to the earlier data (Neif, 1757*), this process commences by March 25; the present information (Curry-Lindahl, 1965) puts the molting of seals in April-May. There is no information at all on the molting of Murman seals. In British waters the molting period proceeds 1.5 to 2 months after the mating period; females molt much earlier (mainly at January end to early February) than males (mainly in March) (Backhouse, 1960; Smith, 1962). Enemies, diseases, parasites, mortality, and competitors. These aspects have been little studied in our waters. In the Baltic Sea the eagle may be an enemy of the pups (Aul, Ling, and Paaver, 1957) but these birds are very rare in the pelagic regions of the sea. The pups are probably threat- ened by large gulls, especially the great black-backed gull (Larus marinus L.), which pecks at the placenta and the newborn (V.A. Zheglov). Evi- dently the pups also suffer from unfavorable ice conditions. Instances of early thawing or hummocking of ice floes can pose mortal consequences for the pups which are yet weak and have not had sufficient time to accumulate the subcutaneous fat reserves requisite for independent sur- vival in water. Ravens pose some danger to the newborn on Murman (Uspenskii, 1941). Recent observations on the breeding sites of the gray seal on Murman have shown that the losses among pups are very few. On Great Ainov Island in 1963, of 93 pups, only two were found dead. Two dead pups were found in the winter of 1960/61 on Veshnyak Island 369 491 (Karpovich, Kokhanov, and Tatarinkova, 1967). In the British Isles pups born on the rocks in high-tide areas perish almost wholly; on northern Rona Island, in two weeks of October, pup mortality was 16% (Boyd, Lockie, and Hewer, 1962*). The mortality among newborns in 1957 on the Farne Islands was in the same range (England; Hickling, 1959*). In calculations of the age-sex structure of the British populations, mortality in the first year was put at 60% of the total population of pups, 30% in the second year, and 12% in the third year; it was 6.7% in each of the subsequent years (Hewer, 1964). The helminth fauna of gray seals inhabiting the USSR waters has yet to be adequately studied. Gray seals are infected by eight species of helminths, among which not a single cestode appears (Delyamure, 1955); among the trematodes, Metorchis albidus, Opistorchis tenuicollis, and Pseudamphistomum truncatum infect the gall bladder and bile ducts; among the nematodes, Anisakis similis infects the intestiae, and Contra- caecum osculatum and Terranova decipiens infect the stomach and the intestine; among the acanthocephalans, Corynosoma strumosum and С. semerme infect the intestine. The gray seal has no competitor in the Baltic Sea. This is the largest of the marine life normally inhabiting this water body and has its own ecological niche, preferentially confined to the more pelagic regions, and feeding at depth. The common seal in the southern part of the Baltic is much smaller than the gray seal. The distribution of both these species in Murman represents the boundary of the range and both evidently live under different biotopic conditions. Population dynamics. In the southern parts of the Gulf of Finland the population is insignificant and apparently is continuing to decline. The total population of the gray seal in the Baltic Sea too continues to drop with no remission, especially in the gulfs of Finland and Bothnia. Thus, the average annual catch of both species of seals in Finland compared with the pre-Revolution period, dropped in the second half of the 1920s roughly by 30% and in the early half of the 1930s by as much as 60%. The situation was similar in Swedish waters. The gray seal in the Baltic Sea is largely unprotected from foreign hunters. Encouraged by mone- tary rewards, they kill it wherever and whenever convenient. Apart from the fact that the seal is intensely hunted in general, significant qualitative changes have occurred in the structure of its population due predomi- nantly to killing of pups in the past as well as now, as they are quite easy to catch (Bergman, 1956). Hence the herd is relatively overloaded with older animals (Curry-Lindahl, 1965) whose natural death cannot be replenished. That the Baltic seal population, which showed some recov- ery after the war years, continues to fall is supported by the magnitude of 369 492 the continuously declining kill in spite of the alluring monetary rewards granted by the Swedish and Finnish governments for every animal killed (Table 27). Apprehensive of the extermination of the seals, Swedish organiza- tions for environmental conservation have turned to the government with proposals for the abolition of rewards for each killed animal, a ban on hunting during the breeding period, and the establishment of a number of sanctuaries (Curry-Lindahl, 1965). The small population of Murman gray seals underwent changes in the current century in relation to hunting activity. In the pre-Revolution period and the initial post-Revolution years, the population was evidently at maximum. In the 1930s, as a result of excessive killing of pups, the population fell but was again restored after the years of the Patriotic War. The establishment of Kandalaksh sanctuary played a positive role in sustaining and raising the population. At present, “the population of gray seals in the sanctuary has restored to the levels prevailing in the 1920s” (Karpovich, Kokhanov, and Tatarinkova, 1967). Field characteristics. The gray seal can be distinguished from other seals by the straight (upper) profile of the head, long snout, and light coloration of the upper portion of the head (Fig. 198). The last feature unmistakably distinguishes from a distance a swimming gray seal from the common seal whose head is invariably dark (Millais, 1904; Collett, 1911-1912). In the Baltic Sea the gray seal can be recognized from a dis- tance by its large size, elongated snout, and coloration, i.e., innumerable dark spots scattered haphazardly on a much lighter gray background; a gray seal protruding above the water can be recognized from the con- trasting color of the throat, neck, and chest with a pattern of larger spots seen prominently on a light background. Murman gray seals can also be identified by these features as well as by the dark, almost monochromatic color of the males. (K.Ch.) Table 27. Dynamics of catch during 1943 to 1953 (Lockley, 1954) Years Catch Average per year Maximum and minimum (number of animals) per year 1943-1945 940 1,345; 625 1946-1949 710 783; 615 1950-1952 528 707; 397 1953-1954 246 290; 203 370 370 493 Е Fig. 198. Semiadult gray seal. Northern spits in the White Sea, September, 1970 (photograph by A.G. Beloborodov). Economic Importance The gray seal commands little economic importance in our animal hunt- ing activities. Its reserves in our waters are insignificant and hence there are no special hunting prospects here. Before the Patriotic War, its kill in Murman did not exceed several dozens (mainly young animals) per year. To increase the population to a level capable of supporting profitable hunting, hunting should be banned everywhere for 10 years and then a thorough study of its ecology done, especially of its feeding habits, to determine whether this seal damages the salmon population.!” If such damage should prove negligible, the gray seal could become a regular source of small numbers of valuable fur. The other ancillary products that could be utilized are the fat (a weaned juvenile can yield about 20 kg of fat and an adult 50 kg or more) and edible meat for animal farms (10 to 12 kg from a young animal and roughly 50 kg or more from an adult seal). In the Gulf of Finland in the Leningrad region there is neither spe- cial hunting of this seal nor is it pursued as a sport. On the Estonian coasts in the prewar years, seals were caught fairly regularly but the gray 125 The available general references indicate that in Murman this seal causes no significant damage to the fishing industry. 371 371 494 seal was not а frequent catch. Outside ше USSR, in the Baltic Sea, the gray seal was caught mainly in the winter-spring in the region of the Aland Islands and the adjoining expanses in the zone of ice fringes and drifting ice floes, in the Baltic Sea itself, the gulfs of Finland and Bothnia, and in smalier numbers, On occasion, in summer and autumn at various other places. The total catch by Sweden and Finland (encouraged by monetary rewards) in the 1940s and 1950s reached an average of 1,000 or slightly more per annum (Table 28). Outside the Baltic Sea, the killing of these seals is rewarded on the Faeroe Islands (some 800 animals including 500 pups were killed in two- and-a-half years) and in Canada. Great Britain has imposed a prolonged seasonal ban during the breeding and suckling periods. Hunting in the Baltic Sea is mainly undertaken by fishing vessels cruis- ing in the period of whelping and molting of seals along the fringes and among fairly thin ice floes. Most of the catch there at this time is of the young; quite a few suckling mothers are also caught. Sweden and Finland have no rules whatsoever pertaining to sealing, nor are there any agree- ments between them regarding it. In the summer-autumn months stray seals close to the coast are killed quite often from launches and boats. Lack of supervision and totally unrestrained killing of the Baltic gray seal population has arisen because of the commonly held belief that this animal is a plunderer of fish. Without doubt, the gray seal does consume fish. But the fish consumed by it is nowhere, and never has been, of much economic importance and its rapacity is not significantly reflected in the fish reserves and hardly so in the catch. No perceptible damage caused by this seal has been recorded either in Murman or the Gulf of Finland. It has been said that it does not generally damage the fishing industry in the Baltic and North Seas (Freund, 1933) although it was earlier blamed as the most dangerous of seals in the Baltic Sea (lort and Knipovich, 1907* ). Furthermore, it has played no perceptible role as a destroyer of economic fish in some Canadian regions (Mansfield, 1966) despite an increase in its population through controls imposed along the lines of experience gained in Great Britain. Table 28. Number of gray seals caught in the Baltic Sea from 1930-1959 (Curry-Lindahl, 1965) Years Caught in Total Finland Sweden Adults Pups Total 1930 - 1939 8,995 17,265 26,260 11,952 38,212 1940 - 1949 3,800 2.715 6,515 6,792 13,307 1950-1959 3,743 2,743 6,465 3,597 10,062 372 495 Nevertheless, the presence of gray seals is undesirable in the larger economic fishing areas, especially in the estuarine sections of seas through which salmon pass into rivers to spawning sites and back. Here their adverse role on the fishing industry is obvious and controls necessitated. The damage caused by the seals to man is not restricted to the consumption of economic fish, i.e., to the adverse effect on fish reserves, but is also manifest in their consumption or partial damage of fish caught in nets, especially salmon, and the damage they do to the nets per Se. In the coastal waters of England and Scotland the damage caused by gray seals is quite significant at places. According to rough but well- argued calculations, gray seals (about 47 head) in 1957 could have con- sumed roughly 10 tons of true salmon and sea trout in the estuary of the Thames River during the entire fishing season covering 182 days. The damage extending to other fish (cod) as well as squids would account for about 3% of the total catch of salmon (the calculations assumed the following daily ration: adult seals 6.8 kg; immature seals aged two to five years 5.5 kg; and yearlings 2.3 kg) (Lockie, 1962). The total dam- age inflicted by the seals to the fishing industry on the Scottish coasts has been put at 67,000 pound sterling per annum (Rae and Shearer, 1965). Fishermen of other countries too (Sweden and the Faeroe Islands) deplore these salmon losses. Nevertheless, there is no unanimous view on this subject. The quantum of damage in some other parts of its range is insignificant (in Norway and along the coasts of the two German states) and the gray seal is a protected animal there. (K.Ch.) Subfamily of Monk Seals or 8-incisored Seals Subfamily MONACHINAE Trouessart, 1904125 These seals are large in size; their body length with tail along the dorsal surface is not less than 200 cm. The hind flippers have a fairly deep median notch. The first digit on the fore flippers is the longest and the rest gradually decrease in size toward the fifth; the claws are well developed on the fore flippers but those on the hind flippers are highly reduced and sometimes almost altogether invisible. The nostrils are mainly disposed on the dorsal side 126 The present structure of the subfamily was established by Gray (1850) long before Trouessart, but Gray named the subfamily Stenorhynchinae from Stenorhynchus, already used in the systematics of marine invertebrates. (K.Ch.) The year 1897 is perhaps more correct as this was the year in which volume I of the Catalogus was published. (V.H.) 496 of the snout. A proboscis on the anterodorsal portion of the snout is lacking. The whiskers are fairly flattened, or smooth, or with slightly wavy edges. There are two pairs of teats. The skull is essentially massive, with thick bones, and there is usually a broad gap between the orbits. The maxillaries anterior to the orbits are not swollen (not convex). The zygomatic arches project laterally to different degrees. The preorbital processes are prominent in most cases. The external opening of the bony auditory meatus is simple, without prominent lobes. The nasal aperture is less open upward and its upper posterior edge does not extend beyond the anterior margin of the orbits at the back; in most cases, in fact, the nasal aperture does not reach the orbits (genus Ommatophoca is an exception). The intermaxillary pro- cesses are usually contiguous with the nasal or almost so. There are two incisors on each side in the upper and lower jaws. The dental formula is: мы. 1 т, Ст, Pa Ms —32. The cheek teeth, except for the first premolar, have two roots. There are no sharp differences in the size and body build and partly in the coloration between males and females. In general, the body build is similar to that of seals of the subfamily Phocinae. The hair coat among newborns is dark toned. Biological affinity for a hard substratum differs: the seals of one genus (Monachus) form rookeries and reproduce on land and the rest of the genera on ice floes. The distribution of the subfamily is broken into different sections (Fig. 199). These seals are partly distributed in the Northern hemisphere: one genus (Monachus) occurs in the subtropical and tropical belt of the Atlantic and Pacific oceans and the rest in subantarctic and antarctic waters, mainly in the zone of drifting ice floes. Monachinae undoubtedly represent a product of the further devel- opment of Phocidae. They deviated from the branch of the 10-incisored seals (Phocinae), lost one each of the upper incisors, and acquired the features of high adaptation to pelagic living. This is manifest particularly in the reduction of claws on the hind flippers, in the llrge notch on the posterior margin of the latter, in the much sharper reduction in length of the digits of the wrist from the first to the fifth, etc. The evolution of the subfamily has been traced to the Miocene of western and southeastern Europe by which time the members of this phyletic branch had not only fully evolved, but differed in some respects by an even greater specialization (Pontophoca). It is highly probable that 497 ‘(uysdeyD yy) эвищовецой ‘$[еэ$ pasosioul-g jo Апшезап$ ay) JO UONNNGUISIG ‘66т ‘9 ELE SSS та ary 0056 0007 0008 0005 60% 0 0007 498 parts of the Tethys or its tributaries served as the birth place of the primary forms of the subfamily. The subfamily has not yet been conclusively systematized. Disagree- ments exist regarding its volume and constituent genera. The old view that the subfamily Monachinae includes only one genus, Monachus, is still quite prevalent (Simpson, 1945; and others); the rest of the (antarc- tic) 8-incisored seals constitute a special subfamily, Lobodontinae Hay, 1930 (= Lobodoninae Kellogg, 1922). However, it would seem more correct to adopt the structure of the subfamily to include a large volume in the form of a single group com- bining all the 8-incisored seals (Chapskii, 1961, 1969). At the same time, differentiation of Monachinae at a much lower level, at the rank of tribes Monachini and Lobodontini (Scheffer, 1948*), is entirely reasonable and rational (Chapskii, 1971*). But the spelling should be Lobodon- inae. Insofar as the composition of the genus is concerned, there is every justification (Chapskii, 1969) that it should be left alone in the same manner as adopted at the beginning of this century, 1.е., not include the genus Mirounga in spite of a recent suggestion to this effect (King, 1966). The subfamily would then contain nine genera (about 41% of all the genera of the family), of which four are extinct and five extant (50% of the contemporary genera of the family): monk seals or white- bellied seals, Monachus Flemming, and the four monotypical genera of seals of the Southern hemisphere, i.e., crab-eater seal Lobodon Gray; Weddell’s seal, Leptonychotes Gill; Ross’ seal, Ommatophoca Gray, and leopard seal, Hydrurga Gistel, 1848. The total number of species is seven, which constitutes about 39% of the total number of species of the family.'’ The species of the subfamily have no economic importance anywhere in the Northern hemisphere because of their negligible population. Until the end of the 1960s, these seals were practically untouched in antarctic waters. The population of some species there, especially of the crab- eater seal, is quite large and the idea of their commercial exploitation is gaining ground. There is only one genus within the USSR, the monk seal, Monachus Flemming, 1822 (20% of the genera of the subfamily), with a single, practically extinct species (roughly 15% of the species of the subfamily). _ Encountered in the USSR waters as a very rare, straying single ani- mal. (K.Ch.) 127 According to the system adopted in this publication, the family consists of 18 species. Si). 499 Genus of Monk Seals Genus Monachus Flemming, 1822 1822. Monachus. Flemming. Philosophy of Zoology, 11, p. 187. Phoca monachus Hermann. 1824. Pelagios'** Е. Cuvier. Mem. Mus. Hist. Nat., 11, р. 196. Phoca monachus Hermann. 1841. Pelagocyon. Gloger. Gemeinn. Naturgesch., vol. 1, no. 34, p. 163. Pelagocyon monachus = Phoca monachus Hermann. 1848. Rigoon. Gistel. Nat. Thierr. flr hoh. Schulen, Х, р. 32. Renamed Pelagios Cuv. 1854. Heliophoca. Gray. Ann. Mag. Nat. Hist., 13, p. 201. Heliophoca atlantica Gray = Phoca monachus Hermann. (V.H.) Seals of large dimensions. Body length from tip of snout to tip of tail 240-295 cm. Similar in body proportions to the species of the subfamily Monachi- nae. There are no sharp external differences between males and females. The skull is quite massive; the much older animals have fairly intensely projecting zygomatic arches. The sagittal crest is quite distinct though not high; the occipital crests are massive and sharply displaced posteriorly on both sides of the sagittal crest along the linea nuchalis terminalis. The squamose-mastoid processes are very well developed; the posterior margin of the mastoid itself projects slightly posterior to these processes when viewed upward. The nasal processes of the premaxillary bones adjoin the nasal bones. The facial portion (apex) of the nasals does not exceed one-half their total length. The anterior margin of their suture is bisected without forming a median angular projection. The petrosal bone projects into the foramen lacerum posterius. The upper incisors posteriorly have a prominent transverse groove merging into a frill. The hair coat is not high but has an extremely prominent nape; the hairs adhere very closely to the skin. The color of the upper side of the body is a dark gray, blackish- brown, turning very light on the underside. Some sex- and age-related differences exist. The skin around the nostrils and in the gap between them is covered with hair. The whiskers are smooth and oval in cross section. The neonatal coat is dense, quite high and smooth, and very dark. The claws on the fore flippers are well developed but highly reduced on 128 Various authors have spelled it differently, viz., Pelagius, Pelagus, or Pelagias. Tif 500 the hind flippers; nonetheless, their narrow and short tips are visible on the latter. The seals of this genus are thermophilic and, among all the pin- nipeds, are the most capable of surviving in subtropical and tropical conditions. They are biologically associated with the coast on which they whelp, suckle the pups, molt, and form rookeries, their size depending on the total population and the nature of the coast. They do not form harems nor do they undertake any significant or regular migrations, being predominantly confined to the coastal zone. The distribution of the genus is highly interrupted (Fig. 200). Its species are distributed in three isolated regions of the subtropics and tropics of the Northern hemisphere. One is the basin of the Mediterranean Sea (up to the Black Sea inclusive) with the adjoining sections of the eastern Atlantic, in the southwest from Gibralter to the Canary Islands and the coasts of northwestern Africa. Another is the now practically uninhabited westernmost part of the Atlantic: the Gulf of Mexico and the Caribbean Sea. The third region of distribution is even more isolated, lying in the Pacific Ocean and covering only the archipelago of the Hawaiian Islands. This genus has evolved through intermediate links from Miocene ancestors found in western and southeastern Europe, including the Sar- matsk formations in USSR territory. A likely direct ancestor of the genus Monachus could be some Lower Pliocene form of the genus Pristiphoca since a member of the recent genus Monachus adjoined the above fossil genus even in the Middle Pliocene. The geological history of this genus binds it to the west of the Old World, especially to the western, south- ern, and southeastern parts of contemporary Europe, to the basin of the Tertiary Tethys Sea. Paleontological proof and morphological data of the contemporary members of the genus Monachus indicate that this genus represents a phyletic base from which other contemporary genetic branches of the subfamily originated and evolved. The genus Monachus, more than any other genus of Monachinae, reveals many features relating it to the sub- family Phocinae (especially the petrosal bone projecting into the foramen lacerum, margin of mastoid visible when seen upward, structure of the zygomatic bones, etc.). It may be regarded as a primary element of the eight-incisored phyletic branch originating from the primary Phocinae and serving in turn as a stage for the evolution of its derivative, i.e., the primary ancestral form of subfamily Cystophorinae. The genus consists of three species, or about 43% of the species of the subfamily: (1) Mediterranean monk seal, M. monachus Hermann, ‘IPUDISUINDYIS “PFW чепемеН— SNYIDUOLU W uesuelss}Ipsf[—Z ‘syooidoa "И чеэаанеэ—1 ‘(irysdeyo МУ) 5пузриои ‘seas Уцош jo snuas 94} JO UOTNQIISIP PsJONISUODIY ‘005 “B14 9/5 aaa 03000$ 020+ OCDE 0505 GO O 0501 502 1779; (2) Caribbean monk seal, М. tropicalis Gray, 1850; and (3) Hawai- ian monk seal, M. schauinslandi Matschie, 1905. Though all of these are allopatric, the species characteristics are independent. They differ mor- phologically in the structure and position of the teeth in the row, in the form of nasal and zygomatic bones, and in other respects. Thus there are two Atlantic and one Pacific species. At present, practically speaking, there are two species (both the species are protected in most places); the West Indian species is apparently wholly extinct. The fauna of the USSR contains one species, M. monachus Her- mann, 1779, or 33% of the species of the genus and roughly 0.3% of the species in the fauna of the USSR. This species was almost extinct in the territory of the USSR even by the second half of the last century. (K.Ch.) MONK SEAL Monachus monachus (Hermann, 1779) 1779. Phoca monachus. Hermann. Beschaf. Berlin. Ges. Naturf. Freunde, 4, p. 501, pl. 12, 13. Lake and coast of Dalmatia; Adriatic Sea. 1785. Phoca albiventer. Boddaert. Elench. Anim. 1, p. 170. Adriatic Sea. 1800. Phoca bicolor. Shaw. General Zoology, 1, p. 254. Adriatic Sea. 1816. Phoca leucogaster. Peron and Lesuer. Voyage aux terres Austr. 2, р. 47. Nimes, southern France, Mediterranean Sea. 1828. Phoca hermanni. Lesson. Dict. Class. d’Hist. Nat., 13, p. 416. Adri- atic Sea. 1838. Monachus mediterraneus. Nilsson. К. Svens. Vet. Ak. Handl., 1837, p. 238. Adriatic Islands and Greek archipelago. 1848. Phoca crinita. Menis. П Mare Adriatica, р. 153. (V.H.) Diagnosis The dimensions are large, evidently larger than many other species of the genus. The color is darker. The nasal bones are relatively short and form 16-20% of the condylobasal length. Their anterior margin has a median notch. The upper incisors are disposed in an arc; there is no more than one additional cusp on the premolars posterior to the main cusp. The preorbital processes are well developed. The uncinate processes of the pterygoid bones are not bent angularly outward. (K.Ch.) Description In general build, the monk seal does not differ greatly from other seals 378 Of similar size (Fig. 201) but is evidently slightly stockier than many 503 ZZ 2..2) — EGA LEE РА РНР ее а. SY LEE = eas ГАР а Sate, ЗЕЕ ZS he, ИР 2277 LLL =i) — a OIA, 2 , ПОР: о —— 4 378 Fig. 201. Monk seal, Monachus monachus (figure by М. М. Kondakov). species of the true seals (Phocinae) with a heavier and apparently slightly shortened anterior portion of the body.!”” The head is flattened on top, as for example among the harp seals (Mohr, 1952). The fore flippers are somewhat shorter than the hind ones. The claw zone on the fore flippers appears as though incised in a strait line while the length of the claws decreases quite uniformly from the first to the fifth digit. The hind flippers have broad terminal lobes due to an angular folding along the inner edge turned toward the apex of the median notch, which is deeper than in the true (10-incisored) seals. The claws on them are highly reduced, very narrow, and barely visible (their length does not exceed 0.5 cm). The claws on the fore flippers are not pointed but broad. The hair coat of adults is rigid, smooth, adheres closely to the body, and very low. The hairs are very thin; their length on much of the body is about 0.5 cm (King, 1959*). On the dorsal side of the body of females from Kilii delta of Dunai River, the longest hairs (5.3 mm) were found on the nape, their length on the back and in the tail section being 3.3 and 2.5 mm respectively. The hairs are longer on the belly: the longest of them are in the midportion of the trunk (10.9 mm) and slightly shorter 129 This impression is gained from some published photographs (Mohr, 1952; King, 1956; Heck, 1966). It is highly possible that this is due to the jugular section of the male actually appearing in the photographs as massive and comparatively short while the head appears large with a slightly swollen snout. Confirmation of this is the wide spacing of the zygomatic bones in the skull of large males exceeding that in females (see p. 506). Quite possibly, sexual dimorphism does exist among monk seals in the size of the body and the dimensions of the skull, although no direct evidence is available (see p. 506). However, the suggestion that the head of this species is unusually large and resembles that of a bear (Sal’nikov, 1967) is evidently due to a misunderstanding and is explained as due to the poor preparation of a stuffed animal. 379 504 on the fore portion (10.5 mm). The hairs are the shortest in the tail section (3.1 mm). The thickness of the hairs is identical in all sections of the body (0.1 mm) (Sal’nikov, 1959).130 The neonatal pelage is at least three times longer, incomparably denser, and altogether without nape. It consists of soft, thin hairs. The labial whiskers are set in five or six rows; their number in a row on each side of the lip decreases gradually from eight to two as the number of the row increases. There is one more lone whisker on top. The total number of whiskers goes up to 70. Their length varies in relation to the position of the row as also the height of the row and probably depending on age and even season (assuming that the whiskers are sloughed during molt). Among the young (less than two years of age) their length varies from 2 to 15 cm. They are oval in cross section, without wavy edges and highly tapered toward the apex. There are hairs like whiskers even in the chin region, their length not exceeding 2 to 3 ст.131 Supraorbital whiskers do exist, contrary to the opinion of some authors, even among the young animals (not less than three). The color!*? of the hair coat of the adult animals is dull and quite often monochromatic. The characteristic predominant shades are gray, brown, and black, as also contaminated tones; the dark color is intensi- fied on the dorsal side of the body compared with the ventral. There is something like a color bloom, creating a faint, bright white or yellowish hue, due to the alternation of light-colored tips of the hairs with compact dark hairs.!* The upper side of the body of adult males is usually a vivid dark gray or slate-black with an admixture of fairly distinct brown shades. On the body flanks, the color turns pale and imperceptibly transits into an even lighter but still vivid gray on the lower side of the body. In this 130 According to some other sources (Schapp, Helwing, and Chizelea, 1962), the length of the hairs among the much younger animals varies from 4 to 7 mm on the ventral side of the body. 131 The description was based on the data of King (1956), Sal’nikov (1959), and Schapp, Helwing, and Chizelea (1962). 132 Description of the color features of the hair coat is very difficult because of the extremely few specimens. The number of skins in our museums is very small and such altogether absent for the other species of the genus. Further, due to prolonged storage, the skins have acquired a fairly distinct yellowish-golden hue. 133 There is one more widely prevalent type of coloration in the form of fine light-colored bands or variegations, quite abundantly dispersed in large patches at different places on the body. These are concentrated most often under the chin, along the flanks, and at the center of the neck, and at various places on the chest and back. Since these variegated spots are visible (seen through) on the reverse side of the skin (on the flesh side), they may be regarded as caused by abrasions. 380 505 background, а very large spot stands out in contrast. This spot is tens of centimeters across, angular, almost white or slightly creamy, rhom- boid or roughly rectangular, and often resembles a butterfly. This is a characteristic feature of an adult male. The spot is usually disposed asym- metrically, most often on the abdomen or on the flanks in the rear half of the body but closer to the fore flippers. Among adult females (King, 1959*; Sal’nikov, 1959), the dorsal side is dark, blackish or dark gray, with a slightly silvery or yellowish bloom due to the light-colored tips of the dark brown hairs. On the flanks, the hair coat gradually turns lighter and becomes light gray on the entire underside of the body, but usually without the large whitish spot characteristic of the males. The yellow bloom often seen on the skins is most likely due to posthumous color changes caused by oxidation of fat remnants present in the skin. The color of the tips of hairs among live animals is mostly whitish or very pale gray, if not white. The pelage of the newborn is a fairly uniform dark brown or dark cin- namon on the dorsal surface; it turns lighter on the flanks and apparently is even lighter in color on the ventral side. Evidently, depending on whether the animal is a male or female, on the underside of the body or on the flanks, posterior to the fore flippers, there is, or is not, a large angular spot of white or almost white color similar to that described in the case of adult males. As the neonatal (juvenile) coat is shed, a rigid, very short hair coat closely adhering to the skin is seen; this coat is an altogether different color and is characterized by sharp dichromatism: dark gray on the dorsal side and very light or almost pure white on the ventral side (or ventral side with yellowish tinge). It may be sharply set off on the flanks, roughly at the level of the base of the fore flippers from the dark gray with brown bloom of the dorsal side (Schapp, Helwing, and Chizelea, 1962). Apparently, at this early age, the color of the first definitive hair coat is identical among males and females. The color of the males of transitional age is highly similar to that of adult animals of the same sex. The former, however, lack the large light-colored spots of the type described above which are characteristic of the adult (and possibly the newborn) males (King, 1959*; Kumerlowe, 1966* ). The skull is somewhat larger than in the other species of the genus (Fig. 202). Its upper contour when seen in profile usually descends quite steeply toward the fore end roughly from the apex of the nasal bones. The narrow and long zygomatic bones have an anteroventral angular margin distinctly restricted outwardly along the suture. Preorbital pro- cesses are distinctly manifest in the form of a nipple. The nasal bones 380 381 506 vise oy о и ий y ИИ MH SN А о М = Fig. 202. Skull of monk seal, Monachus monachus (figure Бу М.М. Kondakov). are anteriorly bifurcated by a median angular notch; the anterior end of each of them is in the form of an angular or tonguelike prominence. The frontal-maxillary suture from the nasal bone to the flexure in the orbit is not shorter than the nasal-maxillary suture and is usually longer. The bony tympanic bullae are roughly triangular, of irregular form, highly elongated forward, lean somewhat toward each other, and terminate con- siderably anterior to the crest of the articular fossa. Their internal margin is concave with the aboral carotid fossa at their very end. The fontanelles in the choanal region are very large. The posterior margin of the bony palate is in the form of an arc with a notch, often asymmetrical. The condylobasal length of the skull of adult males is 280-302 mm, of adult females 260-280 mm. The zygomatic width in adult males is 190-215 mm, in adult females 160-180 mm. The teeth are very massive. All the lower premolars and at least two upper ones are set aslant relative to the tooth row. Their crowns are formed mainly with a single massive main cusp at the base of which is a broad fringe (cingulum) from inside. Anterior or posterior to the base of the main cusp or on both sides of it, quite often an additional denticle, barely raised above the cingulum, appears. All the teeth, except the first premolar, have two roots. The upper incisors are disposed arcuately: the 507 median less large ones are shifted forward slightly compared with the extreme ones. A semicircular notch and fringe in the form of a cingulum is seen On the posterior side of all the upper incisors. The body length of adult males from tip of nose to tip of tail varies from 210-250 cm; some very large animals could be of much larger proportions. Nevertheless, the figure of 300 cm and above cited in the literature (М. Smirnov, 1929; and others) is hardly reliable.!*4 The adult females are probably somewhat smaller although a ges- tating female caught on the coasts of Corsica measured 278 cm (evidently up to the tip of the flippers) (Troitzky, 1953). The length of a female from the Kilii River bed of Dunai (if not sexually mature, quite close to being so) measured 227 cm (Sal’nikov, 1959). The total body weight of adult animals goes up to 300 kg and may even exceed it. (K.Ch.) Taxonomy The monk seal although undoubtedly representing the ancestor from which other species of the genus have evolved, is more specialized in some respects than its descendants. This is manifest in the intensification of the dental apparatus (premolars and molars more massive with a more sharply manifest tendency to a transverse setting, i.e., to the displacement of their anterior edge inward), in the intensification of the maxillary musculature manifest in a broadening of the coronoid process and the lower posterior subcondylar portion of the lower jaw. The enlargement of the zygomatic arches is more significant than among the other species; the transformation of the bony tympanic bullae has proceeded farther, more forward advanced, etc. The phyletic relations of the monk seal with the other species, in a morphological context agree with the probable direction of the dispersal of the species of the genus: the Caribbean seal is closest to the European seal in the structural features of the skull while the Hawaiian seal differs most. The two former species are related particularly in the steep curva- ture of the upper line of the profile and generally the greater height of the skull, the greater mass and tapering disposition of the cheek teeth, enlarged subcondylar portion of the lower jaw, structure of the anterior region, general shortening of the zygomatic bones, and structural fea- tures of the bony tympanic bullae. On the other hand, the Caribbean seal can be considered closer to the Hawaiian seal in the proportions of the skull (low index of its width) and in some plastic features (length of 134 Тре body length of adult males as established from the published literature, makes no claim on adequate accuracy since not a single author has specified the method of measurement. 382 508 the nasal bones and their contact with the premaxillaries, structure of the coronoid process, pterygoid bones, contours of the suborbital apertures, etc.). Some craniological features among the Caribbean seals are of an intermediate character (reduction of the preorbital processes, form of the anterior margin of the nasal bones, length of the zygomatic bones, etc.) (King, 1959*).135 The following general features can be cited: 1. The Mediterranean monk seal, M. monachus, represents a special species. Some structural features which are of a more primary (primitive) nature are characteristic of it (arcuate setting of the upper incisors, upper contour of skull profile steeply dipping forward, presence of sharp pre- orbital processes, etc.), proximating it to the 10-incisored seals (angular form of the lower anterior margin of the zygomatic bones). 2. Of the other two species, the Caribbean seal, M. tropicalis, is more related to it than the other species, while possessing several features bringing it closer to the Hawaiian М. schauinslandi. 3. М. schauinslandi combines in itself craniological features of a more primitive character (bony tympanic bullae and some swelling of the pre- orbital zone of the rostrum) as also features that are more evolved (long, arcuate zygomatic bones that are concave anteriorly and smoother upper line of the profile). (K.Ch.) Geographic Distribution Coastal waters of the Mediterranean Sea along the continental coasts of Europe and Africa and also the coastal islands, including those away from the mainland. Parts of the Atlantic Ocean southwest of Gibraltar up to the Madeira and Canary islands inclusive, mainland coasts of Africa in the south up to 20 to 15° М lat. (Cape Cabo Blanco in the Senegal River estuary), western, Balkan, and southern Asia Minor coastal zones of the Black Sea (Fig. 203). Geographic Range in the USSR Forms a negligibly small portion of the northeastern rim of the range. The reconstructed boundaries of distribution in the last century or earlier covered the western strip of the Black Sea adjoining Romania within the southern part of the coast of former Bessarabia with the region of Zmeinyi Island (Krotov, 1952), the sea along the western coast of 135 When evaluating these features, it should be remembered that the data for all the species are extremely scant and the extent of individual variation of several features is not known. All these could be a source of error. 382 383 509 Fig. 203. Reconstructed distribution of the monk seal, Monachus monachus т the USSR (К.К. Chapskii). Crimea, possibly both sides of the Tarkhankut Peninsula (Zernov, 1913), and farther south and east of the southern coast of Crimea from Cape Kherson roughly to Gurzuf (Nordman, 1840; Керреп, 1883*; Puzanov, 1929). Some small tongues of distribution intrude from the side of Asia Minor waters into the extreme southeastern part of the Black Sea toward Batumi (Sal’nikov, 1959). In the early 1840s, these seals were encoun- tered quite often on Zmeinyi Island (Nordman, 1840; Puzanov, 1969*). Reports on the habitats of monk seals in the region of Sevastopol go back to ancient times although they indicate that seal sightings were generally rare. At the end of the eighteenth century, it was likewise reported that seals were “rarely seen on the coasts including Sevastopol harbor, unlike at other places where they are often sighted” (Gablits!’, 1785*). 38 136 Interestingly, Pallas (1811*), who lived for quite sometime in Crimea at the beginning of the last century, reported that seals in the Black Sea came only from the Mediterranean contd... 510 The presence of these seals on the southern coasts of Crimea in the first half of the nineteenth century was more accurately reported by Nordman (1840) who indicated sightings in the caves on the rocky south- western coast. The seals were few, however, and occasionally became tar- gets for hunters. It is interesting that all the recent published reports on this animal are usually a repetition of the earlier publications. Reports of original finds of these seals are extremely rare. Thus, in the 1830s, one seal was killed between Kuchuk-Lambat and Karabakh (Keppen, 1883*) and, in 1834, a very large seal was “sighted on a cape at the end of the land in a botanical garden” (A. Nikol’skii, 1891). At present, the monk seal lives or reproduces nowhere in the western part of the Black Sea. The appearance of stray animals in the extreme northern portions of the Dunai delta represent extremely rare instances of transgression far beyond the limits of the habitats that are normal for our time. Such instances were noted at the end of the 1930s and early 1940s at some points in the Kilii delta: on Siberian spits, in the narrow straits of Prorva where seals were caught in fishing nets, and also at several other places in the water as well as on sand spits, including the small islets Limba and Kuril (Sal’nikov, 1959). From 1946 through 1951, there were five more instances of seals being caught by fishing hooks (Krotov, 1952), including a female on May 20, 1950. Zmeinyi Island too falls in the same coastal section of the northwestern Black Sea but straying of the monk seal onto this island is as rare an event as in the Dunai delta. There is no positive information whatsoever on finds of this seal farther north along the western coast of the Black Sea barring a reference to an “animal like that of a seal” noticed 30 km east of Odessa in 1950 (Kleinenberg, 1956). There is no positive information on sightings of the monk seal on the Crimean coasts in this century and it may be regarded as totally extinct there (Puzanov, 1929). In Tarkhankut-Bokkal region (a unique place where it was still considered possible to sight an occasional seal early in the nineteenth century and early in this century), the seals were obviously more numerous in the remote past (Zernov, 1913). There is no truth, however, that seal populations lived and reproduced on the Tarkhankut Peninsula in the last century. Significantly, there are no other references in the literature of that time to the existence of this seal. Yet, there are persistent reports of seal sightings in the region of Medvezhii caves near Sevastopol beyond the Kherson lighthouse, where these seals were even hunted at the end of the last century (Zernov, 1913). (“т Pontum Euxinum adscendit em Mediterraneo’”’). However, he was simply repeating the widespread belief of the local inhabitants. 384 511 There are three references in the literature pertaining to the south- ern coast of the Caucasus but they do not inspire confidence. One states that on the coasts of Adler region a whole herd of animals “with promi- nent snout and whiskers” was seen once (Poznanskii, 1880); another con- cerns Batumi region near which some fishermen killed a seal in water but could not get it out because it drowned (Kleinenberg, 1956). In the 1920s, near the rocks in the region of Cape Zelenyi (close to Batumi), a few seals settled for several years. Over the course of time their number dwindled and the last of the animals was killed in 1933 by a dolphin hunting ship cruising in Chernaya River estuary (Sal’nikov, 1959). The greater inhabitation of the coasts and the absence of appropriate biotopic conditions are adequate reasons for disbelieving the appearance of monk seals on the southern fringes of the Caucasian coast. Geographic Range outside the USSR In the past this covered almost the entire coastal strip of the Black Sea along the Balkan Peninsula (from Dunai to Bosfor) and Asia Minor. In the Mediterranean Sea basin the distribution encompassed almost the whole sea coast and the islands except for the long inhabited sections on _ the southern coast of Europe (most of the French and Italian coasts and many parts of Spain and Greece) and also some sections on the eastern coast and African coast. Outside the Mediterranean Sea, the distribution covered a small section of the Atlantic Ocean along northwestern Africa, in the south up to Cabo Blanco and even the estuary of the Senegal River (15° N lat.) and also included the Madeira and Canary islands. Even at the beginning of the present century, much of the coastal waters surrounding the southern coasts of Europe fell outside the distri- bution while the expanses interrupting the distribution along the African and Asia Minor coasts enlarged. In the 1960s, the distribution com- prised a few isolated sites of essentially disconnected diminishing pop- ulations. Some are protected by the government, especially in Bulgaria, Yugoslavia, and Rio de Oro. Only stray wandering animals have been sighted, and extremely rarely, on the Romanian coasts. In the first 50 years of this century, only nine cases of monk seals caught in fishing nets in Romanian territorial waters were recorded and the last such catch (in July 1960) after an interval of 12 years (Schapp, Helwing, and Chizelea, 1962). In Bulgaria there are two zones of reproduction, both with extremely small populations, namely Cape Kaliakra and south of Burgas. The rest of the Black Sea distribution encompasses the coastal belt of Turkey (Ana- tolia). The boundaries and the nature of distribution of the species there 385 512 are not clearly known but the population can be regarded as extremely small and widely scattered, mainly in the western regions. Outside the Black Sea, there are a few places where some small reproducing colonies still exist and also some regions where these seals are encountered but their population and reproduction have not been established despite their presence being known for 20 to 30 years. Repro- ducing populations of the Mediterranean Sea are encountered on some islands in the Adriatic Sea along the Dalmatian coast of Yugoslavia, the Lebanon coast, and possibly here and there the Turkish coast; the existence of one or some small colonies in Greek waters and on the Ionian Islands is quite probable; their presence has been established with certitude on the small islands near Tunisia, in the waters of Morocco, southern Sardinia, and Corsica. In addition to the aforementioned sites, some Stray seals are sighted from time to time in the Sea of Marmara, in the Dardanelles, along the eastern fringe of the Aegean Sea, in Salonik Bay, on the Rodhos, Kipr, and Kriti islands, in the northeastern corner of the Mediterranean Sea, and on the coasts of Syria and Israel; the presence of monk seals is also probable on the Egyptian and Algerian coasts, on the Baleares, and at other places. These seals have been reported southwest of Gibraltar in the oceanic part of their distribution in the region of Medeira (Desert Island) and especially in the continental coastal sections of northeastern Africa to the south of Rio de Oro (in the region of Cabo Blanco). The southern- most point on the coast of northwestern Africa for which information is available about the presence of this seal is the estuary of Senegal River (about 15° N lat.). In general, however, the southern boundary of distri- bution on the Atlantic coast of Africa roughly corresponds to the 20° winter isotherm of water. (K.Ch.) Geographic Variation No information whatsoever is available on the morphological features of the local populations. (K.Ch.) Biology!*’ Population. Right now, it is not appropriate to refer even to the very existence of the monk seal, let alone its population in our ferritorial waters. In the remote past, however (a century or sO ago), as already 137 Information on biological aspects is based mainly on foreign sources which, however, are neither abundant nor thorough. 386 513 stated before (р. 510), the seals were regarded as quite common оп Zmeinyi Island. They were evidently reported in very small numbers on the southwestern coast of Crimea including Sevastopol region although, Over a century ago, it was stated that seals in the Black Sea were “few and only rarely stray animals reach the northern coast” (Kessler, 1861). The population is not much on the coasts of Bulgaria and Anatolia of Turkey. In spite of a total ban on hunting and strict conservation for many decades, the population on the Bulgarian coasts has shown no upward trend. Just 20-30 animals have been reported in the region of Cape Kaliakra (Caspers, 1950). The population of the colony south of Burgas does not even exceed half this number. The Bulgarian population is essentially the only source for the dispersal of this seal in the northern part of the western Black Sea including Zmeinyi Island and Crimea. The rarity of the seal on the Romanian coasts has already been commented upon. It is not possible to draw a picture of the present population of the monk seal on the Turkish coasts of the Black Sea. Some sixty years ago, 3 or 4 seals were encountered on the Anatolian coasts during a 3-week cruise. This gave grounds for the statement that the sighting of this seal on the Anatolian coasts “was not so rare” (Zernov, 1913). Somewhat later, and also during a three-week cruise, “10 to 12 families” were counted on the west coast of Anatolia, at Zonguldak (Caspers, 1950; Mohr, 1952). There is almost no information from Turkey for the 1960s and that available does not provide a complete picture (Mirsaloglu, 1964 from Kiimerlowe, 1966). It is only known that the seals are sighted every year not only on the Mediterranean coasts, but also on the Black Sea coasts and that small colonies probably do exist on the Anatolian coasts as well (Wijngaarden, 1966* ). Since, however, no seal conservation measures have been practiced in Turkey in the last half a century, its population has been considerably depleted and at present is hardly more than the Bulgarian population. Had the situation been different, solitary animals would have been sighted in our waters on the Caucasian coast in the last 30-40 years. The number of Mediterranean populations, not exceeding 20, is extremely small. Their existence is reliable only at three places: 1) island on the Dalmatian coast of Yugoslavia, 2) northern coast of Corsica, and 3) Tunisian islet Galata. Moreover, two populations of indeterminate strength are encountered on the Ionian Islands and south of the Aegean Sea. The existence of colonies on Crete, on the southern coast of Turkey, and also on the coasts of Libya east of Benghazi is doubtful. The largest population of monk seal is found on the Atlantic coast of Africa, south of Rio de Oro, directly around Cabo Blanco, where some 200 animals (or 514 even more) have been counted (Kusto and Dyuman, 1953; Wijngaarden, 1966* ). The total number of all the populations of monk seal equals less than a thousand. This number is so small that the species is threatened with extermination. Habitat. Being an aegialoid [littoral] animal, 1.е., biologically associ- ated with coasts, the monk seal is confined predominantly to the coastal zone and is not found deep at sea. Since, however, the coasts within the distribution zone have been extensively inhabited for a long time now, the seals have taken to vacant or less populated sites for breeding. These are mainly rocky islands and generally those sections of the mainland coast which are not inhabited by man. Thus in Crimea the rockier and more rugged sections of the southwestern coast are regarded as sites of their habitation. The seals rarely take to open and low beaches. At least the seal colonies known at present are not associated with such biotopes. It is possible, however, that in the past they did live on the vacant and open beaches of Zmeinyi Island but these were probably only temporary rookeries not associated with breeding. The shallowness of the coasts and their low level explain the absence of permanent rookeries on the vacant and low coasts in the narrow straits of Dunai (Sal’nikov, 1959). At various sites outside our waters, the seals occupy predominantly the uninhabited and less inhabited islands, more rarely the mainland coasts with complex, highly rugged relief, abounding in rock clusters and various types of rocky recesses. They generally select caves and grot- toes, especially those with an underwater inlet. In such rock crevices and caves, protected from the direct action of the surf, the animals eagerly take to sandy-pebbly beaches. Similar biotopes with caves and grottoes abound in the Black Sea (on the Bulgarian coasts at Cape Kaliakra and on the Turkish coasts of Anatolia), in the Mediterranean Sea, and also on the ocean coast of northwestern Africa (including Cabo Blanco). In this context, the explanation for the absence of this seal south of Cabo Blanco, i.e., the coast there is low and sandy, is rather intriguing (since not wholly correct) (Bétger, 1951*). Food. Almost nothing is known about the food of these seals in the Black Sea waters of our country. The stomach of females caught in Kilii, the narrow strait of Dunai, revealed about 3 kg of semidigested remnants of flounder (Rhombus maeoticus) (Sal’nikov, 1959). The catch of seals coincided with the en masse approach of this fish to the coasts for spawning. Judging from the massive teeth of the monk seal, one may assume that it specializes in consuming large fish. This has been confirmed by seal food preferences in zoological gardens and the stom- ach contents of some specimens caught in the Mediterranean Sea. Thus, 387 SS) on the coasts of Sardinia the stomach of these seals revealed the rem- nants of wrasses (Labrus) and sparid fish (Dentex) (Carrucio, 1893; King, 1959*). The seals were noticed playing with large fish measuring about 70 cm in length. They caught them in their mouth after throwing them up in the air, with the head of the fish turned directly into the mouth (Caspers, 1950). It is therefore hardly correct to suppose the food of the monk seal to be large fish, especially such massive and migratory fish as anchovy and sprat; given the sluggishness of the monk seal, catch- ing such large fish would be difficult (Sal’nikov, 1959). On the African coasts, in addition to fish, the seals consume thorny lobsters (Palinurus). According to fishermen in the region of Cape Kaliakra, a large seal can consume 15-20 kg, with a preference for sturgeons (Caspers, 1950), but this estimate is doubtful. The daily ration of the seals in zoological gar- dens reached 12 kg (Gavar, 1927*) and they consumed all types of fish, such as whiting, eel, sardine, salmon, etc. Home range. Under minimal favorable conditions, monk seals are usually confined year round to the same section of the coast in which they breed year after year, i.e., the seals lead a settled mode of life. The dimen- sions of the selected sections evidently vary depending on such factors as the size of the population, availability of fairly suitable hideouts, food conditions, etc. The actual population regime on Cape Kaliakra con- forms to these factors. The boundaries of the occupied coastal sections could only be tentatively established for each of the given populations. Daily activity and behavior. Almost no observations on the behavioral features of the animals transgressing into our waters have been reported. Data on the behavior of the monk seal outside our country are likewise scant. The seals are regularly encountered outside their hideouts in the day (Caspers, 1950; and others) but may be found sleeping even on the coast in grottoes when the brightness is not sharp (Troitzky, 1953). Contrarily, they exhibit the same mobility even in darkness (Kiimerlowe, 1966). It may be assumed that there is no strict sequence of sleep (night) and activity (day). They can be regarded as more diurnal than nocturnal ani- mals. In this respect, observations on the behavior of these seals on Cape Kaliakra are of interest. With surprising stereotyped regularity, noticed for hours on August 23, 1941, the seals swimming from the southern side dove roughly 20 m from a rock. Later, they remained submerged for brief intervals, obviously in shallow water, often surfacing in between, at times with the head ducked and only the back showing above the water. It appeared as though the animals were searching the bottom for something. This probably was so, considering that the water around Cape 388 516 Kaliakra is exceptionally transparent. Leisurely swimming thus, submerg- ing and surfacing, the animals turned away for some distance from the rocks. They continued the reverse journey in the same manner until they were lost to sight. Nearly eight minutes later they again appeared and repeated the selfsame behavior. Before submerging, the animal raised its body quite high above the water and flexed its forepart so as to assume the required direction at depth. While diving, air was often exhaled even inside the water, which bubbled up in the form of an arc until the head of the animal surfaced. From this arc one could judge beforehand the point of surfacing. On surfacing, the hissing sound of the animal’s expiration was Often audible (Caspers, 1950). Monk seals are generally good divers; they usually throw themselves into water from rocks and are somewhat like sea lions in this respect. They first assume an appropriate posture by heaving the chest. It has been assumed that monk seals are not particularly good swim- mers (Sal’nikov, 1953). Some observations on their swimming capabilities from the coast and from ships (Caspers, 1950) do not justify changing this view. Nonetheless, a slowly swimming animal could hardly seize a large fish. Divers with aqualungs could not approach adult monk seals to photograph them (Kumerlowe, 1966). The relations between individual animals are hardly known but Bulgarian fishermen were witness to scuffles which, in their view, arose when a new animal entered the colony. The reports of fishermen on regular and severe scuffles between adult females sound somewhat strange. Scuffles are accompanied by loud laryngeal cries resembling at times the bellowing of calves, barking, and even the scream of a beaten or bitten dog (Wolf, 1818*; Carrucio, 1893; Caspers, 1950; Morales- Agesino, 1950* ). The voice of young seals is generally similar to that of an adult but fainter and less “fervent” (Morales-Agesino, 1950*) while some have a more characteristic voice. A very young specimen caught close to Livorno, uttered the sound “ovaavaavava” in captivity, sometimes something like a sneeze was heard interspersed with an intense dull sound, and at times the animal emitted a bleat or simply a loud scream (Wolf, 1818; after Mohr, 1952). A newborn seal caught near Split gave out laryngeal sounds similar to “oa” (Priemel and Mohr, 1952*). Observations of the behavior of captive animals have revealed well- developed higher nervous activity. This is manifest in comprehension and great affinity to man, as noted in its ability to respond to his com- mands to some extent. A very young animal from around Livorno became totally domesticated after a few months and would “kiss” its master while producing a sound similar to belching. On being brought to Nurnberg, it became totally silent (Wolf, 1818; after Mohr, 1952). Another pup, 389 S17 which had learned to recognize its trainer, would cry out on sighting her (Ber, 1838*). Seasonal migrations and transgressions. Being generally confined to a single section of the habitat, monk seals do not undertake periodic or distant migrations. At the same time, the transgressions of stray sin- gle animals to fairly long distances from the breeding site are widely known although not frequent. Such animals were sighted in almost all parts of the range and even at its very boundaries. The reasons for these transgressions are not known. It is difficult to say for certain whether the seasonal arrival of fish for spawning plays a significant role in this respect in certain regions. Nevertheless, this phenomenon in particular is used to explain the appearance of some animals and their being trapped in fishing nets in Dunai delta (Sal’nikov, 1959). However, transgressions here are extremely rare and reveal no regularity whatsoever. Were they regular migrations, in spite of the extremely small population, the migrating ani- mals would first have been encountered in somewhat larger numbers and more often, and secondly their departure from Kaliakri would have been perceivable. But, far beyond the boundaries of the colony, on Cape Kali- akri, single animals and almost invariably young ones are encountered (Calinescu, 1936). These phenomena can more naturally be explained as cases of transgression. Reproduction. Very little is known. The period of mating follows the cessation of lactation although it can commence even somewhat earlier and is evidently protracted. Its calendar periods vary for different animals. Most animals evidently mate in the autumn or at the end of summer. The total duration of the embryonic period (from the moment of mating of the female to parturition) has been put at 10 (Caspers, 1950) or 11 (Troitzky, 1953) months. The whelping period falls at the end of summer- autumn. It is July-August for the Bulgarian populations (Caspers, 1950). Much earlier periods have been reported for the Anatolian coast pop- ulations of Turkey: a female caught near Zolgundak delivered a pup in a zoological garden at Ankara on May 5, 1962 (Mursaloglu, 1964 from Kumerlowe, 1966). According to other indirect data (Zernov, 1913), the period of whelping on the coasts of Anatolia is roughly in the second half/end of July. In the Mediterranean Sea the whelping period extends from mid-August through September and even October (Carrucio, 1893; Dathe, 1934; Troitzky, 1953) and on Rio de Oro, even later, possibly around mid-November (Agasino, 1950 from King, 1956). The reference to spring whelping (Postel, 1950*) is evidently not wholly correct. It is impossible to establish the dimensions of the newborn because of the absence of information on the method of measurement and on age. Pups with a fresh umbilical cord, found in Dalmatia on September 518 19, 1933, measured roughly 90 cm and weighed 26 kg (Рае, 1934). A fully mature fetus was removed from the womb of a female killed on September 27, 1947 on the coast of Corsica; it measured 120 cm in length and weighed 17 kg (Troitzky, 1953). Lactation extends for 6-7 or 7-8 weeks (Caspers, 1950; Troitzky, 1953). The total duration of the reproductive cycle according to some authors (Caspers, 1950) is 12 months and according to others (Troitzky, 1953), about 13 months and hence births are not an annual phenomenon. It is difficult to say which view is more correct but nevertheless it may be assumed that births are an annual feature in most cases. Growth, development, and molt. Accurate data are not available on postnatal growth and development. Since the length of the smallest pup at the end of December in Rio de Oro measured 150 cm (Morales- Agasiono, 1950*), it may be assumed that the pups add 25 -30 cm during lactation. The period of molt of the neonatal hair coat is not known. It is also not known whether the pup remains on the coast throughout lac- tation or goes into the water. Some young ones with the neonatal coat fully preserved were caught in the sea. Thus, the Russian hydrobiological expedition in mid-August of 1912 caught on the Anatolian coast a pup that was still completely covered in a firm neonatal hair coat. It molted only after a few weeks (Zernov, 1913). In the second case, a newborn pup with a fleshy umbilical cord was caught on the coasts of Dalmatia. Both these cases point to the ability of even extremely young animals to enter water without molting the primary, neonatal coat. It has been pointed out in the literature, however (Troitzky, 1953), that the pup goes into the water for the first time at the end of lactation and possibly molt. The embryonic hair coat is thus sported for not less than 1.5 months. A newborn in captivity, weighing 26 kg, was bottle-fed for one week with a mixture of half oatmeal and half cow’s milk with a small addition of cod liver oil. This mixture was given six or seven times a day. The pup died due to injuries sustained during transport (Dathe, 1934). The total growth duration before the onset of sexual maturity has not been established. It has been assumed that the monk seal reproduces for the first time at about four years of age (Troitzky, 1953). Enemies, diseases, parasites, mortality, and population dynamics. Six species of helminths parasitizing the gastrointestinal tract were detected among monk seals (Delyamure, 1955): cestodes Diphyllobothrium hians, р. latum, О. lanceolatum, and Diplogonoporus tetrapterus; and nematodes Contracaecum osculatum and Terranova decipiens. These seals face no competition with other pinnipeds. Besides man, sharks pose a threat to the juveniles at some places. 390 59 Field characteristics. This is a large seal of deep dark monochromatic coloration on the dorsal side and a different coloration on the flanks and belly depending on age and sex. A large light-colored spot in the pos- terior half of the body is a characteristic feature among pups and adult males. This is the only species of seal in the Mediterranean and Black seas and also on the Atlantic coast of northwestern Africa. (K.Ch.) Economic Importance This seal has no economic importance because of its small popula- tion. In some countries (Bulgaria, Yugoslavia, and in Rio de Oro), this animal is protected. It needs to be protected throughout the range of its distribution. (K.Ch.) Subfamily of Hooded Seals and Elephant Seals, or 6-incisored Seals Subfamily CYSTOPHORINAE Gray, 1866138 These are seals of large and extremely large dimensions. The body length with tail along the dorsal surface (Lc) of adults varies from 180 to 500 cm or more, the males being slightly larger and heavier than the females. The hind flippers are slightly longer than the fore flippers. The first two digits of the fore flippers are longer than the third and subsequent ones. The claws are quite massive. By the time of maturity, the males grow a dermo-muscular, sac- or proboscislike process on the anterior upper portion of the snout joined to the nasal cavity. This is capable of considerable enlargement on being filled with air. There is one pair of teats. The skull is massive, broad, with a roomy cranium and zygomatic arches markedly protruding laterally; the zygomatic width considerably exceeds the width of the skull between the mastoid processes. The nasal processes of the premaxillae fall far short of reaching the nasal bones. The nasal opening is wide open and its upper and lateral margins extend far backward. The preorbital processes are well developed. 138 The independence of the subfamily Cystophorinae has come under dispute quite recently (King, 1966). Considering the controversial nature of this new interpretation of the family Phocidae, however, its existing structure has been retained. The subfamily is repre- sented in our waters by a single species, which is rather a random find. It nonetheless plays a role in our sea hunting activity in the North Atlantic and hence has been described fairly fully here. The species is also of interest because its biology and distribution were studied quite completely only in the last decade; yet although our scientists put considerable effort into its study, still little is known about it. (У.Н.) 392 520 The incisors in the upper jaw number two and, in the lower jaw, one on each side. The extreme upper lateral incisors are similar to the canines and slightly longer and thicker than the medial ones. At least the first two premolars have a single root and an almost wholly undivided crown. The cheek teeth are relatively small. Biologically, the species of the subfamily are extremely divergent. Some of them are pagophilic with no association with the land (hooded seal) while others have almost no association with ice floes and reproduce on land (elephant seals). All of them undertake extensive migrations. In the period of reproduction, males are confined mainly to one and the same rookery along with the lactating females and offspring. The pups at birth are not covered with a luxuriant embryonic hair coat of nearly white color, characteristic of the pups of most of the 10-incisored seals (Phocinae), but rather with a short firm hair coat. Some mem- bers of the subfamily (hooded seals) live in pairs at the time of mating while others (elephant seals) form harems. The former mate in water and the latter on land. Fairly large cephalopods and fish play an important role in their nutrition. Hooded seals molt far away from their breeding sites. Elephant seals, however, molt in the very same rookeries and with a far more intense scaling of the epidermis, which comes off in large patches. The distribution is interrupted and bipolar (borealnatal). Some members of the subfamily are distributed in the arctic and subarctic Atlantic (hooded seals) and others (elephant seals) mainly in the Southern hemisphere, predominantly along the edge of Antarctica, and enter only insignificantly into the Northern hemisphere on the western American coasts (Fig. 204). The origin of the hooded seal dates back to the Miocene. The seals of this branch originated somewhere in the west of the Old World; the earliest known finds of Cystophorinae have been dated to the lowermost Middle Miocene strata of France (Burgundy). The center of origin of the elephant seals is not known with certainty but possibly falls in the north. But there is a view that they evolved from the Antarctic 8-incisored seals (King, 1966) somewhere in the Southern hemisphere. The occurrence of elephant seals north of the equator in the Pacific Ocean, especially along the Californian coast is quite a recent phenomenon, evidently in the most recent stages of the Quaternary period. The route to this place ran apparently from the subantarctic regions along the Chilean coasts and was facilitated by the cold Peruvian currents. The system of the subfamily in the conventionally accepted form and as adopted here, is simple. It consists of two genera—hooded seals, Cystophora Nilsson with one species (see ahead), and elephant seals, 5211 Mirounga Gray with two species: M. leonine Linnaeus, 1758 and M. angu- stirostris Gill, 1866. Some suggested radical changes in the structure of the family, concerning also the subfamily Cystophorinae (King, 1966), as mentioned above, cannot yet be accepted. One genus, Cystophora, is distributed in the northern part of the Atlantic Ocean; one species of the second genus, Mirounga, in the Antarctic and the cold temperate waters of the Southern hemisphere (M. leonina), and the third on the western coasts of North America, somewhat north and south of 30° N lat. (transgressions into the north up to Vancouver). The ranges of the two species of the genus Mirounga are distinctly separate. Both genera are presently of economic importance but more so the hooded seal, which is a preferential target of sea hunters. Many rookeries of elephant seals destroyed in the last century are now being successfully restored by practicing conservation; at places, as in South Georgia, these are being exploited strictly along economic lines. In the USSR fauna the subfamily is represented by a single genus, the hooded seal, Cystophora Nilsson, 1820. But its presence in our waters is not a regular phenomenon (see below). From the middle of the 1950s, the hooded seal began to play a positive role in the economy of our country, as Soviet hunting ships were able to negotiate the ice floes in the Greenland Sea. (K. Ch.) Genus of Hooded Seals Genus Cystophora Nilsson, 1820 1820. Cystophora. Nilsson. Skand. Fauna. Dagg. Djur., I, p. 382. Cystophora borealis Nilsson = Phoca cristata Erxleben. 1911. Cystophoca. Brass. Aus dem Reiche der Pelze, 668. Substituted for Cystophora Nilsson. (У. H.) These are seals of large dimensions. The skull of the adults is massive, with thick bones, but relatively short and broad. The cranium is greatly enlarged but nonetheless stunted. In spite of the large volume of the orbits, the interorbital space is quite broaa, slightly more than the maximum diameter of the alveolus of the canine. The features of the bony tympanic bullae viewed from below resem- ble a trapezium somewhat; the posterior carotid aperture is disposed on their posterior side along the inner margin. The very large and open nasal aperture is highly enlarged in the posterior upper portion and extends posteriorly beyond the anterior margin of the orbits. The entire bony palate is very long and covers more than one-half the total length of the 522 ‘puluoa] Dsunop ‘[еЭ$ Зие44э]э WIDYyINOS э41 JO UOTINGLNSIP Jo souepunog aieunxoidde эцз 1214эр soul] usyo1g ‘эеи1л0ц401$ АЭ ‘s]eas pososioul-9 Jo Апитезап$ ay) Jo чоппа1“а “p07 ‘314 See Wy 000$ 0009 0006 0005 COCO! 0 001 165 393 573 skull. The posterior margin of the bony palate is fringed with a wavy line for the most part, like two symmetrical gentle notches converging medially and forming a projection set backward. This projection often has an additional narrower notch/bifurcation in the middle. The palatine bones are elongated and form a very broad bony palate. The choanae are broad and high; their longitudinal bony septum reaches or falls just slightly short of the posterior margin of the bony palate. The length of the zygomatic bones (without processes) is usually not more than double the smallest width. The lower (or posterior) processes are slightly longer than the upper. The maxillary bones are contiguous with the nasals in a small section; their length is usually about one-third that of the nasals and usually less than their width. The jugular (paroccipital) processes are well developed and bent backward. The nasal bones are considerably advanced forward and are very prominently suspended above the nasal opening. Their anterior portion is fairly enlarged and usually terminates in an obtusely angled median projection (see Fig. 206). The rostral portion of the skull has no bulges anterior to the orbits and has a concave and not convex (as in 10-incisored Phocinae) profile when viewed from above. The angular process of the lower jaw is weakly manifest. Cranial sutures not persisting until old age, in addition to occipital (including the lambdoidal) and basal sutures and, in rare cases, even the anterior palatal suture, are characteristic. The dental formula is: 2 1 4 1 La) Ст» Pig Mi. The canines are very massive; the lateral incisors in the upper jaw are slightly larger than the inner ones and similar to the canines; the molars and premolars are relatively small, with low crowns, slightly flat- tened from the sides and almost not differentiated. As long as they are not worn out, only a shallow indentation is visible on some as though separating the rear section of the crown in the form of a denticle which, however, may not be present at all. Sometimes, a similar additional den- ticle is seen in the front. The true molars of the upper and lower jaws have two independent roots. The rear premolars to quite often have two independent or partially used roots. Adult males are perceptibly larger than females. The claws are well developed on the fore as well as hind flippers. The color of the adults is bright and spotted. The whiskers are horn or brownish in color, flattened, and with wavy edges. On the upper side of the snout from the nostrils to the forehead, the adult males have a hollow dermo-muscular saclike 394 524 process formed by an overgrowth of the nasal cavity. At rest, it is barely visible and hangs limply with its fore edge above the tip of the snout. It can be highly inflated during excitation. These seals are biologically associated with drifting ice floes. They do not form harems. They feed on fish and cephalopod mollusks. Distributed in the subarctic and arctic Atlantic from Canada to the western parts of the Barents Sea, insome years up to the White Sea inclusive. These seals undertake distant migrations from the wintering and breeding sites to the regions for molt and feeding. Their main biotope is the marginal strip of drifting sparse pack ice. In the period of feeding they cover even much farther boreal regions to the south. The origin of the genetic branch has not been traced conclusively with the finds of teeth reported thus far dated to the Middle Miocene formations of France and the Pliocene formations of Belgium. Mesotaria ambigua, described from these finds, is regarded as a possible predecessor of the contemporary genus Cystophora. An immediate problem for taxonomists is to provide a more thorough explanation of the morphological differences and similarities between the hooded seal on the one hand, and species of the subfamily of true seals (Phocinae) and the subfamily of monk seals, Monochinae, on the other. The genus consists of a single species, the hooded seal, Cystophora cristata Erxleben, 1777. This seal is a random find in the USSR where it penetrates into the northern part of the White Sea (its inlet and neck sections). As one of the most important targets of sea hunting, this seal is caught predominantly in the international waters near Jan Mayen, Iceland, Green- land, and Canada for its skin, fat, and other byproducts. (K. Ch.) HOODED SEAL!?? Cystophora cristata (Erxleben, 1777) 1777. Phoca cristata. Erxleben. Syst. Regn. Anim., I: 590. Southern - Greenland and Newfoundland. 1785. Phoca cucullata. Boddaert. Elen. Anim., p. 107. 1820. Cystophora borealis. Nilsson. Scand. Faun. Г: 383. Southern Greenland and Newfoundland. 139 This seal is sometimes referred to as the Atlantic gray seal (“tevyak’”’), which is not correct as this name has been assigned to another species (Halichoerus grypus). Hunters sometimes call it the “klappmyss,” a word borrowed from the Norwegians. 525 1824. Phoca leucopla.'*° Thienemann. Reise im Norden von Europa, р. 102. Grimsey Island, north of Iceland. 1825. Phoca mitrata. G. Cuvier. Oss. foss., 5, p. 210. 1843. Phoca isidorei. Lesson. Rev. Zool., p. 256. d’Oleron Island, France. (V.H.) Diagnosis Only species of the genus. Description In general appearance, the hooded seal is similar to the large species of the subfamily of true seals, 1.е., common or harp seal, except that it is larger (Fig. 205). The body is elongated and the fore flippers have a significantly trun- cated posterior upper margin. The hind flippers do not form a deep recess ON maximum expansion. Пре WZ = Е SE: FP PLCS: р I a CE, ae =F = и ae 2 _ = и Fr р Wye 7 395 Fig. 205. Hooded seal, Cystophora cristata, mature male and female (figure by М.М. Kondakoy). 140 Ellerman and Morrison-Scott (1961*) assign this name to an individual variant of the harp seal. (V. H.) 395 526 The length of the anterior portion of the body from tip of nose to base of fore flippers is about 31% in females and 34% in males; length of fore flipper in both males and females about 14%, and maximum body girth about 65% of body length (L. Popov, 1960). Adult males differ sharply from all the other seals of the North Atlantic in having a special saclike formation on the anterior upper part of the head. It represents a spacious paired enlargement of the nasal cavity between the skin cover and skull and nasal cartilage longitudi- nally divided for the most part by an elastic mucous septum. In the normal (collapsed) state, it is comparatively less recognizable because it is externally bound on the nose bridge by a transverse fold and anteri- orly [it] forms a pouch of the rostrum hanging above the mouth opening. When the animal is in a state of excitation, this saclike cavity becomes highly inflated under the pressure of air generated by the closure of the nostrils and stands out prominently above the anterior portion of the snout in the form of a high and thick semicircular balloon with a length up to 30 cm and height up to 25 cm (Fig. 207). In addition to inflating this subcutaneous sac, the males are also capable of blowing a typical balloon out through either nostril. Such a balloon is formed by inflating with exhaled air the elastic longitudinal mucous membrane separating the saclike cavity.!*! The hair coat of adults is distinctly dichromatic. Dark spots of extremely variable size and shape are scattered quite haphazardly on a light gray background (sometimes darker on the dorsal than ventral side). These spots are slightly angular and quite often of fanciful shapes that frequently merge when the contours are particularly ornate. The spots in the midportion of the body are considerably larger, with more complex outlines, but more sparsely scattered than in the anterior portion, where they become very small close to the head and look like dabs. The spottiness on the dorsal surface is somewhat denser than on the ventral side. The spots are dark brown, often brownish-black or almost black. The anterior portion of the head, hind flippers, and tail are more deeply pigmented and often monochromatically dark. The speckles on the fore flippers are usually quite distinct. Among males with this type of coloration, much darker animals are sometimes encountered. The spots in them are so large and numerous, especially on the upper part of the body, that there is virtually no space left for the main background; hence such animals appear to be 141 Brondsted, 1931*; Olds, 1950; Trenze, 1950*; Berland, 1958; Mohr, 1958*; Sierts, 1958*; L. Popov, 1960; К.К. Chapskii. 396 397 527 Fig. 206. Skull of the hooded seal, Cystophora cristata (figure by N.N. Kondakov). Fig. 207. Schematic structure of the nasal cavity of the male hooded seal, Cystophora cristata, at rest (A) and in a state of excitation, filled with air in one of the paired chambers (B), causing extension and inflation of the elastic nasal septum (С). 1—posterior portion of the nasal cavity; 2—eye; 3—cartilaginous section of the nasal septum; 4—elastic nasal septum; 5—anterior portion of nasal cavity; 6—nostrils; 7—inflated balloon (after Berland, 1966). a monochromatic grayish-black from a distance. The females are practi- cally indistinguishable from males in coloration though there are refer- ences that the main background among females is lighter and the dark spots more contrasting (Collett, 1911-1912). The age-related color changes of the skin are significant. The first coat with which the pup is born is characterized by the total absence 396 397 528 of spots and sharp color differences between the ventral and dorsal surfaces. The upper part of the body from the tip of the nose to the tail is a monochromatic bluish-gray with a bluish-olive tinge. This col- oration changes sharply on the flanks to almost total white on the lower side of the body. The color of the inner and lower sides of the flip- pers is similar. The boundary between the dark and light colors runs from the lower margin of the nostrils 1.5-2 cm below the eyes almost through the ear opening or slightly above it, diverging onto the sides of the [shoulder] blades, and forms a long narrow projection in the mid- portion of the outer surface of the fore flippers. Posterior to them, the boundary extends along the flanks of the body (above the axillae) and reaches, almost without flexure, the hind flippers, terminating in their upper part, and onto the tail. Such an unusual coloration of pups is seen in the last stages of embryonic growth, a few weeks before birth (Reinhardt, 1865*). Another characteristic feature of the hair coat of the newborn is that it is not typical of seal pups born on ice floes, 1.е., soft, wavy, and very dense; instead the embryonic coat is short, rather dense, and somewhat stiff. This type of coat is seen among other pagophilic seals after the embryonic coat has been shed, 1.е., the coat of the newborn corresponds to the second coat of other seals. The embryonic coat is seen in the later stages of fetal growth but is shed in the womb itself sometime before birth. The discarded embryonic hair, in the form of dense matted feit and round disklike clumps, is often detected in the amniotic fluid (Olds, 1950; L. Popov, 1959; Yakovenko, 1959; V.K. Shepeleva). The mech- anism of matting of. the embryonic hair is not wholly clear but this is hardly due to fetal movements. It is more probable that the sloughed hair is swallowed by the fetus and becomes rolled into clumps during passage through the intestine under the influence of peristalsis and during excre- tion (Mansfield, 1963). The hair clumps consist of fairly homogeneous light gray hairs, thus revealing the color of the embryonic coat. The hooded seal is among the largest of the true seals. The body length along the dorsal surface (Lc) of adult males from the Greenland Sea measures 210-250 cm, the largest reaching almost 280 cm; the body length of most adult females is 180-200 cm but some measure almost 227 cm. The average length of males is 235.7 cm, of females almost 195 cm (L.A. Popov, 1959, 1961).!42 12 According to Degerbol and Freuchen (1935), the largest males reach a length of 350 - 400 cm, which is rather doubtful (the method of measuring the length was not specified; it was probably with the hind flippers). According to Mansfield (1963), the average length of males from the western Atlantic is 233.7 cm and that of females 203.2 cm. 398 529 The weight of adult males (length along the dorsal surface Lc 225 - 240 cm) is 260-300 kg; adult females (200-215 cm long), however, weight 145-160 kg (L. Popov, 1959, 1961). The average weight of Canadian-Newfoundland male hooded seals is considered as 317.5 kg (Mansfield, 1963). A large male measuring (Lc) 275 cm long caught on October 31, 1954 on Faeroe Islands weighed 375 kg (Ehlers, Sierts, and Mohr, 1958). The condylobasal length of the skull in adult males (Fig. 206) varies from 241.5 to 293, 0, average 271.5 mm; in females 218.0 to 238.0, average 225.3 mm; zygomatic width in adult males 158.0-223.0 mm, females 155.0-176.0 mm; mastoid width in males 151.0-171.0 mm, females 144.0- 156.0 mm; length of upper tooth row in males 58.0-85.0 mm, females 50.0-65.0 mm (L. Popov, 1959, 1961). The os penis in adult males is 20.5-21.0 cm long, 2.5 -2.6 cm wide, 2.0 cm thick, and weighs 32-38 в (Mohr, 1958*). (К. Ch.) Taxonomy Only species of the genus. Geographic Distribution Predominantly the pelagic regions of cold streams along the southern fringe of the arctic zone and its adjoining northern sections of the boreal strip of the Atlantic. Geographic Range in the USSR Only the extreme northeastern branch of distribution extends to the ter- ritorial waters of the USSR, not beyond the northern sections of the White Sea covering its inlet and neck sections. Stray hooded seals are sometimes encountered here; family groups are rarer. These are more in the nature of stray transgressions of mothers with pups than regular vis- its. The hooded seals transgress into these regions possibly together with herds of harp seals migrating to breeding sites. Such instances, though not arising every year, are well known to the White Sea hunters who sometimes sight and catch a few animals while hunting for harp seals. Geographic Range outside the USSR The southern distribution of the range passes slightly south of the winter boundary of drifting ice floes from the coastal zone of America in the region of the Gulf of St. Lawrence and Newfoundland Island, encom- passing the extensive pelagic expanse to the south and east of the latter and extends (with some bend toward Davis Strait) toward the waters 398 530 of southern Greenland. Encircling its southern extremity, it runs farther northeast, covering Denmark Strait, and extends north and northeast and especially east of Iceland through the Greenland Sea and northern part of the Norwegian Sea, then descends close to the Scandinavian Penin- sula considerably below the boundary of winter drifting ice floes. Along their fringes and receding from the peninsula to the south, evidently up to the coastal belt, the boundary extends to the southwestern portion of the Barents Sea to the threshold of the White Sea into its northern part. The northern boundary of distribution is the fringe of dense accumu- lations of perennial arctic pack ice between Spitsbergen and Greenland and evidently the same latitude in the straits between western Greenland and the eastern fringe of the Canadian archipelago (Fig. 208). Within these limits of the main distribution, the populations of the hooded seal are unevenly distributed. Associated in distribution with the fringes of fairly sparse drifting ice floes, the hooded seals in the course of their annual cycle drift alternately north and south, partly due to the seasonal variations in the position of the ice floe fringes. The southern boundary of distribution in the winter extends from the southwest, roughly from 45° N lat. at Newfoundland, to the north- east up to the Polar Circle (perhaps slightly south along the Norwegian coasts). In summer, on the contrary, it shifts into a more arctic position, 1000 2000 3000 4000 5000 km | | jee ie Fig. 208. Distribution of the hooded seal. 1—breeding zone; 2—molting zone; 3—probable courses of migration; 4—including with pups (after Rasmussen, 1960, with additional information); 5—distribution boundaries including probable boundaries (broken line). Dots in the White Sea represent sites of transgression into the USSR waters (К.К. СварзКи). 399 aye) toward 60° in southern Greenland and Labrador to the Polar Circle in the northeastern regions of Iceland and in the much higher latitudes toward Spitsbergen and the western part of the Barents Sea. Some pop- ulations migrate in the course of the year from some sections of the distribution to others in search of food or sites for breeding, molting, and wintering (see “Seasonal Migrations and Transgressions”’). The pas- sive migrations caused by these factors and the active migrations in a relatively short period sharply modify the boundaries of distribution. This reduces the overall zone of distribution into a tentative concept, defined as the maximum distribution range. Thus the distribution is extremely dynamic. The distribution can be described in greater detail as follows: its western boundary could be drawn along the Atlantic coast of Canada to north of Nova Scotia, encompassing the Gulf of St. Lawrence and the extensive zone of open waters of the Atlantic around Newfoundland, along the coast of Labrador to Hudson Bay, which is intersected by it in the westernmost section, and farther north along the entire coast of Baffin Land with almost the whole Strait of Lancaster and the Admi- ralty Strait opening into it included. Even more northward, the distribu- tion includes the entire eastern rim of Baffin Land, Smith Strait, Kane Basin, and Robson Bay. This represents the northern limit of distribution between Canada and Greenland. Subsequently, following the coastline of western Greenland, the boundary descends to its southern extremity. Thus Baffin Bay and Davis Strait fall wholly in the distribution. Along the eastern coast of Greenland, the boundary runs from Cape Farewell roughly up to 75° М lat., is separated there from the land usually by broken ice floes, tends northeast up to Spitsbergen, descends along its western side and runs east into the Barents Sea reaching 30° (Nansen, 1924) and even 50° E long. (L.A. Popov, 1960). From this extreme east- ern limit, the distribution turns southwest (White Sea) and later, entering the southern boundary traces back in the direction described above. Since breeding among hooded seals is largely localized, some exten- sion of their spatial division is possible. Apparently, there are three fairly isolated geographic groups, which vary in population and importance. Two of these groups inhabit the western part of the range from the Gulf of St. Lawrence and Labrador to the northern region of Davis Strait and southern Greenland in the east to Denmark Strait. In the breeding period the hooded seals of these groups are concentrated in two regions: on both sides of Newfoundland Island, mostly north and northeast of it, 43 п compiling the geographic distribution, the data of @ritsland (1959), Rasmussen (1960), Mansfield (1963), and @ynes (1964) were used. 400 532 and to а lesser extent westward in the Gulf of St. Lawrence. The third population is distributed in the northeastern part of the range on the ice floe edges between Denmark Strait and Spitsbergen. Jan Mayen serves as the region of whelping for this group (for more details, see “Seasonal Migrations and Transgressions’’). (K.Ch.) Geographic Variation Not studied. Biology Population. The population of the hooded seal has yet to be precisely determined. At the end of the 1950s, the world reserves were put at 300,000 - 500,000 (Scheffer, 1958). Aerial surveys were made in 1959 and 1960 in Denmark Strait. The total population of the species was put at 500,000 (Gritsland, 1960). Some higher estimates, also assumptions, put it at 500,000 - 700,000 (Rasmussen, 1960). The long-term steady hunting in Denmark Strait could serve as a rough index of seal reserves. On average, 62,500 hooded seals were caught there annually from the 1870s. Only a very large herd of hun- dreds of thousands could have sustained such large catches. Considering this phenomenon, a figure of 500,000 seals was cited even in later years (Chapskii, 1966). Habitat. Mostly similar to that of the harp seal with which the hooded seal shares much in ecology. Both species are confined to drifting ice floes, mainly to their fringe zones. But while the herp seal prefers com- paratively young and low ice floes, the hooded seal selects contrarily very stable perennial pack ice for reproduction. The sites of the hooded seal invariably have frequent waterways since this seal does not make air holes in the ice floe (Nansen, 1924; L. Popov, 1960). As it avoids dense accumulations of ice floes, the hooded seal is not encountered on shore ice. It usually does not enter the deeply protruding bays and straits or narrow straits between islands packed with ice floes. Ц does not enter deep into the Canadian archipelago, transgressing only into the more southern broad straits opening into Davis Strait and Baffin Bay. For the same reason, it does not approach coasts densely packed with ice floes, remaining confined to their outer fringes. During the breeding period hooded seals are usually encountered inside the ice massif (which the hunters’ schooners cannot always pene- trate), on stable arctic pack ice, and around very young ice floes alternat- ing with open water pools. The accumulation of ice floes with open water 401 583 pools is particularly characteristic of regions with a slow, often circular drift at the confluence of opposing currents. Jan Mayen constitutes such a region of relatively stable, coarsely and finely broken ice floes. Here, as a result of the interactions between the cold eastern Greenland streams with the northwestern branch of the Gulf Stream, favorable conditions arise for feeding as well as for prolonged residence on the drifting ice floes. The hooded seals find very similar conditions in Denmark Strait, which represents their molting region. To a much greater extent than the harp seal, the hooded seal is a pelagic inhabitant of open expanses and is seen close to coastlines only in the period of migrations; at this time it prefers the coastal strip of southern, mainly southwestern Greenland. In winter and early spring it is dispersed sparsely in pelagic regions, far from ice fringes, especially in the Norwegian Sea (see “Seasonal Migrations and Transgressions’’). In this very period of winter fattening it is also confined to the western Atlantic in the vicinity of the Newfoundland coasts and evidently close to Iceland (Wynes, 1964). Like other pagophilic seals, the hooded seal requires ice floes for reproduction and molt. This need is also felt when the animals are not fat enough and hence suffer excessive heat loss in water. Ice floes play some role also as a substratum for resting and provide a means of escape from aquatic enemies. The affinity for a strip of ice floes is evidently caused by food conditions, promoted by the polar front zone in general, and the available strip of spring-summer ice floe fringes in particular. Food. This aspect has not been studied. Only very general refer- ences are available regarding specialization of the dental apparatus for grasping and tearing large adible objects including fish (Nansen, 1924; L.A. Popov, 1961). These are essentially the sea perch, flounder, cod, and cephalopod mollusks; the seasonal changes in the distribution of the hooded seal are probably related to the migration of cod. In the period of suckling pups and even during molt, adults and others in the . transitional age group do not obviously cease to feed and go out to sea quite often. Their stomachs revealed food remnants, in particular squids (L.A. Popov, 1959; М.Уа. Yakovenko). Nevertheless, feeding in these periods is very poor and highly irregular. During lactation and molt the animals become quite emaciated (Rasmussen, 1960); intensive feeding occurs after mating, before congregation in the molting region, and also at the end of molt. On transition to independent feeding, the young seals initially feed on amphipods, small squids, and other edible objects. An adult male in a 534 zoological garden was fed on herring, whiting (hake), cod, coalfish [pol- lack], flounder, witch flounder, turbot, and sea perch; this seal refused to eat the latter, however, when a choice was available. Initially, when live fish were fed, it consumed eel, carp, and tench. There was a distinct preference for herring, with up to a hundred consumed daily (Ehlers, Sierts, and Mohr, 1958). Home range. It is practically impossible to establish the home range of such a widely migrating and wandering animal as the hooded seal. The area of drifting ice floes which the seals select for reproduction is also not amenable to precise calculations, albeit the animals are confined to the same icy regions at this time. Much depends on the actual ice conditions of a given year, type of drifting ice floes, features of approach, and the initial disposition of the animals. Moreover, the density of animal disposition varies even during the short duration of lactation under the influence of ice floe movement. The animals live in “families” in the regions selected for reproduc- tion. The whelped female is confined to a definite ice floe with her pup; the adult male also stays closeby on the same ice floe, forming a mating pair with the chosen female. Such pairs with suckling pups are disposed at different distances from each other. Generally, each pair occupies a separate ice floe. But when the floe is very large, other pairs occupy it but at distances of tens of meters or sometimes far less. Thus occasion- ally the hooded seals are so densely disposed at sites of reproduction that the latter resemble the nursieries of harp seals (Rasmussen, 1954*; A.V. Potelov). In general, however, very uneven distribution of the seals is a char- acteristic feature during the nursery period. In the large concentration north and northwest of Jan Mayen Island on March 13, 1962, on aver- age five mating pairs with pups were encountered per mile of traverse. On March 24 of the same year, slightly eastward, up to 15 “families” were counted on some ice floes measuring roughly 100 x 100 m; 15 to 25 animals were sighted from a ship on March 26 on the spits of a coarsely broken ice floe (Khuzin and Potelov, 1963). In general, in the winter-spring season some mating pairs occupy from 670 m? to roughly 0.6 km? at reproduction sites. Sometimes such mating pairs with pups are typically interspersed among masses of harp seals (R.Sh. Khuzin). During molt on the ice floes in Denmark Strait the disposition of hooded seals is denser and more stable. The distance between animals is just a few tens and sometimes even a few meters. During local move- ments and more so during migrations, the concept of “home range” is 402 535 no longer applicable due to the wide scattering of the animals and their uninterrupted movements. Hideouts and shelters. The hooded seal does not seek hideouts and shelters on the ice floe or in the snow. As a rule, it does not make air holes in the ice floe but utilizes the natural openings and open water pools for emerging onto and exiting from the ice. When, however, the open water pools have a thin ice cover, the seal can pierce it for res- piration. In 1956, a hooded seal under observation pierced a young ice cover 4.5 cm thick with its head, thrust up out of the water, surveyed the surroundings, filled its lungs, and submerged never to be seen again. Perhaps the seal managed to breathe at other places in a similar manner as Open water was nowhere visible, the entire surface was frozen and calm, and frost prevailed (M.Ya. Yakovenko). Daily activity and behavior. These aspects have not been studied. During the molting season, the animals rest long periods on the ice floes interspersed with brief spells of submergence in water. Males and females enter water even more often during the period of suckling the pups. The animals spend time predominantly or exclusively in water for the purpose of feeding and moving about. Seasonal migrations and transgressions. Hooded seals are among the widely migrating seals. Although not much is known about their migra- tions, their general characteristics have been established. During the breeding season the productive males and mothers are grouped in two widely separated regions. One group concentrates in the southwestern part of the range, on the southern extremity of Labrador opposite the Strait of Belle Isle and the northernmost extremity of Newfoundland. A small population locates in the Gulf of St. Lawrence. The other region occurs on the eastern boundary of the range, northwest of Jan Mayen Island. The hooded seals reproducing in the westernmost section of the dis- tribution, having completed this biological cycle by early April, abandon the ice floes for sites abounding in food. They first travel along the edges of Labrador north toward Davis Strait and later, turning east, approach the western Greenland coasts rich in fish and feed for sometime right at the banks as also in the coastal regions of southwestern Greenland. Further, some groups of seals move higher and approach Greenland at latitudes 65 to 66° and others (the majority apparently) remain in the feeding grounds, not moving north of 60 to 61° N lat. and are seen roughly at the same time in May on the southern coasts of Greenland. Having recovered their fat reserves to some extent by the end of May to early June, the hooded seals reach the southernmost tip of Greenland, move round Cape Farewell, and continue their course along 403 536 eastern Greenland (instances are known of hooded seals marked near Newfoundland being recovered at Cape Farewell). Later, the seals migrate farther northeast along the ice edges to Denmark Strait, the main (though not the only) site where hooded seals gather in large numbers and remain for molting in June and July. In its first year, the juvenile does not undertake this migration east- ward; it migrates evidently farther north into Baffin Bay and remains there until August. Having completed molt, the adult hooded seals begin to leave Denmark Strait in the first half of July and scattering widely, return by the same route to the southern tip of Greenland, and passing it repeat their journey in a reverse direction. Roughly from mid-July through mid- August, hooded seals emaciated during molt are caught on the coasts of southwestern Greenland. Returning to their birth place, the Canadian- Newfoundland population again visits the western Greenland fish banks and feeds well. These seals then pass through Davis Strait and are seen on the Labrador coasts in early autumn, proceeding south into the region of the Great Newfoundland banks for feeding. They return north in February to the whelping zone and concentrate again on ice floes before commencing a new annual cycle.!*4 The populations reproducing in the region of Jan Mayen Island, after whelping and mating also scatter quite extensively but their migration does not attain the scale that is characteristic of the Canadian-Newfoundland populations. The pups abandoned by their mothers migrate initially passively with the moving ice fringes and are encountered at April end to early May as highly scattered solitary animals in the fringe zones from 69 to 76° N lat. (L.A. Popov, 1960; and others). From April to June the adults are scattered even more widely among thin ice floes and beyond the fringe zones. They are then capable of dis- tant migrations in search of food. These migrations sometimes extend right up to the White Sea. A large collection of hooded seals, some- what like a nursery in the Greenland Sea (on Jan Mayen Island), was encountered around May 10, 1962 in the middle of the Barents Sea (73° 38’ -74°00'N lat. and 24°28’ -35° 20/E long.) (V.A. Potelov). Much of the population nevertheless tends southwest into Denmark Strait where it molts in June. The rest is scattered on the fringes of the Green- land Sea to north of Jan Mayen roughly up to 75° N lat. and molts there on pack ice (Wolleback, 1907; L. Popov, 1959; R.Sh. Khuzin). 144 According to Allen (1880), Nansen (1924, 1939), Bartlett (1928*, 1929), @ritsland (1959), Rasmussen (1960), Mansfield (1963), and others. 404 557 At the end of the molting period, the Jan Mayen hooded seals are again scattered over an extremely wide expanse whose boundaries were only recently deciphered. It was assumed that they were confined to the high latitudes and appeared near the mainland only very rarely. There were frequent reports from fishermen using nylon nets to catch cod and halibut in the Norwegian Sea, of hooded seals being caught in them at Olesund, northern Tronnelag, Vesteralen region, and other places along the Norwegian coast. Hooded seals were seen there even in winter and were very Common in spring. Such instances were particularly numerous in the spring of 1959 (@ynes, 1964). These facts compel us to recognize that, in winter and even in spring, the seals of this species stay regularly at relatively low latitudes not only in pelagic zones, but also close to the coasts, entering even into the Norwegian fjords. Instances have been recorded of scattered solitary animals at differ- ent points on the coasts of Scotland and England; one find has been reported even from Biscay coast of France. In most cases these were young animals, some of which had only just been born. Along the Amer- ican coast of the Atlantic Ocean south of Nova Scotia, instances have been reported even recently of the appearance of solitary animals in the region of New Jersey and on the coast of Massachusetts; a dead female was found right on the coast of Florida in the winter of 1916 (Miller, LSI): The transgression of a hooded seal into the eastern zone of the Yenisey estuary is wholly exceptional in the matter of distance involved and is inexplicable (N. Smirnov, 1908, 1929). Reproduction. The period of mating sets in immediately after the completion of lactation and extends roughly for two weeks. Even by around April 10, ovulation has occurred among almost all the mature females; the bulk of the animals mate in the first week of this month (L.A. Popov, 1959; Oritsland, 1964). Precise information is not available about the copulation process but animals were observed mating in water. The pair slowly floated close to the surface, periodically thrusting out for respiration. The extremely large size of the males, larger than the females, suggests that they woo the females anew each year. Traces of fights in the period of heat have been detected on the skin of males (Yakovenko, 1959). The lag in implantation of the blastocyst has been established as 3 to 4.5 months from the time of mating, i.e., up to July end to early August (Oritsland, 1959, 1964). The hooded seals whelp in different parts of their range almost simultaneously, in most cases in mid-March; pups appear only very rarely at a much later period. 404 538 Fig. 209. Pup of hooded seal with mother. Jan Mayen region (photograph by A.V. Yablokov). Twins have not been reported among the females studied but some- times two pups have been seen near the same female. Unlike other seals, sexual maturity among hooded seals sets in rel- atively early; in rare instances, the first ovulation occurs at the age of two years but in most cases does not result in gestation (Oritsland, 1964; Yakovenko and Khuzin, 1965*). Even at the age of three years, only half of the females are pregnant. Only 80% of the 4-year-old females whelp while 10% of the five-year-olds still remain nonproducers (@rits- land, 1964). Subsequently (most often from seven to eight years), females whelp every year (number of barren animals is not over 5%). The ability to become fertilized and whelp is maintained right up to the end of their lives, which can be taken as 35-36 years. The onset of sexual maturity among males occurs somewhat later. Until three years of age, they are mostly immature.'® At four years some males evidently become mature but number about 30%. This rises to 80% among five-year-olds (L. Popov, 1960). Commencing from six years of age, all males are mature (Khuzin and Potelov, 1963) though, 145 According to Yakovenko (1959), 3-year-old males (with four layers on the cross section of the canine) are already mature and evidently come into heat since the skin of one such animal revealed wounds inflicted in a fight. 405 539 according to other authors (L.A. Popov, 1960), immature animals are still encountered at this age, their proportion reaching 13%. The sex ratio among the newborn is 1:1 as observed from the numer- ical ratio between the males and females in the molting rookeries. Of 1,125 pups examined, 49.2% were males (@ritsland, 1964). Growth, development, and molt. The length of the newborn along the dorsal surface, judging from the size of the fetuses measured in the period of whelping (about mid-March and even in the first few days of April), varies from 79 to 112 cm, on average 100 cm (Collett, 1911-1912; Nansen, 1924146 L.A. Popov, 1959; Potelov and Yablokov, 1967*; V.A. Potelov). The newborn weighs 12-27 kg, on average 19.3 kg (V.A. Potelov). Other figures too have been cited (23 - 30 kg) but the pups were probably already full grown and better fed.!*’ After 10-15 days of lactation, pups (Fig. 210) grow to 117 cm (Lc) on average while the weight increases up to 37.5 kg on average, the range being 23-55 kg (V.A. Potelov). The fat content of the milk has been reported as 43% (Khuzin and Yablokov, 1963) and on average 60% (with a protein content of 4-9%) (V.A. Potelov). Toward the end of the period of lactation, the thickness of the fat layer in the skin increases to 4-5 cm from 1-1.5 cm at birth (L. Popov, 1961; Shepeleva, 1971); of this, the thickness of the fatty tissue is 3.5-4.0 mm. The subsequent overall growth of the animal proceeds with the same intensity: the body length of yearlings measured along the dorsal sur- face (Lc) increases to 135-150 cm, up to 160-170 cm at two years, and 170-185 cm at three years of age (Yakovenko, 1959). According to others (Rasmussen, 1960), the body length measured along a straight line (Lcv) among yearlings of males and females is identical at an average of 133 cm!48; sex begins to exert a pull in the two-year-olds; the average length of males then is 155 cm, of females 152 cm. At three years of age, the males measure on average up to 172 cm, females up to 164 cm. The same intensity of growth continues up to 10 years of age (when the males grow to an average of up to 218 cm and females up to 193 cm). 146 Nansen and Collett indicate the length of the newborn as 90 cm. The measurement was evidently along a straight line, i.e., Lev. 147 The much older data of 3-5 kg (Ognev, 1935) or 6-8 kg (М.Р. Vinogradov, 1949) are undoubtedly erroneous. 148Треге is a view that sex-related differences in body length exist even at the time of birth and increase slightly during lactation (Potelov and Yablokov, 1967*) but the data in support of this view are not very convincing. 406 540 Fig. 210. Pup of hooded seal. Greenland sea (photograph by V.K. Shepeleva). Growth continues, although at a slower rate, up to at least 20 years of age. The age-related changes in color of the hair coat reveal some char- acteristic features. The original, neonatal color remains unchanged for a year. After the first molt (i.e., at the beginning of the second year), the first spots, not yet vividly pigmented, appear on the anterior portion of the back and partly on the neck (Nansen, 1924; Yakovenko, 1959). In the course of a year their number increases and these spots extend over much of the back and are also seen on the chest but not in the belly region. In the first few months of the fourth year (after the third molt), the main background on the upper side of the body turns olive-gray and is cov- ered with the innumerable dark brown spots characteristic of the color of adult animals. The underside of the body also acquires a brownish tinge and is speckled with spots scattered haphazardly. The snout becomes dark. On attaining the age of seven years, the main background turns lighter in color while the spots become darker (Yakovenko, 1959). This skin pattern evidently remains unchanged until the death of the animal. According to some authors, the immature animals of both sexes are dis- tinguished from the adults only in the number of spots, which increase far more with age; the main background, however, remains uniformly light in color (L. Popov, 1959b). 406 407 541 The nasal skin зас is not manifest at all among the newborn males in the first year. In the second year it becomes prominent as a “tiny cushion” slightly enlarging when the animal is excited (L. Popov, 1959b). The sac as such is manifest in the fourth year (Yakovenko, 1959). A real “hood” was seen in a five-year-old male (age established based on marking) (Rasmussen, 1957). The hooded seal begins to molt three months after the end of the whelping period. Denmark Strait is the main site of congregation of hooded seals of all ages (except under-yearlings) for molting; they gather there from around June 10 and leave toward July 20. Thus the molting period extends for 1.5 months. The molting rookeries of the hooded seal are less dense than those of the harp seal but are far denser than the nurseries of the former. There are no distinct age- or sex-related differ- ences whatsoever among molting hooded seals: the animals are disposed alternately in close proximity. Outside Denmark Strait, no other such region of en masse collection of hooded seals in the molting period is known, but small numbers of molting hooded seals are seen in Jan Mayen and Spitsbergen regions and generally almost all along the fringes of ice floes in the Greenland Sea (Qritsland, 1964; R.Sh. Khuzin). Enemies, diseases, parasites, mortality, competitors, and population dynamics. The young seal spends its early life on ice floes in an almost sterile icy environment under the protection of adults without exposure to any factor of natural mortality to which this population is sensitive. Later, when the juvenile enters the water and fends for itself on the fringe of ice floes, it may fall prey to the Greenland shark or killer whale. The latter possibly attacks rather frequently even the adult seals. Among other enemies is the polar bear but the hooded seal is much less threatened by this animal than the other arctic seals; furthermore, the population of this predator in the Jan Mayen region is not much. In July, 1956, the stomach of a polar bear killed in the Greenland Sea revealed the remains of a hooded seal pup (M.Ya. Yakovenko). This seal is parasitized by 15 species and two larval forms of helminths.!*? From among the trematodes, Metorchis albidus infects the gall bladder and bile ducts of the liver, Opisthorchis tenuicollis and Pseudamphistomum truncatum the bile ducts of the liver; from among the cestodes, Anophryocephalus anophrys and Diplogonoporus 49 The review of helminths was compiled by scientists of the Department of Zoology of Crimean State University based on the following data: Rudolphi (1819), Siebold (1848*), Brown (1893*), Price (1932), Berland (1963), Skryabin (1950*), Delyamure (1955), Delyamure, A. Skryabin, and Treshchev (1965*), Delyamure, Skryabin, and Alekseev (1964*), Treshchev, Zavaleeva, and Potelov (1967), and others. 407 542 tetrapterus infect the duodenum and the small intestine while Diphyllobothrium latum, D. pterocephalum, Pyramicocephalus phocarum, and Diphyllobothriidae g. sp. infect only the small intestine; from among the nematodes, Contracaecum osculatum, Terranova decipiens, Phocascaris phocae, Ph. cystophorae, and Anisakidae g. sp. parasitize the stomach and small intestine while Skrjabinaria spirocauda parasitizes the heart, lungs, and blood vessels; from among the acanthocephalans, Corynosoma strumosum and C. semerme infect only the intestine. The dissection of 293 hooded seals (Treshchev et al, 1967) established a high rate of infectivity; helminths were detected in 61.4% animals. If, however, the newborn (96 pups) are excluded from the total number of dissected animals and only the adults (197 seals) are taken into consideration, the percentage of infectivity rises to 91.4. The most severely infected seals were aged 1 to 12 years. The most frequently infected organ was the small intestine (in 58.02% animals); less frequently infected were the stomach (29.35%), duodenum (27.9%), and even less the heart (3.4%), lungs (1.02%), and liver (0.68%). Parasites sometimes cause severe diseases. The nematode Ph. cystophorae is capable of perforating the intestinal wall of the host. 5. Fig. 211. Head of an adult female hooded seal (photograph by A.V. Yablokov). 408 543 spirocauda, a parasite of the heart, probably kills the hooded seals at six to seven years of age (Delyamure and Treshchev, 1966). The seal louse Echinophthirius horridus parasitizes the skin and the seal mite Halarachne halichoeri the nasal cavity. Competition in the feeding grounds can arise only from the harp seal. In practice, however, there is no such competition as the food reserves suffice to meet the needs of both species; furthermore, the hooded seal, more than the harp seal, lives on deep-water nektonic life. Had there been an acute competition, the two species would not probably have formed nurseries in the same regions. Most of the feeding grounds of these two species are nonetheless distinct. Some idea of the population reduction can be gleaned from hunt- ing statistics, especially a perceptible increase in younger animals in the herd, i.e., a relative reduction in the number of mature animals and an increased percentage of young ones. In 1956, the young constituted 27.5%, which rose to 34% in 1957 and to 49.5% in 1958. It was consid- ered that the killing was exceptionally high in the 1950s and the state of the herd at the beginning of the 1960s caused anxiety (Rasmussen, 1960), necessitating a ban on hunting in Denmark Strait. In turn, during the years of World Wars I and П the forced cessation of hunting using ships promoted the restoration of reserves. The reduction at present is, however, quite perceptible. Hunting is the main factor responsible for the population dynamics of the hooded seal. Field characteristics. The under-yearlings are easily distinguished by the two-toned coloration of the skin, 1.е., bluish-gray on top and white below. The adult animals are recognized by their large size and con- trasting spottiness. The hooded seal can readily be distinguished from the harp seal with transitional (gray spotted) coloration by the former’s fairly large blackish-brown and dark brown spots often fusing into fanci- ful patterns on a gray background with densely scattered spots and dabs in the anterior portion of the body. A positive feature of recognition is the saclike process on the head of adult males, which is quite often inflated through the nostrils (Fig. 212, A, B). Distribution in separate “families” on ice floes during the nursery period is also a useful species characteristic. (K.Ch.) Economic Importance The economic importance of the hooded seal in the world hunting indus- try is quite high. As an object of hunting using ships in the North Atlantic, it occupies second place after the harp seal. The main region of hunting now falls in the international waters near Jan Mayen. Sections 544 1 i Try yyy МЫ РР! > 409 Fig. 212. Head of an adult male hooded seal. A—with a “cap” slightly inflated by the nasal cavity (after Mohr, 1963); B—schematic depiction of fully inflated nasal cavity (“сар”) (figure by К.К. Chapskii). 409 410 409 545 of the “polar front” to the north of Newfoundland are of much less importance. In both these regions the hooded seal is caught together with the harp seal. Until recently, hunting was practiced in Denmark Strait and half a million hooded seals were caught there in the 1860s. In good hunting seasons at the end of the last and early in this century, 15 to 20 hunting ships caught 40,000 to 57,000 seals. In the region of Jan Mayen, however, even in the first two decades of this century relatively few hooded seals were caught. Large congregations were not found in this area. The catch rose markedly from the 1920s. In 1926, Norwegians caught 53,000 hooded seals in the expanse from Spitsbergen to Denmark Strait (Sivertsen, 1941). In the postwar years (from 1956 through 1958), hunting notably decreased in Denmark Strait and was totally banned in 1961. In the region of Jan Mayen, however, it continued at a high level. From 1946 to 1958, it averaged at different times 35,000 - 59,000 - 44,000 seals per annum. The catch of hooded seals in the Newfoundland area (in which Norwegians and Canadians hunt) represented about 10% of the hunt in the Jan Mayen region. In these years the catch averaged 5,700 animals, mainly pups. The Soviets took to hunting the hooded seal in 1955 and continued to do so up to 1967. The ice floes of the Greenland Sea in the region of Jan Mayen constituted the hunting region. Here the harp seal was the main target. The extent of hunting is depicted in Table 29. Relative to the harp seal, also hunted in this region, the catch of the hooded seal varied from 10-15% of the total catch in the early years of hunting to almost 70% in some subsequent years. There is nothing specifically different in the hunting of hooded seals using ships compared to harp seals except that the ships have to operate under more severe icy conditions, often approaching each “family” group separately. The hunting operations for some reason or the other involve Table 29. Catch of hooded seal by Soviet hunting ships in the Greenland Sea (R.Sh. Khuzin) Year Total catch Pups in total catch 1960 4,575 1,472 1961 11,781 7.101 1962 9,562 3,874 1963 7,303 3,470 1964 8,861 4,706 1965 8,380 5,326 546 great difficulties as well as much time. Hunting itself is not а very com- plicated affair due to the slower self-preservation reflexes of the animals © in the period of reproduction. The main problem is locating the rookeries since aerial assistance is not used. Although the hooded seal represents an enticing target (an adult animal yields more fat than a harp seal, while the skin of a juvenile hooded seal is highly valued as a commercial fur), it was not specially sought in the early years of hunting and only whatever came in the way of a ship was taken. Later, however, hunters laid greater emphasis on the hooded seal. After World War II, the Norwegian hunters invariably sought the juveniles so that the catch of pups exceeded that of adults, amounting to 55.5-86% of the total catch. On average, in the first 13 postwar years the Norwegian hunters killed 70% of the juvenile hooded seals available in the Jan Mayen region. The fashion for seal skin over- coats played no mean role in this regard. The hair coat of the pups is distinguished by denseness, moderate luxuriance, excellent color and luster, while the skin is quite durable. The skin with the subcutaneous fat layer when removed by the hunters (without the flippers) weighs 5 to 11.5 kg (average 9.1) in the case of a newborn but averages 20 kg at the end of lactation. The weight of an adult skin with the fat in early spring (during the breeding season) in the case of males (length along the dorsal surface, Lc, 225-260 cm) varies from 45 to 60 kg. The weight of the blubber in males constitutes roughly 60% of the total weight and in females impoverished during lac- tation 30-40% (L. Popov, 1961). The meat is used as feed in fur animal farms. To implement rational exploitation of the resources of the Jan Mayen hunting region and to ensure conservation at a stable level for a long time, agreements are imperative between the countries involved in hunting the hooded seal. Efforts in the field of research should be directed primarily toward an accurate census of the pups of the hooded seal; a comprehensive study of its ecology, including intensity of reproduction; migration char- acteristics; degree of isolation of the Jan Mayen herd from the western Atlantic (Canadian) populations; etc. The major task is to establish ratio- nal norms of permissible kill to prevent the exhaustion of hooded seal resources. Some measures restricting hunting have already been implemented in Norway and the USSR. These measures provide: (1) restricted hunt- ing in the breeding season in the Jan Mayen region (commencement of hunting not earlier than March 20 and its cessation not later than 547 May 5); (2) banning the reentry of a ship for hunting in the same sea- son; (3) banning hunting in Denmark Strait; and (4) banning hunting in summer оп drifting ice floes north of Jan Mayen Island. (K.Ch.) 411 ORDER OF WHALES Order CETACEA Brisson, 1762 413 COHORT OF WHALES COHORT Mutica ORDER OF WHALES Order CETACEA Brisson, 1762 These are highly specialized mammals, totally adapted to aquatic life. Dimensions vary from moderate to large and these animals are the largest among mammals. Body length and weight range from 1.1 m and 30 to 300 kg in some large dolphins to 33 m and 100 to 120 tons in blue whales (Fig. 213). The body is streamlined, spindle-shaped or teardrop-shaped, elon- gated or somewhat shortened: Such a shape offers the least resistance to swimming in water (Fig. 214). In common dolphins the body is better streamlined than in such fast-swimming fish as gray mullets and boni- tos (Shuleikin, Luk’yanova, and Stas’, 1937). Resistance to swimming in water is also reduced by the reduction of ear pinnae and the placement of mammary glands and teats in special pouches in females and the penis in a special slit in males. The forelimbs are modified in the form of fin-shaped flippers, which essentially serve as rudders. The hind limbs are totally reduced. At the end of the laterally flattened caudal crest there are paired horizontal caudal flukes without a skeleton. The caudal section of the trunk serves as the main locomotory organ. In most species an unpaired dorsal fin without a skeleton performs the role of a stabilizer while swimming. The head is often massive, sometimes about one-fifth of the body length, and is usually fairly elongated. It terminates obtusely or acutely, or has an extended rostrum, i.e., the so-called “beak”. The head in almost all these animals merges without a visible neck into the trunk, which gradually narrows with no sharp boundaries into the caudal crest. Whales do not have a compact hair coat. Baleen whales have some stray bristles on the snout, typically structured like the whiskers of land mammals. These hairs evidently play a specific role in searching for 414 Fig. 213. Comparative size of whales (figure by М.М. Kondakov). A—toothed whales: 1—common dolphin, 2—pilot whale, 3—killer whale, 4—Baird’s beaked whale, 5—female sperm whale, 6—male sperm whale; B—baleen whales: 7—blue whale, 8—fin whale, 9—sei whale, 10—humpback whale, 11—Minke whale, 12—gray whale, 13—bowhead whale. massive plankton collections (up to 400 nerve tips present in the follicle of one such hair). In the adult state, among the toothed whales, only the river dolphins possess whiskers: the Ganges River dolphin—Platanista gangetica and the Amazon dolphin—Jnia geoffrensis living in muddy river water. In most of the other toothed whales (except for the white whale and narwhal), whiskers are present only in the embryo. 414 Cutaneous glands are lacking. The skin consists of an epidermis, der- mis, and a very thick layer of subcutaneous fatty cellular tissue [blubber] with no distinct demarcation between the last two layers (Fig. 215). The 553 415 Fig. 214. Characteristic body form of whales (figure by М.М. Kondakov). 1—right whale; 2 and 3—rorquals; 4 and 5—dolphins; 6—white whale; 7 and 8—sperm whales. 554 ) ме eR S = SN 416 Fig. 215. Structure of the skin of whales according to У.Е. Sokolov. 1—epidermis; 2—dermal papillae; 3—dermis; 4—subcutaneous fatty cellular tissue; S-—subcutaneous musculature; 6—bundles of collagen fibers; 7—fat cells. epidermis in the body sections exposed to the maximum resistance of water while swimming (anterior margins of the fins and anterior part of the head) attains maximum thickness. The inner surface of the epidermis has numerous cells in which dermal outgrowths (papillae) grow vertically upward toward the kin surface. The pigmentation of the epidermis deter- mines the body color of whales. At the boundary of the epidermis and dermis, pigment cells—melanophores!—are visible. The color of some ‘Numerous white or weakly pigmented spots (scars) of varying size are usually found on the skin of whales. These are scars from bites during fights, attacks by sea lamprey, parasitization by some Protozoa, crustaceans, etc. 555 cetaceans has been correlated with age-related variations (for example, among white whales and narwhals). The dermis consists of a thin layer of 416 dense intertwined bundles of collagenous fibers and papillae rising from this layer into the epidermal cells. In the subcutaneous fatty cellular tis- sue, Straight fascicles of collagenous fibers are considerably larger than in the dermis and widely separated from each other, and the entire space between them is filled with accumulations of fat cells. The thickness of the subcutaneous cellular tissue varies in different seasons: for example, among female fin whales, it is 7.8 cm in December and grows to 12.7 cm by March (Slijper, 1936). The peculiar structure of the skin of whales evidently promotes development of a laminar flow of water around the swimming animal, with a minimum expenditure of energy when developing high speed. The caudal flukes (Fig. 216) under the skin cover consist of three lay- ers: one deep and two superficial. The superficial layers are formed of the tendons of m. longissimus and m. hypaxialis lateralis? (Narkhov, 1937) and spread fanlike into the flukes (Felts, 1966). The deep layer consists С р Е 417 Fig. 216. Figure showing structure of the caudal flukes in various species of whales (figure by N.N. Kondakov). A—Atlantic right whale; B—gray whale; C—fin whale; D—common dolphin; E—sperm whale. 2 According to A.V. Yablokov (1959), the tendons of these muscles do not reach the caudal flukes. 417 418 556 of a fairly dense bunch of large bundles of collagen fibers between which groups of fat cells occur. The caudal flukes are capable of minor indepen- dent movements (Zenkovich, 1952). The internal structure of the dorsal fin is similar to that of the deep layer of the caudal flukes. In the fins, along with normal arteries and veins, complex vessels exclusively typical of whales are seen. Each of these vessels consists of a large artery with a very thick middle layer (tunica media) surrounded by 10 to 13 very fine thin-walled veins. The base of the dorsal fin has more than 20 such ves- sels, which are fewer in number in the caudal flukes and flippers (Tomilin, 1957). These typical vessels are seen in the body parts but differ in struc- ture; their formation is evidently determined by functional requirements. The elasticity of the caudal flukes can evidently vary during the move- ment of the animal due to blood pressure controlled by complex vessels and the general distribution system present in the principal supply vessel (Pershin et al., 1970). It is possible that the effect of hydroelasticity in the caudal flukes has a bearing on the swimming of whales. Skeletal bones essentially have a typical spongy structure and con- tain a large amount of fat. Clavicles are lacking (Fig. 217). The sternum is highly variable in size, shape, and articulation, depending on the num- ber of ribs attached. The scapula is broad, flabellate [fan-shaped], and has a poorly developed crest. The humerus is extremely reduced. The radius and ulna are also highly reduced, flattened, and greatly broadened (Fig. 218). Flattening of the limb bones is evidently associated with the limb performing a typically new function similar to that of the aileron of an airplane wing. In view of the fact that the load on the limb is per- pendicular to it, the limb is flattened in the same direction (Druzhinin, 1924; Yablokov, 1959). All the bones of the free limb are firmly inter- connected and sometimes even fused, and with a common integument. Only the shoulder joint is movably articulate. Nevertheless, antebrachial muscles are preserved in the majority of whales. More often than among v 3 BAB Е 56 чопрдурорие оч“ * ^ Fig. 217. Skeleton and body contour of the common dolphin, Delphinus delphis (figure by N.N. Kondakov). 418 557 Fig. 218. Skeleton of the limbs of whales (schematic). A—bowhead whale; B—fin whale; C—blue whale; D—common dolphin (figure by N.N. Kondakov). other mammals, the fusion of the carpals is noticed. The number of digits is four or five. The number of phalanges in the middle digits is more and in the outer ones less. The number of phalanges in the embryo is more compared to the adult, 1.е., they reduce in number during the course of ontogenesis (Kiikenthal, 1889; Yablokov, 1959). The hind limbs form in a fairly early embryonic stage but soon dis- appear. Their rudiments are very rarely preserved. Among adult whales, only the rudiments of the pelvis arranged in the musculature, in the form of two bony shafts (fused ilium, ischium, and pubis), are preserved - of the hind limbs. Their articulation with the vertebral column has dis- appeared but the muscles of the urogenital system are attached to them and, obviously for this reason, their total reduction has not taken place (Yablokov, 1959). The vertebral column consists of only four sections: cervical, tho- racic, lumbar, and caudal. Sacral vertebrae are lacking due to the disap- pearance of the hind limbs. The cervical section is represented by seven highly reduced vertebrae which may be fused into a single unit or sev- eral groups. In the rest of the sections, the number of vertebrae varies. Transverse and spinous processes attain large proportions. Hemal arches, similar to those in the lower vertebrates, appear in the caudal section. Е ИЕ = = — И: 2727 ПРЕ. 2 ge i PN 419 418 419 S\N \ \ He WN м PO ро М Ny Wi у Bai № oN -\ а N 4 Sins а \) RTS Sas т.е Fa \ \ К Fig. 219. Rudiments of the hind limbs of the sperm whale. A—outer view, В and C—X-ray pictures (figure by М.М. Kondakov, after А.А. Berzin, 1970). The number of ribs varies from 10 to 17 pairs. The anterior two to eight pairs are normally joined to the vertebrae; the posterior ribs have neither capitulum nor collum and are articulated only to transverse verte- bral processes. The anterior eight pairs of ribs are joined to the sternum. The thoracic cage, unlike in most land mammals, is not laterally com- pressed but round in cross section or slightly flattened dorsoventrally. The bones in the skull of whales are arranged symmetrically or asym- metrically. In the latter case, the bones on the right side are broad- ened and shifted leftward while the left ones are thickened. Some bones may overlap others. The cranial section is shortened and rounded. The supraoccipital bone is highly developed and joined with the frontal bone anteriorly while the parietal bones are shifted laterally. The nasal bones are small and the outer bony apertures are shifted far backward and lie directly anterior to the neurocranium. The nasal passages are short and run fairly vertically. Maxillary and premaxillary bones and the vomer are highly extended and form a rostrum. The bony palate is elongated due to the large pterygoid bones, as a result of which the choanae appear displaced backward. The mandibles are long and have a rudimentary coronoid process, which is sometimes lacking. In the adult state, only the members of the suborder of toothed whales possess teeth, which are homodont and serve only to hold and kill the prey. In the embryo of baleen whales, teeth begin to form but soon 420 959 cease to grow and are resorbed. Among adult baleen whales, the teeth are functionally replaced by special, characteristic horny formations of the palatal epithelium, the so-called “whalebone” [baleen] which serves to strain the tiny planktonic organisms on which these whales feed. The triangular plates of the whalebone are fixed along the edge of the upper jaw parallel to each other in such a way that the smaller side of the triangle falls toward the upper jaw while the larger side is turned out- ward and the hypotenuse is inside the oral cavity. The inner part of the plate of the whalebone is split into innumerable fine and long keratinous filaments. The baleen plates are slightly separated from each other and the filaments of the neighboring plates are intermeshed to form a gauze- like structure. When the whale swims with its mouth open, water passes through this structure continuously while the planktonic organisms are retained on its surface. By closing its mouth, the whale pushes out the water from the oral cavity with its tongue, which also helps to push the food particles into the esophagus: whales do not masticate food. The tongue is well developed and soft lips are absent. Salivary glands are either altogether lacking or are rudimentary. The stomach is complex and consists of three to fourteen sections (Fig. 220). This complexity is explained by the absence of food mastica- tion. The length of the intestine varies: it may be five or six times the body length as, for example, among baleen whales (Mysticeti) or among species of the family of beaked whales (Ziphiidae); 15 to 16 times among sperm whales and bottlenose dolphins; and up to 32 times among La Plata dolphins (Pontoporia blainivillei). The liver is relatively small and a gall bladder is absent. The pancreas is elongated and faintly lobular, far more rarely disjunct; its ducts open into the bile duct. External nares (blowhole) are paired among baleen whales and unpaired among toothed whales, shifted to the top of the head, and have valves that close them during diving. Among the toothed whales, the nasal passage above the skull has special air sacs. Elongated laryngeal cartilages of toothed whales enter the choanae and thus maintain a constant contact between the larynx and the air sacs located above without interruption even during swallowing. The trachea and bronchi are shortened. The short passages transporting air accelerate respiration. The lungs are single-lobed with highly developed smooth muscles. The bronchioles have a series of smooth muscle sphincters (among toothed whales) or contain a large number of muscle fibers in the upper portion of the alveolar septa (among baleen whales). The connective tissue of the lungs is highly elastic. Lymphoid tissue is almost totally absent. The number of alveoli is relatively large and their dimensions somewhat larger than among land mammals. The epithelium of the bronchi and 560 420 Fig. 220. Internal organs of whales (figure by М.М. Kondakov). 1—lungs; 2—heart; 3—diaphragm; 4—stomach; 5—Пуег; 6—intestine; 7—kidney; 8—urinary bladder. 421 561 glands is almost devoid of mucous cells, which is evidently associated with the absence of dust in the air intake (ЗШрет, 1958a). Alveolar septa contain two layers of capillaries and a very large amount of clastic fibers. All the bronchi have a ciliate epithelium and are surrounded externally by cartilage of irregular shape. These cartilages are also seen in the terminal bronchioles (not found among beaked whales, Berardius). Whales can survive without breathing for quite sometime and remain under water from 2-10 to 30-40 min (up to two hours, according to some authors). They are insensitive to large amounts of carbon dioxide in the blood; its accumulation does not regulate their respiration, as in land mammals, but an oxygen deficit in the blood does. The long duration of diving is ensured by the large capacity of the lungs, insensitivity of the respiratory center of the brain to the accumu- lation of carbon dioxide in the body, high content of myoglobin, and slightly higher oxygen-holding capacity of the blood due to its higher hemoglobin content and its high concentration of erythrocytes (Irving, 1938; Taarra, 1950; Korzhuev and Bulatova, 1952; Kleinenberg, 1956a, 1957). Hemoglobin content in 100 ml of blood, g . Sei whale 15.6 Sperm whale 15.8 Cow 12.4 Man 13.0 Respiration in whales can usually be divided into several stages: (1).exhalation after prolonged diving; (2) intermediate brief inhalation and exhalation; and (3) deep inhalation before prolonged diving. Dur- ing intermediate inhalation and exhalation, the whale does not dive very deep, surfaces almost always vertically, and breathes at regular intervals. The number of intermediate inhalations/exhalations varies among differ- ent species; the longer the animal remains under water during diving, the greater the number. There is a particular reflex movement during inhalation: the animal strikes the water with its caudal fin in an up and down movement which invariably keeps the top of the head with the blowhole afloat (Tomilin, 1957). Exhalation can commence even under water, as a result of which the air expired under pressure forms a water jet that is generally characteristic in shape and size for various species.? 3 Apparently, the jet or fountain may also be formed as a result of the condensed vapor present in the warm air expired (in high latitudes) and also as a result of holding and spraying of the water which remains, as the whale surfaces, in the cavity in which the blowhole is located. 422 562 Some whales are capable of diving to great depths (Юг example, sperm whales can sink deeper than 1,000 meters). The quick surfacing of the animal from a great depth should liberate the gaseous nitrogen dissolved in the blood under the high pressure generated during diving. Otherwise the blood vessels would become clogged with nitrogen bub- bles, leading to caisson disease and even to the death of the animal. But whales do not suffer from this disease. This is probably explained by the fact that the amount of nitrogen present in the lungs slightly exceeds the nitrogen-holding capacity of the tissues while no more air enters the lungs during diving (unlike what happens in divers). The brain in most respects is highly differentiated but in many other respects has preserved extremely primitive features that are not found among the members of other orders of mammals (Fig. 221). These differ- ences are mainly: significantly narrow cortex in cross section while layers Гапа V-VI, the oldest in evolution, are predominantly developed; overall high cellular content of the cortical plate; and developmental features and ratios of the zones of old and intermediate cortex (Filimonov, 1949). The weight of the brain in absolute terms is maximum in mammals but is insignificant in relation to the body weight (especially among large whales). The brain of white whales weighs 2,180 to 2,340 g (Kleinenberg et al., 1964), fin whales 6.5 to 7.2 kg, and sperm whales 7 to 8 kg (Kojima, 1951; Janes, 1952). Large convolutions and longitudinal fissures are seen in the short rounded form of the brain. Intense development of the cortex of the hemispheres of the forebrain is a characteristic feature. Without exception, the cortex houses the paleocortical, archicortical, and inter- stitial zones (Filimonov, 1949). The cerebellum attains large dimensions (up to one-fifth of the weight of the entire brain). The hypophysis is highly developed. Fig. 221. Brain of the common dolphin, Delphinus delphis (figure by N.N. Kondakov). 422 563 The structural peculiarities of the cetacean brain are associated with the high degree of development of the higher nervous activity of these animals. Many species of whales are easily tamed and trained. With respect to higher nervous activity, some whales (for example, the bot- tlenose dolphin) may be placed among the higher mammals although lower than the anthropoid apes (Voronin, 1970). Reduction of the rhinencephalon is a characteristic feature. The bul- bus olfactorius is lacking in most whales (Kiikenthal and Ziehen, 1893)4 and the lobus hippocampi are poorly developed. The V and VIII pairs of cranial nerves associated with the sensitivity of the facial region and auditory organs are highly developed. Among bottlenose dolphins (Tur- slops truncatus), the following features of the brain have been noted (Kruger, 1959): a prominent flexure between the mesencephalon and the diencephalon; reduction of tactile thalamic region, evidently asso- ciated with the absence of hairs in adult animals (this is true of most toothed whales); intense development of n. ventralis medialis, definitely suggesting that whales have a sense of taste (although many investiga- tors refute this); and so forth. Amony some toothed whales (narwhal, pilot whales, white whales, and common propoises), a small number of typical pits with minute processes at the bottom have been noticed on the surface of the root of the tongue. They evidently function as organs of chemical perception (Yablokov, 1957, 1961a); taste buds are found in them (Yablokov et al., 1972) as well as many efferent ducts of the albumen and mucous glands. The eyes are small (Fig. 222). The cornea and sclera attain consider- able thickness and the optic muscles are well developed. The crystalline lens has a characteristic spherical form. Eyelids are not developed. Vision is evidently monocular and myopia is typical (Kellogg, 1928; and others). The lachrymal glands are reduced and the nasolachrymal passage is lack- ing. An oily secretion from Harder’s gland protects the eye from the mechanical and chemical action of water. Conjunctival glands, not found among other mammals, are present (Weber, 1886). The olfactory organs and Jacobson’s organ are reduced. Sense organs among baleen whales and some toothed whales (for example, the Amazon dolphins, Га) are represented by vibrissae dis- posed on the snout and sometimes all over the trunk (Fig. 223). Numer- ous nerve ends approach the roots of the vibrissae surrounded by blood lacunae (Japha, 1910). Among all whales, the sensory nerve endings rise 4 The bulbus olfactorius has been described (Flatau and Jacobson, 1899; Filimonov, 1949) in the bottlenose whale, Hyperoodon rostratus, the blue whale, Balaenoptera musculus, and the fin whale, B. physalus. 422 423 564 Fig. 222. Eye of the fin whale, Balaenoptera physalus (figure Бу М.М. Kondakov). along the dermal papillae almost directly at the outer surface of the skin so that the entire surface of the body appears well innervated. The auditory organs are highly modified (Fig. 224). The concha is reduced. The external auditory meatus opens outward behind the eye as a tiny aperture; its lumen is invariably covered. It is possible that the rudi- mentary auditory meatus may serve as an independent organ perceiving pressure changes (Yamada, 1953). The tympanic membrane bends out- ward (among baleen whales) or inward (among toothed whales), and is externally covered among baleen whales by a typical ear plug of ker- atinous epithelium and сегатеп.> In view of the fact that the petrous tympanicum is not only isolated from the other skull bones, but also sur- rounded by special air cavities, it is possible that whales perceive sound signals through the auditory meatus. It has been suggested that sound is perceived through the body surface (especially the head), transmitted to the auditory zone, and that the petrous tympanicum, which is capable of vibrations, acts as a resonator. It has also been suggested that the lower jaw is also capable of perceiving sound from the external environment and transmitting it to the auditory organs. Whales are capable of receiving a wide range of sound waves: from 150 to 120-140 thousand Hertz (Kellog, 1953; Slijper, 1955, 1960; Bullock et al., 1968), i.e., even ultrasonic vibrations. The high degree of development of the auditory section of the brain of toothed whales points to their particularly acute sense of hearing, which is almost unique among mammals. Among baleen whales, hearing is inferior to that among toothed whales but better than that among land mammals (Ogawa and > The age of whales can be determined by counting the layers in a slice of this plug (Purves, 1955). 565 423 Fig. 223. Vibrissae of whales. Top to bottom: Atlantic right whale, fin whale, and gray whale (figure by N.N. Kondakov). Arifuku, 1948). Whales, like bats, are capable of echolocation (Kellogg, 1958). Since vocal cords are lacking, whales cannot produce sound by the methods ordinarily used among mammals. However, that they do produce diverse sounds has long been known. Evidently they result from vibrations of the lower portion of the nasal septum (Tomilin, 1957) or vibrations of the folds of the outer valve due to passage of air from the dorsal nasal sacs (Yablokov, 1961b). It should be noted that dolphins have at least two systems of sound production: they are capable of simultaneous production of location signals and whistles. It has been 424 suggested that whistles are produced by means of the larynx; impulse 424 566 Fig. 224. Structure of the auditory organs in the fin whale, Balaenoptera physalus. Top—outer view; bottom—section (figure Бу М.М. Kondakov). 1—outer opening; 2—membrane; 3—tympanic membrane; 4—ligament between tympanic membrane and malleus; 5—ear plug; 6—auditory ossicles. signals are apparently produced by the air sacs (Evans and Prescott, 1962; Norris, 1964). The typical structure of the frontal section of the skull and the disposition of the fatty “lens” anterior to it enable the animal to judge the direction of sound waves (Romanenko et al., 1965). The blood circulatory system of whales has several characteristic features, some of which are also encountered in other aquatic and ter- restrial mammals (Slijper, 1959): (1) arterial rete mirabile is present on the inner side of the vertebral column and particularly well developed in the regions of the neck, thorax, between the ribs, at the base of the brain, and around the spinal cord. It is formed of a large number of sinu- ous arteries of the muscular type interconnected by several anastomoses. The histological structure of the vessels points to their capacity to modify considerably the volume of blood flowing in them; (2) the venous rate mirabile is present at the base of the skull and is particularly massive in the abdominal zone; (3) an expansion occurs in the hepatic vein (lack- ing in baleen whales); (4) two vertebral veins running along the ventral side of the vertebral column are almost identical in diameter throughout their length and lack valves, indicating that blood can flow in them in both directions: through the major уу. costocervicales they run into the 425 567 anterior vena cava and through the veins between the ribs into the pos- terior vena cava. Since all these features of the blood circulatory system attain maximum development among smaller whales, they are evidently associated not with deep and prolonged diving of the animals, but with frequency of diving and respiration. A reference has already been made to the veinous vessels of fins typical of whales (arteries surrounded by small veins). In addition to the very high (compared to terrestrial mam- mals) oxygen capacity of the blood, the considerable concentration of sugar and sodium chloride in it is a characteristic feature of whales. This is evidently associated with the deposition of additional reserves of food required by the animal during prolonged underwater residence. The diameter of erythrocytes in whales varies from 6.6 to 10.0 microns (Lenfant, 1969). The body temperature of whales is similar to that of land mam- mals and varies from 35 to 40°C (the upper limit was recorded among wounded whales or dolphins caught after chasing). The maintenance of a high body temperature in water, which conducts heat many times better than air, is facilitated by the thick layer of subcutaneous adipose tissue. Killed whales 60 h after the moment of death and in cold water recorded a temperature close to 30°C®, 1.е., only 6-7°C loss of heat during this time (Zenkovich, 1938). The temperature of the outer layers of the skin among whales is close to the temperature of the surrounding water. This is achieved by minimal heat dissipation by the body as required for nor- mal activity in such a highly heat-conducting medium as water. In the case of over-heating of the whale body (for example, on rapid and prolonged swimming), dissipation of heat occurs from the entire body surface but more intensely from the surface of the fins. Compared to land mammals, the musculature of the trunk and limbs has undergone the most intense modification. As a result of losing the locomotory function, the following muscles in the forelimbs are very weak: m. romboideus, m. serratus, m. levator scapulae, and m. infraspinatus; m. trapezius and m. teres minor are totally reduced; and m. deltoideus, m. subscapularis, and m. paniculus carnosus are highly developed. In the trunk, m. latissimus dorsi and m. ileo-costalis passing through the dorsal side of the vertebral column as also m. hypaxialis lateralis and m. hypaxialis medialis passing along its ventral side are particularly well developed. The former are attached anteriorly to the skull and the latter to the lateral processes of the posterior thoracic 6 The high body temperature of the killed whales may also have been partly main- tained by the process of putrefaction; nevertheless, this fact points to the considerable heat-insulating capacity of the subcutaneous adipose tissue. 427 568 vertebrae and to the last of the ribs; posteriorly, these muscles are attached to the caudal vertebrae and (Narkhov, 1937) partly transit into massive tendons running into the flukes and diverging fanlike there. The kidney of whales is multilobulate and relatively much larger than in land mammals (Slijper, 1958a). The urinary bladder is small; the absence of a sphincter in its neck facilitates frequent urination in small quantities which possibly may serve as a signal for other whales (Yablokov, 1961a). Whales evidently do not swallow sea water and the required moisture is provided by the food intake in addition to the water of metabolism. The absorption capacity of the kidneys of whales is not so high as to extract fresh water from the swallowed sea water. The concentration of sea water was found to be more than that of the urine (if whales swallowed sea water, this ratio would be reversed) (Table 30). The testes are located in the abdominal cavity. In blue whales, the length of the testes may reach 45 cm and the weight 45 kg (Slijper, 1966). Of the other glands, only the prostate is present. The penis is long and narrows gradually toward the tip; at rest, it lies in a special sheath. At its base, the penis bends in an S-shape. An os penis is lacking except in the right whale, Balaena glacialis. The cavernous tissue is poorly developed in the penis. The crus of the crura penis are attached to the rudimen- tary pelvic girdle; here, mm. bulbo- and ischiocavernosi are separated. Among adult females, the ovaries are partly or wholly surrounded by an extensive infundibulum of the oviduct. The oviducts extend into the bicornuate uterus, which has longitudinal folds on the inner surface. The portio vaginalis enters the vagina. The outer slitlike genital opening is surrounded by small liplike bulges between which the boneless clitoris is located. Ureters open into the vulvar region; there is no sinus urogeni- talis. The vagina contains several ringlike folds which probably perform the role of valves, protecting the genital system from water entering into it during copulation and whelping. In female baleen whales, the genital Opening and the anus are separated from each other by a considerable distance while in toothed whales, these are located in a common Sac and are surrounded by a common sphincter (Fig. 225). The males are capable of fertilizing throughout or for much of the year. Ovulation among whales is perhaps stimulated by copulation (Sokolov, 1950, 1954; Sleptsov, 1952; Tomilin, 1957; Yablokov, 1959) but the recovery of spermatozoa in the vagina of female common dolphins having mature but not opened-up follicles in the ovaries casts doubts on this assumption (Sokolov, 1961). On fertilization, the corpus luteum of pregnancy develops in the ovary and, after resorption, traces of it are preserved for a long time (presumably throughout life) in the form of a corpus albicans. The age of the female can possibly be determined by 569 — КИГИ / “LD, 77772 22222772 2 ly РРР Е: EE LLL РРР ИРИ 77 РРСРР. й РР РРР й ат - 4% Z Os SEES рр. Я. - 27 jp eee a ——<— Segoe eon ee oe te IP ie CLE 1 2 A “ 7) РРР Я ПАР Ре РОРЕРРРРРРР , ИР 2777; GLEE Ze 22222 i ne. We ИИ SOLIS РРР Ce; 72 АА: РР = й 4 di a VE a LEED J РЕ i << cere lw = === He НИЕ Е Liza > Zig — = <->» oe = Е пе 22 В 427 Fig. 225. Zone of disposition of the urogenital organs in the fin whale, Balaenoptera physalus. A—male; B—female (figure by N.N. Kondakov). 1—genital opening; 2—anal opening; 3—sac for mammary teats. counting these bodies. Among pregnant females, two or three fetuses can be found in the uterus at the commencement of pregnancy but shortly thereafter only one is found. The placenta is diffuse, of the epithelio- choreal type (Zhemkova, 1965), and may partly extend into the second horn of the uterus. In the late stages of pregnancy, the embryo is usually situated with the tail toward the womb exit. The lobes of the flukes are convoluted, the dorsal fin (when present) bent down towards the back, and the flippers turned towards the tail. The sex ratio of embryos is roughly 1:1 (Table 31). Birth takes place under water. The calf is born fully developed and capable of independent movement; its body proportions are highly sim- ilar to those of adults (Zemskii, 1958a) but its length up to 1/4 to 1/2 that of the mother. Some female whales can be fertilized soon after par- turition, during the lactation period. The calves are suckled under water, the duration of each suckling being short (a few seconds). The calf holds the teat of the mother (Fig. 226) between its tongue and upper palate. 428 An exception is the sperm whale calf, which holds the teat in the corner of its mouth (Slijper, 1966). Milk is sprayed into the oral cavity of the calf by the contraction of special muscles in the female. Newborn calves feed very frequently; for example, calves of bottlenose dolphins suckle 570 one TTS 6 LY Bes (On) СИТ STS pss a4 Crs ССР OLY STS a3e]U9019g $цэци9э@$ 179 ILS 887 Lvz LI 61 79 05 6 18 LSI PLT Jo лэашам o[RUIS.J SIN этешэ ae a[ewi3y ae эгешэН эти эешэ Te o]ewsy aye [е1оТ, ЭТЕЧАА 11э4$ эгеЧчм yoeqduinyy ЭГеЧм эп[а ЭТЕЧА 19S ЭГеЧм и злетоплтеа (ISs6T ‘озелай рце эп21 {0661 ‘одип[ рие anziy) зоАлашо э[еЧм Зиоше OBI хэб “TE э[ает, = LE Ors bs от 8'6 06? = ЕСС тотем BIS bE 6'5 6$ 1Z 81 = == OLS ETE if STEM SITY A = = ObZ-OLT от OL = = 078-08? — 15-855 с urydjoq = = O8T = = = == 08$ 10°Z т этечм зопа 19 = Ove = = = == OZ 095 I yey onig 1-64 0-1 065-055 9'9 ЕЕ [9 0/5 057-085 = t эГечл uly ТУТ = 0/5-09%% 85-6966 6531 CL-9S 055-056 05%-055 == с ayeym uly Iy-9T 6-95 OLE-O81T TPr-9T Z-€0 58-9 06Z-OFZ 055-088 9r'7 Ф ayeym 125 LCZEG= ВСЕ Т 10/9095 = oe == = 059-05 = ф этеЧл 125 1Z 8$ OLE = 6 ps 05$ 00$ 67< I этецм 19g ’Od 705 5) ЗИ i) Я ем 6211] Do suoneuIwajap oTeUM Jo} [/WIu ‘и0ел11э940) jo Joquinyy (6€61 ‘1эЦэуэЯ) sofeyA Jo autin ay} Jo ион 15040) ‘0$ тает, 9th 9¢L 427 429 ЙЕ Fig. 226. Teat of a lactating female fin whale, Balaenoptera physalus (figure by N.N. Kondakov). roughly once every 26 min day and night (McBride and Kritzler, 1951). The mammary glands of the female lie along the sides of the genital opening. Two teats (one on each side) lie in slitlike folds and project outward only during lactation. Female whales produce a large amount of milk daily: from 200 to 1200 g among dolphins, up to 90 to 150 liters among fin whales, and 200 liters among blue whales (Sleptsov, 1955). The milk is thick and usually cream-colored. Its surface tension is 30 times more than that of water, which is particularly important consider- ing that suckling takes place under water (the milk spray does not spread on water). The nutritive value of whale milk is very high (Tables 32, 33, and 34). A different composition of the milk of fin whales has been given by Ota et al. (1955) (Table 33). During the period of suckling the growth of calves proceeds very rapidly: for example, by seven months of age the calf of a blue whale had grown to 16 m from 7 м at birth, 1.е., the average daily increment of length was 4.5 cm (Slijper, 1960). Sexual dimorphism among whales is manifest mainly in the differ- ences of body length between males and females. Among baleen whales, the females are larger than males while the converse is true in most toothed whales. Whales are mainly herding animals and live in schools of a few animals to hundreds and thousands. They are encountered 428 428 428 572 Table 32. Composition of the milk of large whales (2, averages) (Zenkovich, 1952) Species Fat Dry residue Water Fin whale 43.57 13.50 42.93 Blue whale 40.25 13.70 46.05 Humpback whale 39.93 14.02 46.05 Gray whale 53.04 6.38 40.58 Sperm whale 37.30 8.00 54.70 Cow 3 РА TUS) Table 33 Fat Protein Water Lactose Inorganic matter 33.0 13.3 53.4 0.3 1.0 31.8 12.3 55.0 0.2 0.7 32.5 10.5 54.1 1.4 1.4 Table 34. Composition of the milk of small whales (2) (Ural’skaya, 1957) Species Fat Protein Sugar Dry Ash Water residue Common dolphin 43.71 5.62 1.45 7.53 0.45 48.76 Common dolphin 41.56 4.88 1.49 6.82 0.45 51.62 Common porpoise 45.80 11.19 1633 13.09 0.57 41.11 Common porpoise 33.90 5.22 1.28 7.10 0.60 59.00 Bottlenose dolphin 46.10 11:55 1.5 13.50; 0.38 40.50 along coasts as well as in the open sea. Some are capable of ascend- ing along major rivers opening into the sea while some species are regularly confined to rivers. The food of many is specialized; plank- tophagous, teuthophagous, ichthyophagous, and sarcophagous animals are known (Tomilin, 1957). Whales include fast swimmers (for example, killer whales and many dolphins) and relatively slow-moving animals (gray whales). Most whales regularly remain on the surface while some (for example, sperm whale) can dive to great depths. The population of the various species of whales is not identical. Some whales are most numerous and can be found in thousands of schools (common dolphin) while others are very rare and only a few sightings have been reported (some members of the genus Mesoplodon 573 (beaked whales) dwarf sperm whale, and false killer whale). Natural fluc- tuations in population have not been studied but can hardly be signifi- cant. As a result of senseless hunting, the population of many species of whales has sharply decreased. Thus, right whales and gray whales have come close to total extinction. International conventions have imposed a total ban on the hunting of these whales and their population is now slowly recovering. The hunting of even sperm whales and fin whales, numerous in the recent past, has to be totally banned to maintain their population at the present level. A sharp reduction in whale hunting or its total ban for several years, followed by rational and strictly controlled hunting, should be considered. Periodic migrations are characteristic of many whales. Among some species, the distance covered during migration is relatively small (Azov- Black Sea common porpoise migrates from the Azov Sea into the Black Sea and back) while migrations among others cover long distances: some large whales travel from tropical waters to high latitudes. Most whales are monogamous. Periods of mating and whelping are usually prolonged. They give birth to one (rarely two) calves. The mate- rial instinct is strongly developed among females. Apart from man and the killer whale, there are no enemies of cetaceans. Cetaceans are universally infected with endo- and ectoparasites (crustaceans). The latter are particularly typical of large whales. Whales are found in all the oceans, in most of the world seas, and in some rivers and lakes. Two factors determine the distribution of whales: food and water temperature. Some species are widely distributed and are encountered in warm as well as cold seas (some species of the dolphin family), others have a small range (gray whales inhabit the subtropical moderate and cold waters of the North Pacific Ocean and the Chukchi Sea), still others are even more restricted (narwhals do not leave the Arctic waters), and finally, the ranges of river, lake, and estuary forms are altogether insignificant. At present, relatively few fossil remains of whales are known. Remains of the more primitive of the known whales have been found in the Middle Eocene of North Africa. There is no unanimity of opinion regarding the ancestors of whales. Some scientists support the origin of whales from ungulates, with which they share many common features, such as diffuse placenta, bicornuate uterus, and complex stomach. The number of chromosomes and the reaction of the precipitation of serum proteins of whales confirm this hypothesis (Makino, 1948; Boyden and Gemery, 1950). Based on the structural similarity of the skeleton and the dental system of extinct whales with the primitive carnivores (creodonts), 430 574 some scientists assume that the whales originated from the latter. А view has been expressed that the ancestors of whales were much older than creodonts, i.e., cretaceous insectivorous forms (Slijper, 1958a). Some authors (Beddard, 1900; Kukenthal, 1900; Hosokawa, 1950; Kleinenberg, 1958; Yablokov, 1964; Anderson and Jones, 1967) suggest a diphyletic origin of the contemporary order of cetaceans, i.e., that the baleen and toothed whales evolved from different ancestors and that their evolution proceeded by convergence and not divergence; hence it would be more correct to regard these suborders as orders. In confirmation of this hypothesis, numerous morphological differences between the baleen and toothed whales are cited but such evidence cannot be regarded as sufficiently convincing. The ancient whales, Archaeoceti, are sometimes regarded as conver- gent with other whales although they have nothing common in phylogeny and hence should not be included in the order of cetaceans (Yablokov, 1964). But even this assumption is not supported by adequately convinc- ing data (Mchedlidze, 1970). The systematics of Cetacea has not been properly worked out. As mentioned earlier, even the content of this order has been disputed by some. The composition of some families and subfamilies has also not been conclusively established. Usually (Simpson, 1945), three suborders are distinguished in the order Cetacea: modern toothed whales (Odon- toceti Flower), baleen whales (Mysticeti Flower), and extinct whales (Archaeoceti Flower). Sixteen families (nine are extinct) and 173 genera (of which 137 are fossil forms) are usually recognized in the order. The total number of contemporary species of the order is 81 (Hershkovitz, 1966). The USSR is host to 6 contemporary families, 19 genera (roughly 52% of the world fauna), and 23 species (roughly 28% of the world fauna) (Tomilin, 1957). The suborder Archaeoceti comprises three families: Protccetidae Stromer (Middle Miocene), Doridontidae Miller (Eocene and Miocene), and Basilosauridae Cope (Eocene and Oligocene), and 14 genera (Simpson, 1945). The family Protocetidae includes short-bodied forms such as Protocetus with a length of not more than 2.5 m and also long-bodied animals (in the available long bodied form of Eocetus, the skull alone measures 120 cm in length). Evidently, Eocetus was the source of the family Basilosauridae while Protocetus and Pappocetus gave rise to the members of Dorudontidae (Deshaseux, 1961). The family Dorudontidae comprises animals 8 to 9 m in length with an elongated and flexible cervical section and serrate cheek teeth while the family Basilosauridae includes animals of 8 to 22 m in length with some skeletal characteristics attaining a high degree of development within the suborder (Romer, 1966). 431 432 55 The economic importance of whales is quite significant although уаг- 105 representatives are not of equal importance in this respect. Some whales are caught in large numbers every year (fin whales and sperm whales) while others are hunted only at random. Valuable edible and commercial products are obtained from almost all organs of the whale. Oil is rendered from the skin and bones (commercial and edible). The meat contains 20 to 26% protein and can be a source of diverse pro- tein concentrates (frothing and emulsifying agents, substitutes for egg protein and deficient peptone for preparing nutrient media in microbi- ology) (Faingersh et al., 1953). The pancreas of whales can be used to produce insulin (1 kg of pancreas contains from 1,000 IU in the blue whale to 3,000 in the sperm whale; Egorova, 1953) and commercial pan- creatin (skin-softening agent; Bodrov et al., 1958); the liver can be used to produce vitamin A (1 g of liver contains from 400 IU in the hump- back whale to 5,800 in the sperm whale; Mrochkov, 1953), vitamin В, and campolon (Bodrov et al., 1958). The brain of whales can serve as raw material for producing cholesterol (frozen brains of fin whale, blue whale, and humpback whale contain roughly 2.5% or about 11% of it in terms of dry matter; Egorova and Lebedeva, 1953). The skin of whales, apart from providing fat, is useful as leather. The upper denser sections of the sperm whale are used to produce sole leather while the lower more porous ones are used as soft leathers (Zaikin, 1953). (V.S.) Key to Suborders of Cetacea 1 (2). Teeth absent. Numerous horny plates on upper jaw form straining apparatus. External nares bipartite. Skull symmetrical. Middle sec- tions of movable halves of lower jaw flexed outward. Only first pair of ribs joined to sternum. ... Suborder of baleen whales, Mysticeti. 2 (1). Teeth present. Straining apparatus absent. External nares not bipartite. Skull asymmetrical in frontal section. Middle sections of immovable halves of lower jaw straight or flexed inward. Not less than three pairs of ribs joined to sternum.................. уе ие 8 Suborder of toothed whales, Odontoceti (See below). SUBORDER OF TOOTHED WHALES Suborder ODONTOCETI Flower, 1867 Size small, medium, and large (body length up to 2 m, up to 4 to 6 m, and up to 21 m). The body is torpedo- or teardrop-shaped. 432 433 576 Fig. 227. Skull of the common dolphin, Delphinus delphis (figure by М. М. Kondakov). In the skin, the dermal layer and, in most cases, its fat-free portion are well distinguished. The network of elastin fibers is relatively poorly developed (except in small dolphins). A hair coat is lacking in most species. The body color is monochromatic, sometimes with typical white spots and bands. The skull bones (Fig. 227) are sharply asymmetric although, in the early stages of embryogeny, the skull is characterized by all features char- acteristic of the skull of land mammals (Sleptsov, 1940). But this similar- ity disappears very soon as a result of the intense growth of the supraoc- cipital and premaxillary bones and also the supraorbital processes of the frontal bones, and the skull of the embryo then resembles that of an adult. Uneven development of the skull bones is also seen in the late stages of embryonic growth and extends into the postembryonic stage (Sleptsov, 1940c). The reasons for the genesis of asymmetry have not been accurately ascertained thus far: some researchers suggest that the unequal pressure of water on different sections of the skull while swim- ming is probably responsible while others regard this asymmetry as a consequence of the reduction of olfactory nerves. The nasal bones are poorly developed and do not cover the posterior part of the bony nares. The nares are displaced leftward and open into a common chamber. Max- illae, premaxillae, and nasal bones are notably shifted onto the frontals and almost wholly cover them. Primitive features of structure are seen in the skull: presence of os parasphenoideum, tabularia, and postfrontalia,; the number of bony components in the skull of embryos is more than in juvenile and adult animals (Sleptsov, 1949c). Cervical vertebrae may be 433 577 fused. Among the fast-swimming species, the number of caudal vertebrae is more. Among the deep-diving animals, because of the reduced num- ber of ribs (in the sperm whale, their number decreases to three), the latter are joined to the sternum by flexible joints or attached by cartilage (beaked whales). The dental formula varies from о or : to roughly 2. The teeth are homodont and monophyodont. Three types of teeth are distinguished (Fig. 228) (Yablokov, 1958a): i Simple peg-shaped teeth with highly developed pulp cavity and thin cement and enamel layers among adult animals. This type of dental sys- tem is found in the common dolphin, common porpoise, pilot whale, etc. Except for the pilot whale, all these animals have a large number of teeth uniformly distributed in the jaws. . Teeth with highly developed layer of cement; enamel lacking on the teeth of adult animals; the tooth crown in the young animals has a thin layer of enamel (Lenberg, 1911). Teeth are simple, peg-shaped, larger than in the first type, and their number goes up to 30-50. The pulp cavity is well developed or absent. This tooth type is seen among sperm whales, white whale, apparently in dwarf sperm whale, Risso’s dolphin, dwarf killer whale, Irrawaddy River dolphin, and false killer whale. . Teeth are flat, wedge-shaped, with highly developed enamel layer and cement filling the pulp cavity and thus covering the entire tooth with the exception of the crown; further, the cement layer typically adheres to the enamel layer in the midportion of the tooth. The number of teeth is small and they occur only in the lower jaw. This type is characteristic of Baird’s beaked whale, beaked whales, and apparently the rest of the members of this family except for the Tasman beaked Fig. 228. Teeth of whales. A—common dolphin, Delphinus delphis; B—white whale, Delphinapterus leucas; C—Baird’s beaked whale, Berardius bairdi (figure by N.N. Kondakov). 434 578 whale. The teeth of the killer whale occupy an intermediate position between the first and second types. They are similar to the teeth of the first type but differ from them in their large dimensions, comparatively small pulp cavity, and large amount of dentine. In these respects, they resemble the teeth of the second type but, in contrast to them, have a poorly developed cement layer. In most of the toothed whales, the number of teeth is subject to indi- vidual variation. Among some species, teeth are fewer in the upper than in the lower jaw (Baird’s beaked whale, sperm whale, Risso’s dolphin, etc.); among others, on the contrary, the number of teeth in the upper and lower jaws is identical (narwhal, white whale, common porpoise, and black finless porpoise). Annual layers are deposited on the teeth and the age of toothed whales can be determined by counting these layers in a section of a tooth. A unique feature of some cetaceans is that the left (rarely the right) upper tooth has transformed into a tusk, as seen in the narwhal males (less commonly, in females also), reaching a length of 2.7 m. The digestive system among all the species is totally separated from the respiratory tract and has several characteristic features (Yablokov, 1958b). The tongue, unlike in land mammals, has a different structure and function. It is highly mobile and is covered with a thick sheath consisting of many layers of horny epithelium and a massive layer of connective tissue. The tongue positions the captured quarry in the oral cavity, pushes it into the gullet, and prevents water from entering. The soft palate is absent. The initial sections of the digestive tract are covered inside with many layers of horny epithelium, which evidently protects the tract from damage by hard portions of food swallowed whole. The multichambered muscular stomach (Fig. 229) among species feeding predominantly on cephalopods (sperm whale and Baird’s beaked whale) has a lining of glandular epithelium in the first section (Sleptsov, 1955). The subsequent sections of the stomach apparently take part in the process of absorbing the food; their structure is similar to that of the intestine. A cecum is lacking (seen in Platanista; Weber, 1927) and the intestinal sections are not distinctly separated from each other. The relative length of the intestine varies markedly from species to species. Ratio of length of intestine to body length (Yablokov, 1958b) Bottlenose whale 1:6.0 Sperm whale 1:15.0-17.5 435 433 579 Common dolphin 110451225 Common porpoise 1:12.2 13.8 Bottlenose dolphin 1:14.4- 15.8 Chinese finless porpoise 1:6.5 White whale 1:61 5951 The respiratory system too reveals several characteristic features. The external nares are not bipartite (Fig. 230) and are supported by dense dermomuscular folds, such that the projections of one fall into the depression of the other. Several special paired air sacs lie above the skull: the upper ones dorsally, the lower ones proximally, the rear ones nasofrontally with additional cavities, and the lateral ones enterolaterally (Kleinenberg and Yablokov, 1958). Their function consists in supporting the nasal passage while diving; the greater the pressure of water, the more intense their action. Moreover, they produce sounds. The larynx projects into the internal nares in the form of a tube formed by elongated arytenoid cartilages and the epiglottis (Fig. 231). This tube is covered by a special muscular sphincter. Bronchioles contain annular sphincters of smooth muscles (lacking in bottlenose whales). These are best developed in smaller species which dive often and thus breathe often and have a relatively large lung capacity. Apparently, the system of sphincters enables adaptation to pres- sure changes in the lungs during frequent dives and powerful inhala- tion/exhalation (Slijper, 1958a). 2 Lk 4 > А № Ху In? \ ie EMSS NAS \ Sy h АИ Ne ies SY ASA 1 S NSE yy SS << RSI 3 WS WEY REY Fig. 229. Structure of the stomach of toothed whales (figure by N.N. Kondakov). 1—esophagus inlet; 2—first section of stomach; 3—second section of stomach; 4—third section of stomach; S—sphincter; 6—duodenal ampule; 7—commencement of duodenum. 580 The brain has well-developed ventral nuclei in the auditory nerve, anterior Olives, etc. (Ogawa and Arifuku, 1948). The auditory organ dif- fers significantly from that of baleen whales in the shape of the tympanic bulla and the mode of its attachment to the skull (Yamada, 1953). Sexual dimorphism is manifest among many species and is most dis- tinctly seen in the dimensions of the animals: the males of most toothed whales are larger than the females. There are other differences too in some species. Thus, the dorsal fin among male killer whales is much higher than in females while a tusk is a feature of male narwhals (mainly). In the course of postnatal development, changes occur in body dimensions and its sections. Some species undergo color changes too (for example, white whales which are dark colored at birth turn lighter and become white with advancing age). > < < < 3 ~ СХ А В Fig. 230. Unpaired blowhole among toothed whales, viewed from above (figure by N.N. Kondakov). A—bottlenose whale; B—sperm whale. Fig. 231. Structure of the upper respiratory passages (figure by N.N. Kondakov). 436 581 Almost all toothed whales are fast swimmers, which has а bearing on their food characteristics. Some are capable of deep and prolonged submergence. They live on the coasts as well as in the open sea. Some are found in rivers and species of the family of river dolphins, Susuidae, regularly inhabit large rivers (Ganges and the Amazon). All the toothed whales, unlike baleen whales, swallow prey entire. Depending on the predominant food, several adaptive groups can be distinguished (Tomilin, 1957). 1. Ichthyophagous whales, feeding mainly on pelagic schools of fish. Their rostrum is long and narrow and the teeth numerous and covered with enamel. The common dolphin and dolphins of the genus Stenella belong to this group. 2. Bentho-ichthyophagous whales consuming mainly bottom- dwelling fish and invertebrates. The rostrum is considerably shortened and the number of teeth less. Common porpoise, white dolphin, bottlenose dolphin, etc. belong to this group. 3. Teuthophagous whales, feeding mainly on cephalopods. These have a small number of teeth but none at all on the upper jaw. Sperm whale, beaked whales, bottlenose whale, etc. belong to this group. 4. Teutho-ichthyophagous whales (pilot whale and false killer whale), consuming cephalopods and fish. They occupy an intermediate position between the second and third groups. The rostrum is reduced in length, but broadened because of the premaxillae. Although the number of teeth is less, they are well developed. 5. Sarcophagous whales (killer whales), along with fish also feed on large marine mammals. They have short but massive jaws with few but very strong teeth. Toothed whales are distributed in all the oceans, in almost all the open seas of the world, and in some rivers and lakes. Among the primitive Odontoceti of family Agrophiidae Abel, 1913 (Upper Eocene), the early stage of bone extension in the skull has been observed: the maxillae began extending posteriorly and covering the frontal bones while the nostrils occupied a position above the orbit. The members of Agrophiidae apparently represent the ancestors of Squalodontidae Brandt, 1873 (Oligocene-Miocene) which in the Lower Miocene gave rise to contemporary dolphins (Delphinidae) and sperm whales (Physeteridae) and during the Miocene to beaked dolphins (Ziphiidae). The systematics of the suborder of toothed whales has not been adequately studied, not only with respect to morphology and diagnosis of the species, but even the number and composition of the families. 582 Most scientists have по doubts about the correctness of recognizing river dolphins, beaked whales, and sperm whales as individual families (Kellogg, 1928; Nishiwaki, 1972 placed the dwarf sperm whale in the fam- ily Kogiidae). Views differ regarding the white whale and the narwhal, dolphins per se, and the common porpoise (for more details, see p. 586). Combining all the three above-mentioned groups of whales into two families is evidently more correct (Hershkovitz, 1966; Yablokov et al, 1972). Thus the suborder of toothed whales includes three contemporary superfamilies and four families: river dolphins—Platanistoidea Simp- son with one family of river dolphins, Platanistidae Gray (= Susuidae); dolphins—Delphinoidea Flower with two contemporary families: dol- phins, Delphinidae Gray and narwhals, Monodontidae Gray; and sperm whales—Physeteroidea Gill with two families: sperm whales, Physeteri- dae Gray and beaked dolphins, Ziphiidae Gray (= Hyperoodontidae). The suborder comprises 31 genera and 71 species (Hershkovitz, 1966). In the USSR, the members of two superfamilies and three families (not river dolphins), 15 genera (roughly 48% of the world fauna) and 17 species (roughly 23%) are found (Tomilin, 1962). The suborder includes five extinct families: Agrophiidae Abel (Upper Eocene of North America); Squalodontidae Brandt (Upper Oligocene-Upper Miocene of Europe, Lower Miocene of South America, Australia, and New Zealand, and Middle and Upper Miocene of North America); Eurhinodelphidae Abel (Lower Miocene of South America, Lower and Upper Miocene of Europe, Middle Miocene of North America, and Upper Miocene of Japan); Hemisyntrachelidae Slijper (Middle Miocene of North America and Lower Pliocene of Europe); and Acrodelphidae Abel (Lower and Upper Miocene of Europe and Middle Miocene and Lower Pliocene of North America). Some toothed whales have acquired economic importance in the recent past. Sperm whales are even now caught in large numbers. At the same time, many of the scarce and less numerous species are practically of no importance. Not all the species of toothed whales which could serve as valuable targets of hunting are caught in sufficient numbers in the waters of the USSR (for example, killer whales) and some are not caught at all (for example, dolphins in the waters of the Far East). (V.S.) Key to Species of the Suborder of Toothed Whales Inhabiting and Probably Found in USSR Waters 1( 6). Anterior part of snout strongly projecting beyond tip of lower jaw. Functional teeth absent or not more than three pairs in 2768}, 3) (12). 4 (5). 5 ( 4). 6( 1). 7 (16). Sec): 9 (10). 10 ( 9). 583 upper jaw and not less than seven pairs in lower jaw. Symphysis of lower jaw halves covers more than one-third their length. Body dimensions large (3-5 m in newborns and up to 15-21 m in adults). Head accounts for one-third body length. Blowhole at end of snout. Dorsal fin low, hump-shaped. Lower jaw with 18-28 pairs of massive teeth. First cervical vertebra freenMemaAINGeMUSEG HET. ом аа able t os eae: MIDE TOR fade ne 2 Sperm whale, Physeter catodon (p. 801). Body dimensions small (not more than 4 m in adults). Head accounts for roughly one-fifth body length. Blowhole at center of head. Dorsal fin concave along posterior margin. Lower jaw with 9-16 pairs of small curved teeth. All cervical vertebrae fused. Body length 2.7-3.4 m. Lower jaw with 12-16 (less often, 10 Onli) spairssOf teeth eR VERT ORs PN TIA rte. 2 on S30 PEMA, ДЕН Pygmy sperm whale, Kogia breviceps (p. 841). Body length 2.1-2.7 m. Lower jaw with 8-11 (less often, 13) pairs of teeth. ... Dwarf sperm whale, Kogia simus (p. 845). Anterior part of snout slightly projecting or not projecting beyond tip of lower jaw. Functional teeth present in both upper and lower jaws. If absent in upper jaw, their number in lower jaw does not exceed seven pairs. Symphysis of lower jaw halves does not exceed one-third their length. Teeth absent in upper jaw, not more than two pairs in lower jaw. One or two pairs of furrows on neck converging anteriorly. Notch not present on posterior edge of caudal flukes. Dorsal fin closer to flukes than to flippers. Width of midportion of straight, long, thin rostrum not less than one-seventh its length. Only one pair of teeth in lower jaw, located far behind anterior end of jaw. Teeth highly flattened laterally (their smaller diameter not less than one-half larger diameter). Fully developed tooth laterally trapezoid. In skull, antorbital notches relatively poorly developed. Aperture of pair V of nerves in maxilla anterior to aperture in premaxilla or at the same level. Palatine bones not contiguous. Ratio of length of tooth crown to width more than 6:1 ...... .... Stejneger’s beaked whale, Mesoplodon stejnegeri (р. 866). In skull, antorbital notches relatively well developed. Aper- ture of pair V of nerves in maxilla posterior to aperture in premaxilla. Palatine bones contiguous. Ratio of length of tooth crownto width notwmore Вай 4 ei. lees teresa iaielerst 438. 584 Te: 12 (13). 13 (12). 14 (15). 15 (14). 16, (7): 17 (26). 18 (23). 19 (22). 20 (21). 21 (20). 22 (19). 23 (18). 24 (25). Width of midportion of rostrum not more than one-sixth its length. Teeth located on anterior end of lower jaw and conical (larger diameter exceeds smaller by not more than 1.5 times). Length of head about 20% of body length. Massive crests running along maxillae reaching level of skull apex. Maximum diameter of anterior tooth usually not more ап 2 ст ...... Northern bottlenose whale, Hyperoodon ampullatus (p. 885). Length of head about 13-17% of body length. Low crests run- ning along maxillae not reaching level of skull apex. Maximum diameter of anterior tooth does not exceed 2 cm. Two pairs of teeth present. “Beak” flattened dorsoventrally. Rostrum from above not triangular ......................4. te eer Baird’s beaked whale, Berardius bairdii (p. 850). One pair of teeth present. “Beak” not flattened dorsoventrally. Rostrum) from, abovetniangular.., Aisa) ана в oe eee eRe TING ET. Cuvier’s beaked whale, Ziphius cavirostris (p. 876). In most species, teeth present in upper and lower jaws. Teeth absent in upper jaw only in Risso’s dolphin, and in lower jaw in narwhals. In latter case, however, width of midportion of rostrum 2/5 - 1/3 its length. Furrows not visible on neck. Notch present between caudal flukes. Dorsal fin lacking. Head rounded, without “‘beak”’. Number of teeth not more than 44. Neck distinctly visible. Horny protuberances absent on dorsum. Body monochromatic (white or yellow in adults). Up to 44 teeth in upper and lower jawsiit 32 при Benue Ot tec A ees SLUNG, EELS. ee White whale, Delphinapterus leucas (p. 757). Body of adult animals spotted (monochromatic, dark, in juve- niles). Teeth lacking or not more than one pair оп upper jaw. In males and sometimes in females, left upper tooth in form of Spirallyatwistedy tusk ооо sR di. лены: Dy AEROS ее. Narwhal, Monodon monoceros (p. 791). Number of teeth up to 100. Neck lacking. Horny protuberances presenton dorsum 2. ive: ase oie 12 Ae Rees ome .. Black finless porpoise, Neophocaena phocaenoides (p. 750). “Beak” well developed. Number of teeth exceeds 160. Entire body black, with narrow light-colored band only along abdomen from neck to tail; band forms rhomboidal patch on 439 25 (24). 26 (17). 27 (42). 28 (29). 29 (28). 30 (31). 31 (30). 32 (37). 33 (34). 34 (33). 35 (36). 36 (35). 37 (32). 38 (39). 39 (38). 40 (41). breads tae. обои rors sR ее О ЗН о. .. Northern right whale dolphin Lissodelphis borealis (р. 646). Body black except for white lower sections and abdominal flanks .. Southern right whale dolphin, Lissodelphis peroni (р. 651). Dorsal fin present. “Beak” absent or barely perceptible. Body length exceeds 4 m. Flippers broad (length not more than twice width) and rounded. Teeth massive (diameter more than 2.5-2.8 cm). Body dark in color. White band on abdomen extending as tongue on flanks in region of anal opening, its tip directed anteriorly. Dorsal fin high, especially in males (up to Пети ай ее Bee aid Mid fo ke Killer whale, Orcinus orca (p. 680). Body length less than 4 m; if more, flippers narrow (length exceeds width by more than 2.5 times) and pointed. Tooth diam- eter not more than 2.5-2.8 cm. Body color different. Dorsal fin smaller. Teeth absent in upper jaw (sometimes one or two pairs seen); two to seven pairs present in lower jaw .................... sys APCs es EEE Risso’s dolphin, Grampus griseus (p. 696). More than six pairs of teeth in upper jaw and more than seven pairs in lower jaw. Teeth in upper and lower jaws not more than 8- 13 pairs each. Low dorsal fin in anterior half of body. Head anteriorly obtuse with high forehead. Teeth only in anterior half of upper jaw овал AGEL Pilot whale, Globicephala melaena (p. 702). Relatively high dorsal fin roughly midbody. Head anteriorly rounded. Teeth in upper jaw not exclusively in anterior half. Entire body black. Roughly 50-51 vetebrae ................. а False killer whale, Pseudorca crassidens (р. 674). Body dark gray, edge of jaws and region around anal opening white: Roughly/67 -7Ojvertebrae! 335. ae. 5 ek. ВВ LS nS. Se AEP Dwarf killer whale, Feresa attenuata (p. 719). Teeth in upper and lower jaws more than 15 pairs each. Body size of adult animals exceeds 2m ..................... ан Broadsnout dolphin (melon-headed whale), Lagenorhynchus electra (p. 672). Body size of adult animals exceeds 2 m. Dorsal side of caudal crest with high keel. White field on body flanks sharply set off from surrounding dark background. 586 41 (40). 42 (27). 43 (44). 44 (43). 45 (52). 46 (47). 47 (46). 48 (49). 49 (48). 50 (51). 440 51 (50). Number of teeth in upper jaw (15-24 pairs) usually less than in lower(2 2228 pairs). ОЕ О sty ots eae нана BA ROS Dall porpoise, Phocoenoides dalli (p. 738). High keel on dorsal side of caudal crest lacking. Body color dif- ferent. Number of teeth in upper jaw (16-30 pairs) usually more than. in lower, jaw:(16 227 (pairs) $0.20 JA. hee: 2 he В ORE Common porpoise, Phocoena phocoena (p. 722). “Beak” distinctly perceptible. Symphysis of lower jaw more than one-fourth its length. Tooth CLOWNS! BTOOVEG NEAL AE ARNT NO ЗОО ран... И ALLL Rough-toothed dolphin, Steno bredanensis (p. 591). Symphysis of lower jaw less than one-fourth its length. Tooth crowns smooth. “Beak” of moderate length or short. Body length exceeds “beak” length (from tip of snout to beginning of corpus adiposum) by more than 25 times. Diameter of teeth in middle of tooth row exceeds 3 mm. : Teeth not more than 20 pairs in upper jaw and 26 pairs in lower. Diameter of teeth in middle of tooth row exceeds 6 mm.“Beak” dark in color. Keel absent on dorsal side of caudal Стел nuns Bottlenose dolphin, Tursiops truncatus (p. 632). More than 26 pairs of teeth each in upper and lower jaws. Diam- eter of teeth in middle of tooth row less than 6 mm. If number of teeth less (up to 22 pairs), their diameter more (up to 7 mm). Beak white. Keel present on dorsal side of caudal crest. Tip of “beak” white. Trunk dark-colored above right up to joint with flippers. Number of vertebrae exceeds 84. Number of teeth in upper and lower jaws less than 28 pairs. Pterygoid bones separated by gap, broadened anteriorly ..................... .. White-beaked dolphin, Lagenorhynchus albirostris (р. 660). Tip of “beak” dark. Dark color of trunk does not descend to joint with flippers. Number of vertebrae less than 84. Number of teeth in upper and lower jaws more than 28 pairs. Pterygoid bones adjacent or separated by gap, not broadened anteriorly. Number of vertebrae 74-78; ribs 12-14 pairs; teeth in upper jaw usually 30 -32 pairs. Narrow dark band extending along body flanks from base of flippers to lower caudal crest ........... ELE ie WOON OLR Rn ten Da Pacific white-sided dolphin, Lagenorhynchus obliquidens (p. 664). Number of vertebrae 77-82. Ribs 14-15 pairs. Teeth in upper jaw usually 35 -38 pairs. Dark band along body flanks from base of flippers to lower caudal crest absent ...... тан Se Atlantic white-sided dolphin, Lagenorhynchus acutus (p. 656). 52 (45). “Beak” long. Body length exceeds “beak” length (from tip of snout to beginning of corpus adiposum) by not more than 20 times. Diameter of teeth in middle of tooth row not more than 3 mm. 53 (54). Dark band extending from flippers to chin. Palate with two longitudinalfgroOvesime HI BAO AB, RR OSL MIS ОЛ. Common dolphin, Delphinus delphis (р. 607). 54 (53). Dark band extending from flippers to chin absent. Palate with- out grooves. 55 (56). Body dark gray above, white below. Length of rostrum of skull twice length of cerebral section. Number of teeth 148258 ...... .. Long snout [spinner] dolphin, Stenella longirostris (p. 604). 56 (55). Body black or blackish above, white, whitish, or ash-gray below. Length of rostrum less than twice length of cerebral section. Number of teeth 32-50. 57 (58). Body black above, white below. Number of teeth 4-2? ....... .. Blue-white [striped] dolphin, Stenella coeruleoalba (р. 594). 58 (57). Body blackish above, whitish or ash-gray below. Number of teeth м. 59 (60). Body ash-gray below. Maxillary teeth small, less than 3 mm in diameter, Number of vertebrae 79 8. Malay [Pan-Tropical spotted] dolphin, Stenella dubia [= айе- пиащ] (р. 601). 60 (59). Body whitish below. Maxillary teeth large, more than 5 mm in _ diameter. Number of vertebrae not more than 70 ........... а: Bridled dolphin [Atlantic spotted dolphin], Stenella frontalis (p. 602). (V.S.) SUPERFAMILY OF DOLPHINS Superfamily DELPHINOIDEA Flower, 1864 Cetaceans of small and medium dimensions. The body of most of the dolphins is well proportioned. The caudal fin bears a deep notch between the flukes. Many species have a large dorsal fin (sometimes lacking) in the midbody. Furrows not seen on the neck. Rostral part of head stretched into a well-distinguishable “beak” which, however, may be poorly developed or altogether absent. The “beak” may 441 588 be sharply demarcated from the frontal portion of the head or may pass smoothly into it. The body is monochromatic (dark or light in color) or dark above and light-colored below, or dark with light-colored bands that vary in disposition, shape, and number. The zygomatic and temporal bones are poorly developed in the skull. The supraoccipital and frontal bones overhang a small temporal fossa. Crests are lacking on the maxillae. The pterygoid and nasal bones are relatively small. The palatine bones are joined along the midline of the palate and are not separated from the vomer. The zygomatic process of the squamosal bone is highly reduced. The symphysis of the lower jaw is not more than one-third the jaw length. The width (spacing) of the lower and upper jaws is nearly similar. The number of teeth varies from © to 5. These are migratory animals living in schools. Their food is diverse: fish (ichthyophagous whales), cephalopod mollusks (teuthophagous), cephalopods and fish (teutho-ichthyophagous), and warm-blooded vertebrates (sarcophagous). The species of the superfamily live in almost all the seas of the world right up to the icy seas in the high Arctic. Some are very widely distributed, almost throughout the world (common dolphin and killer whale) while others are more localized. Some species (genus Sotalia) live in rivers (South America and southern Asia). Members of the family are known from the Lower Miocene, having separated from the ancient Squalodontidae. The oldest members (Argy- rocetus, Schizodelphis, and Delphinavus) had a long and narrow rostrum and a large number of teeth. The contemporary form of the dolphin can be traced to the end of the Miocene-Early Pliocene. The systematics of dolphins has been differently interpreted by var- ious scientists. In the superfamily of dolphins, Delphinoidea, one family—Delphi- nidae (Tomilin, 1957), two families—Delphinidae and Monodontidae (Hershkovitz, 1966; Yablokov её al., 1972), or three families—Monodon- tidae, Delphinidae, and Phocoenidae (Simpson, 1945) have been distinguished. In more detailed systems, the number of families has risen to four—Monodontidae, Stenidae, Phocoenidae, and Delphinidae (Andersen and Jones, 1967), or even six—Monodontidae, Delphinidae, Grampidae, Globicephalidae, Orcaelidae, and Phocoenidae (Nishiwaki, 1966; Ridgway, 1972). It would appear to be more correct to combine all these groups into two families: dolphins (Delphinidae) and narwhals (Monodontidae). 442 589 Representatives of both the families are found in the USSR fauna. The extinct forms are represented in the superfamily of dolphins by the family Eurinodelphinidae (three genera), family Hemisyntrachelidae (two genera), and family Acrodelphidae (six genera). Moreover, 24 gen- era belong to the family Delphinidae (Simpson, 1945). Some representatives of the family are of economic importance and are hunted in large numbers. Others are almost not caught because of their small number or for other reasons and have no economic importance. (V.S) Family of Dolphins Family DELPHINIDAE Gray, 1821 Animals of small, medium, and large dimensions. The dorsal fin in most cases is well developed. There is no neck between the head and the trunk. The rostrum of the skull is usually longer than the cranium. The petrous temporal bone does not grow toward the skull but is attached to it by ligaments. The cervical vertebrae may be fused (atlas and axis more often fused). The anterior ribs have a double articulation with the vertebrae. The ulnar process is well developed in the forelimb. Open seas, coastal zone, and rivers are inhabited. The geographic distribution of the members of the family covers the range of the superfamily; only the northern limit falls more southward. The family comprises 18 genera,! of which 14 have been reported (or may still be found) in USSR waters: rough-toothed dolphins, Steno Gray; spotted dolphins, Stenella Gray; common dolphins, Delphinus Linnaeus; bottlenose dolphins, Tursiops Gervais; right whale dolphins, Lissodelphis Gloger; shorthead dolphins, Lagenorhynchus Gray; false killer whales, Pseudorca Reinhardt; killer whales, Orcinus Fitzinger; Risso’s dolphins, Grampus Gray; pilot whales, Globicephala Lesson; dwarf killer whales, Feresa Gray; common porpoises, Phocoana G. Cuvier; Dall porpoises, Phocoenoides Andrews; and black finless porpoises, Neophocaena Palmer. The remaining four genera inhabit waters outside the USSR: Sotalia Gray, Lagenodelphis Fraser, Cephalorhynchus Gray, and Orcaella Gray. The species of some genera are of economic importance. (V.S.) 1 According to P. Hershkovitz (1966), the placement of Lagenorhynchus electra Gray in an independent genus, Peponocephala Nishiwaki and Norris (Electra auct—nom. praeocc.), requires further justification. 444 442 590 Genus of Rough-toothed Dolphins Genus Steno Gray, 1846 1846. Steno. Gray. Zoology Voyage Erebus and Terror. I, p. 30. Delphinus rostratus Cuvier = Steno bredanensis Lesson, 1828. (V.H.) Small dolphins reaching up to 2.5 m in length (Fig. 232). The “beak” on the head passes smoothly into the slope of the fore- head. The dorsal fin is triangular, high, with a notch along the posterior edge [falcate]. The body is dark gray on the dorsum, light gray with yellowish-white spots on the flanks, and white on the abdomen. The rostrum is long and narrow (Fig. 233), its length more than three times its width. The symphysis of the lower jaw constitutes over 30% of its length. The pterygoid bones are adjacent. The postorbital processes of the frontal bone are small. The teeth are large, with grooved crowns. Teeth 20=27 (usually 3). Vertebrae 65 - 66. The first and second vertebrae are fused. The biology of the dolphins of this genus is almost unstudied. Their population is insignificant. They live in groups and feed on squids and fish. The geographic range covers the temperate and warm waters of the Atlantic and Pacific oceans and also the Indian Ocean. They have not been reported in the waters of the USSR. However, they might be found in the region of Kuril Islands and the Sea of Japan. The genus comprises one species: the rough-toothed dolphin, S. bredanensis Lesson, 1828. Sometimes the rough-toothed dolphin, together with the genera Sotalia Gray and Sousa Gray, is placed in a separate family, Stenidae Fraser and Purves, 1960 (Anderson and Jones, 1967). (V.S.) Fig. 232. Rough-toothed dolphin, Steno bredanensis (figure by N.N. Kondakov). 591 442 Fig. 233. Skull of rough-toothed dolphin, Steno bredanensis (figure Бу М.М. Kondakov). ROUGH-TOOTHED DOLPHIN? Steno bredanensis Lesson, 1828 1817. Delphinus rostratus. Desmarest. Nouv. Dict. Hist. Nat., 9, p. 160. Bretagne, France. Nom. praeocc. 1823. Delphinus frontatus. G. Cuvier. Rech. ossemens foss., 5, p. 278. Por- tugal. Description does not conform to the rules of nomenclature. 1828. Delphinus bredanensis. Lesson. Hist. Nat. mamm. oiseaux decon- verbés depuis 1788. Substituted for Delphinus rostratus Desmarest, 1817. (У.Н.). Single species of the genus. The body is well proportioned. The “beak” is somewhat compressed laterally. The dorsal fin lies roughly midbody. The flippers are broad at the base. * Russian name corresponds to the English name. (V.H.) 446 592 A sharp demarcation of different colors is a characteristic feature of body coloration. The snout is sometimes white. The adult male measures 2.23 m in length and weighs 102 kg. (У.5.) Transgressions and habitation in our waters of the Sea of Japan and near the Kuril Islands are possible (Fig. 234). Outside the USSR, this dolphin is found in the Pacific Ocean south of the strip Japan—Hawaiian Islands—California; in the Atlantic Ocean from the southern part of the North Sea (Holland) and waters of France in the east and Virginia in the west to roughly 40° $ lat. (Tristan da Cunha, Argentina); and in the Indian Ocean from the northern coasts and Java to the Cape of Good Hope. Geographic variability has not been established. Biology not known. Lives in small schools. (V.H.) Genus of Spotted Dolphins Genus Stenella Gray, 1866 1864. Clymene. Gray. Proc. Zool. Soc. London, p. 237. Delphinus euphrosyne Gray, 1846. Nom. Ргаеосс. 1866. Stenella. Gray. Proc. Zool. Soc. London, p. 214. Steno attenuatus Gray, 1846. 1866. Euphrosyne. Gray. Ibid., p. 214. Clymene euphrosyne Gray. Nom. ргаеосс. 1880. Prodelphinus. Gervais. In: Van Bénéden et Gervais. Ostéographie des Cétacées, p. 604. Substituted for Clymenia Gray, 1864. (V.H.) Small dolphins; body length up to 2.7 m. The body is well proportioned. The dorsal fin is located midbody and its apex curved backward. The flippers are crescent-shaped. The “beak” is long and narrow. The lower jaw is slightly longer than the upper. The body coloration differs markedly from species to species and, further, is subject to considerable individual variation. The dorsum and upper sections of the body flanks are usually dark; the lower portion of the body flanks and abdomen are light in color. The premaxillae are curved in the rostral part of the skull. The ptery- goid bones are adjacent. Longitudinal grooves are absent or very small on the palate. The symphysis of the lower jaw constitutes less than one- fifth the length of the jaw itself. The teeth are small, sharp, and 34-65 in each half of the jaw. The cerebral section of the skull is broad in the occipital region and flat in the frontal region. The structural features of the skull of the spotted dolphins on the one hand are similar to the common dolphins, Delphinus, and on the other to the genus of bottlenose dolphins, Tursiops. ul uy yy et ЕЕ В Tas: Ant Peay int]! Ih ПИН ОКА ОА il | Lent 1 ий | | | |. || Л 1 | 1000 0 1000 2000 3000 4000 5000 km О НИТИ : НИ НН Fig. 234. Range of rough-toothed dolphin, Steno bredanensis (У.А. Arsen’ev). 443 594 The common features of spotted dolphins and common dolphins are the long rostrum, slightly exceeding the cerebral section of the skull, the large number of very small teeth, and premaxillae usually confluent with the rostral part of the skull. However, as in bottlenose dolphins, spotted dolphins do not have longitudinal grooves on the bony palate. Long, thin, spinous and transverse processes are characteristic of the vertebrae. The cervical vertebrae may be fused. The sternum is T-shaped and consists of four sections. Ribs 15 - 16 pairs. In the flippers, the second and third digits are the longest (2nd one longer). Biology almost not known. Its distribution (Fig. 235) ranges from the cold to the tropical waters of the Pacific and Atlantic oceans (including the Mediterranean Sea), from the Bering Sea and Greenland to the Cape of Good Hope, Cape Horn, and Australia, and the Indian Ocean. Not caught in our waters. The composition and systematics of species of the genus have not been adequately studied. Ten species are usually recognized but their number seems to have decreased: (1) S. asthenops Cope, (2) S. clymene Gray, (3) S. coeruleoalba Meyen, (4) S. crotaphiscus Cope, (5) S. dubia G. Cuvier, (6) S. frontalis G. Cuvier, (7) S. graffmani Lonnberg, (8) S. longirostris Gray, (9) S. malayana Lesson, and (10) 5. pernettyi Blainville.* Only the blue-white dolphin, S. coeruleoalba, is encountered in the waters of the USSR while the appearance or residence of S. dubia, S. frontalis, and S. longirostris is possible. (V.S.) BLUE-WHITE [STRIPED] DOLPHIN Stenella coeruleoalba Meyen, 1833 1833. Delphinus coeruleo-albus. Meyen. Nova. Acta Leop.-Carol., 16, 2, р. 609. Near La Plata estuary. 1846. Delphinus styx. Gray. Zoology Voyage Erebus and Terror, 1, p. 39. Atlantic Ocean around South Africa. 1846. Delphinus euphrosyne. Gray. Ibid., p. 40. North Atlantic, waters of England. 1848. Delphinus lateralis. Peale. U. S. Explor. Exped., Mammalia, p. 34. Pacific Ocean at 13°58’ N lat. and 161°22’ W long. (V.H.) Diagnosis Body length up to 260 cm. The dorsal surface is black and the ventral sur- face white. A narrow black band runs from the eyes to the anal opening 3 In alphabetical order, after Hershkovitz (1966). a т, TT | ТИ. ih i es И мн a el i О ИИ р. И, | i i! yt i! 1 И Nh! . м iN И it 0 1000 1000 2000 3000 4000 5000 Кт Fig. 235. Range of spotted dolphins, Stenella (V.A. Arsen’ev). 445 447 447 596 and from the eyes to the base of the flippers. The flippers are small. The length of the rostrum is less than double the length of the cerebral section of the skull. Teeth 4-28. Vertebrae about 76-79. (V.S.) Description The main body measurements of female and male blue-white dolphins (Fig. 236) caught in the waters of Japan (Tomilin, 1957) are respectively (in cm): body length 200 and 196 (may go up to 2.7 m); distance from tip of snout to base of flippers 48 and 50, up to blowhole 34 and 35, up to anterior edge of dorsal fin 93 and 94, and up to anal opening 137 and 138; height of dorsal fin 17 and 15, length of the base of dorsal fin 31 and 28; width of caudal flukes 67 and 44; and length of flippers 27. Of the more than 3,000 blue-white dolphins caught at the end of November, 1967 at village Kawana (Honshu Island), a random sample of 75 adults (body length 170 to 256 cm) comprised 59 females and 16 males; 71 newborn calves (body length 97 to 144 cm) comprised 36 females and 35 males (Sokolov and Kasupa, 1969). Among the females studied, six with a body length ranging from 184 to 217 cm (184, 199, 206, 208, 213, and 217) were immature; two with a body length of 139 [sic.] and 213 cm had not given birth before but were ready for fertiliza- tion (with large follicles in the ovaries); and nine had calved before (they bore traces of the corpus luteum of pregnancy in the ovaries) but were now barren or had lost their calves. The body length of these last nine females was: 217 cm (two), 224, 226 (two), 229, 231, 232, and 239 cm. The ovaries of two of them (body length 232 and 239 cm) contained large follicles. Of the remainder, 40 females were lactating (the status of two females could not be ascertained) and the ovaries of three con- tained mature follicles. The body length of these 40 females was (in cm): 209, 216 (two), 218 (two), 219, 221 (three), 222, 223 (four), 224 (four), Fig. 236. Blue-white dolpin, Stenella coeruleoalba (figure by М.М. Kondakov). 597 225, 226 (four), 227 (three), 228 (two), 229 (three), 230, 231 (three), 232 (two), 233, and 234 (two). The body length of immature males varied from 170 to 214 cm (170, 176, 184, 191, 195, and 214 cm) and of mature males from 224 to 256 cm: 224, 228, 232 (two), 233, 235, 237, 239, 241, and 256. The condylobasal length of the skull (Fig. 237) of these dolphins 448 from La Mancha was (in cm): 47, length of rostrum 29, width of rostrum at base 12, length of lower jaw 41, and length of symphysis of lower jaw 4 (Tomilin, 1957). Phalangeal formula: Ц, Пу-1о, Ша, [V4, and У, _, (V.S.) 4 ра of РА Я Я 7 4 ERAS А х 447 Fig. 237. Skull of the blue-white dolphin, Stenella coeruleoalba (figure Бу М.М. Kondakov). 598 a Be Aue ‘oe {| и 250 0 250 500 750 1000 km ЕЕ BN] 150 160 170 130 140 448 Fig. 238. Range of the blue-white dolpin Stenella coeruleoalba in the USSR (V.A. Arsen’ev). Geographic Distribution This dolphin inhabits the warm and temperate belts of the Atlantic and Pacific oceans. Studies of its distribution are highly schematic. Geographic Range in the USSR Waters of the Pacific Ocean around the Kuril Islands and possibly the waters of the Bering Sea (Fig. 238). Geographic Range outside the USSR (Fig. 239) Strays in the Atlantic Ocean northward up to the coasts of Great Britain, Shetland and Orkney Islands, and southern Greenland; known in the waters of Quebec and Nova Scotia. Not reported in the Northern, Baltic, 1000 0 1000 2000 3000 4000 5000 km le \ ` 150 Fig. 239. Species range of the blue-white dolphin, Stenella coeruleoalba (V.A. Arsen’ev). 449 450 600 or Barents seas.* Distributed southward up to Rio de La Plata along the South American coast and up to the southern tip of Africa. Lives in the Mediterranean Sea. In the Pacific Ocean, known from the coasts of Japan and the Kuril Islands in the west and from the Bering Sea, British Columbia, states of Oregon and Washington in the east up to New Zealand. [51с.] (V.A.) Geographic Variation Our knowledge of the geographic variation of the blue-white dolphin is extremely scant and this aspect has generally not been discussed in the world literature. In our literature on the fauna of the USSR, the existence of two subspecies is sometimes acknowledged (Tomilin, 1957, 1962). These are: the nominal form in the seas of the Far East and the Atlantic form, S. c. euphrosyne Gray, 1946, for “the Baltic Sea, possibly the Barents 5еа”. The main difference between them, in addition to some skull features, lies in coloration (band from eyes to base of flippers single in the former and double in the latter). Sometimes these two forms are treated as independent species. Their nomenclature differs according to different authors. As pointed out above (see “Geographic Distribution”), the existence of the Atlantic form (euphrosyne) in our waters is just not possible. According to the convention prevailing in our literature, the name S. c. coeruleoalbus Meyen, 1833 may be retained only tentatively for the blue-white dolphin found in our Far Eastern waters. However, it would be better to use the binary name. It is quite possible that the use of the name styx Gray, 1846 might be more correct. The systematics and nomenclature of the species and forms of the genus Stenella are complex and in a high state of confusion (compare, for example: Ellerman and Morrison-Scott, 1951, 1962; Tomilin, 1957, 1962; Hall and Kelson, 1959; Hershkovitz, 1966; and others). A radical review of the subject, based on substantial evidence, is required. (V.H.) Biology The biology of the blue-white dolphin has almost not been studied and practically no observations have been made on the behaviour of these animals at sea. ‘Information and assumptions on habitation in these seas (Tomilin, 1957, 1962) have not been confirmed in the latest compilations (Ellerman and Morison-Scott, 1951, 1966; von den Brink, 1958; Koval’skii, 1964; Bobrinskii et al., 1965;- Hershkovitz, 1966; Siivonen, 1967). (V.H.) 451 601 The largest of the measured embryos was 103 cm long. All the young dolphins 120-150 cm long were comparatively recently born since their stomach contained exclusively milk or milk together with small quanti- ties of the remnants of squids. The smallest female with a corpus luteum in the ovary measured 215 cm in length; apparently females of this size attain maturity. Among animals of this length (or slightly larger), the ovaries and testes begin to lengthen rapidly. Among dolphins caught in November, 1956, 12 were immature females, 72 gestating, and 27 simul- taneously gestating and lactating. The oldest of these dolphins was 18 years of age (Nishiwaki and Yagi, 1953). Sufficient numbers of the blue-white dolphin are caught in the coastal waters of some regions of Japan and China. Hunting is done in small crafts and with small-bore harpoon guns. The kill is sold in fish markets. Economic importance is very negligible. (V.A.) MALAY [PAN-TROPICAL SPOTTED] DOLPHIN Stenella dubia [= attenuata] С. Cuvier, 1812 1812. Delphinus dubius. G. Cuvier. Ann. Mus. Hist. Nat., Paris, 19, p. 9. French waters. [nomen dubium] 1826. Delphinus malayanus. Lesson. Voyage autour du monde..., Zoologie, I, p. 184. South China Sea, strait. between Java and Kalimantan (Borneo). [nomen dubium] 1846. Steno attenuatus. Gray. Zoology Voyage Erebus and Terror, Г, p. 44. Type locality not established. (V.H.) Body length up to 183 cm. The color is blackish above and ash-gray below (including the caudal flukes). The flippers are large. The length of the rostrum is less than double the length of the cerebral section of the skull. Teeth 3-44. Vertebrae 79-81. The body is well proportioned, resembling the common dolphin (Fig. 240). The forehead is low and long. The color is pale bluish-black 451 Fig. 240. Malay [Pan-tropical spotted] dolphin, Stenella dubia (figure by М.М. Kondakov). 602 above, aSh-gray ог grayish-white below. Numerous minute whitish ог grayish, sometimes pinkish spots are scattered over the body. A dark band runs from the flippers to the corner of the mouth. The upper jaw and lateral sections of the lower jaw are black. The tip of the “beak” is white. A band runs forward from a black spot around the eye. The main body measurements (as percentage of body length) of males with a body length of 165 - 208 cm and females 180-201 cm (Nishi- waki, 1966) are respectively: from end of upper jaw to blowhole 14.2 - 17.1 and 14.2-17.7, from fork between caudal flukes to posterior edge of dorsal fin 39.2-40.5 and 39.6-42.8, from caudal fork to anal opening 25.2-26.5 and 25.2-27.2; length of flippers 14.4-15.1 and 12.0- 15.2, maximum width of flippers 5.0-5.3 and 4.2 -5.3; length of base of dorsal fin 12.1-14.2 and 12.4-14.2, height of dorsal fin 8.2-9.7 and 7.8-9.4; and width of caudal flukes 21.6 -22.5 and 23.5. The main dimensions of the skull (as percentage of condylobasal length) of these dolphins with a condylobasal length of skull 381 - 428 mm (Nishiwaki, 1966) are: length of rostrum 58.5 - 61.5 and its width at the base 21.5 - 23.3; interorbital width 38.8 - 42.8; length of lower jaw sections 84.5 - 90.3, and length of symphysis of lower jaw 16.0 - 17.3. (V.S.) The presence of this species in USSR waters has not been estab- lished. Presence in or transgressions into the waters of the Sea of Japan and around the Kuril Islands are possible. Outside the USSR, this species is found in the South Atlantic Ocean up to Cape Horn and Cape of Good Hope, the Indian Ocean, the Pacific Ocean from New Zealand to the South China Sea, Hawaiian Islands, and waters of Japan. (V.H.) Biology is not known. BRIDLED [ATLANTIC SPOTTED] DOLPHIN? Stenella frontalis G. Cuvier, 1829 1829. Delphinus frontalis. G. Cuvier. Regne Animal, I, р. 228. Cape Verde Islands, Atlantic Ocean. (V.H.) Body length about 180 cm. Color of the dorsal section blackish, changing over to grayish on the flanks. Abdomen whitish. The dorsum has light- colored and the abdomen dark spots. A dark-colored band runs from the corner of the mouth toward the flippers (Fig. 242). The eyes are encircled 452 by black rings. The flippers are small. The length of the rostrum is less 5 The Russian name is given here. 603 452 Fig. 242. Bridled dolphin, Stenella frontalis (figure by М.М. Kondakov). than double the length of the cerebral section of the skull (Fig. 243). 35-44 37-38 Teeth 32-7 (usually 35-35). Vertebrae: 7 cervical, 15 thoracic, 19 lumbar, and 36 caudal; total 77. Phalangeal formula: I,-3, По, Шу, 1V3, and V5. (V.S.) Its residence has not been established in the waters of the USSR. Habitation in or transgressions into the Sea of Japan are possible. Outside the USSR, it is found in the Atlantic Ocean (from Cape Verde Islands to Cape of Good Hope and from North Carolina to 22777 777 a RUIN SS Us 453 Fig. 243. Skull of the bridled dolphin, Stenella frontalis (figure by М.М. Kondakov). 455 604 Florida), Mediterranean Sea, and Indian Ocean. It has been reported in the waters of Japan and the Korean peninsula (Cheju-do Island and Sea of China). (V.H.) Biology is not known. LONG-SNOUT [SPINNER] DOLPHIN Stenella longirostris Gray, 1828 1828. Delphinus longirostris. Gray. Spicil. Zoologica, 1, p. 1. Type locality not established. 1846. Delphinus alope. Gray. Zoology, Voyage Erebus and Terror, 1, р. 52. 1846. Delphinus roseiventris. Wagner. Schrebers Sauge thiere, 7, pl. 360. (У.Н.) Body length roughly up to 213 cm. Dorsal side of body dark gray. The abdomen is white with random gray spots. The flippers are small (Fig. 244). The length of the rostrum is double the length of the cerebral section of the skull (Fig. 245). The bony palate has very small longitudinal grooves. Teeth 2-56 (usually 22-25). Vertebrae 72-73. (V.S.) Its residence has not been established in USSR waters. Habitation in or transgressions into the Sea of Japan are possible. Outside the USSR (Fig. 246), it is reported in the Atlantic Ocean south of the Bahamas and Senegal up to Cape Horn and Cape of Good Hope, the Indian Ocean (Ceylon), and the Pacific Ocean from Mexico and Japan to Australia. (V.H.) Biology is not known (Fig. 247). Genus of Common Dolphins Genus Delphinus Linnaeus, 1758 1758. Delphinus. Linnaeus. Syst. Nat., ed. X, 1, p. 77. Delphinus delphis Linnaeus, 1958. Small cetaceans with body length up to 2.6 m. The “beak” is long. The dorsal fin is high and crescent-shaped. The narrow flippers are also curved. Body coloration is subject to considerable individual variation but is usually black above, gray with dark- and light-colored bands on the flanks, and light-colored below. The rostrum is 1.5 to 2 times longer than the cranium and bifurcated anteriorly. Two longitudinal grooves occur on the bony palate. The pterygoid bones are adjacent throughout their length. The supraoccipital 453 453 605 Fig. 245. Skull of the long-snout delphin, Stenella longirostris (figure by М.М. Kondakov). crest is asymmetric: its right half is larger than the left. The lower jaw is slightly longer than the upper. The teeth are conical and number 2=%. Vertebrae 70-75. Phalangeal formula: I,_3, IIg_,, III;_7, IV>-4, and V,-2. The 2nd and 3rd digits are the longest. Ribs, up to 15 pairs. The sternum is complete and T-shaped. These are pelagic forms living on fish. The females are capable of annual reproduction. Gestation extends for roughly 10 months and lac- tation for four months. The females may ovulate and can be fertilized during the lactation period. These animals are distributed in all the oceans and most of the seas in the temperate and warm parts of the world. The genus comprises a single species, the common dolphin, D. delphis Linnaeus, 1758. Some scientists (for example, Nishiwaki, 1972) recognize two more species: D. capensis Gray, 1828 and D. Бата Dall, 1873. These animals could be of economic importance. (V.S.) 606 = © 5 —— | 1 ТИТ и ин ht 4:4 ' Г, о 9 НЕ Nhe 1 wit И И atures AA yyy АМАН НИИНИИ ВОЙ Г ие 1600 2000 3000 4000 5000 km Wey rely! НИИ yu Vi! У al il itil 8 = Fig. 246. Range of the long-snout dolphin, Stenella longirostris (V.A. Arsen’ev). 454 455 456 607 Fig. 247. Long-snout dolphin, Stenella longirostris, at sea (figure by М.М. Kondakoy). COMMON DOLPHIN Delphinus delphis Linnaeus, 1758 1758. Delphinus delphis. Linnaeus. Syst. Nat., ed. X, I, p. 77. Waters of Europe (“Oceano Епгорео”). 1860. Delphinus algeriensis. Loche. Rev. Mag. Zool. Paris, 12, p. 474. репа. 1873. Delphinus bairdii. Dall. Proc. California Acad. Sc., 5, р. 12. Point Arguello, Santa Barbara, California. 1883. Delphinus delphis var. curvirostris. Riggio. Nat. Sicil., 2, p. 158. Mediterranean Sea. 1935. Delphinus delphis ponticus. Barabash-Nikiforov. Byull. Mosk. Obshch. Ispyt. Prirody, 44, p. 249. Yalta, Black Sea. (V.H.) Diagnosis Single species of the genus. Description Well-proportioned animals with a moderately elongated body, distinctly demarcated long beak, and relatively high dorsal fin (Fig. 248). The body color is a combination of black and white and intermediate shades of the same. The upper part of the body is dark-colored and the underside is white; two gray elongated fields and one to three gray bands extend from the zone of the anal opening to the anterior half of the body along the flanks. One dark-colored band runs from the chin to each of 608 ИЕ Ms < ИИ / YI: Wy = Pas LY pope ИИ 7772 456 Fig. 248. Common dolphin, Delphinus delphis (figure by М.М. Kondakov). 457 Fig. 249. Variation in coloration of the common dolphin, Delphinus delphis (figure by N.N. Kondakov). the flippers. A narrow dark-colored strip joins the eyes at the bridge of the nose. The flippers, caudal flukes, and dorsal fin are dark-colored. Color combinations of Black Sea dolphins vary markedly (Fig. 249). Far Eastern common dolphins are differentiated from their Black Sea and Atlantic counterparts by distinctly visible lateral and diagonal bands and by the fact that the dark color of the upper part of the body flanks is sharply demarcated from the light-colored underside (Tomilin, 1957). 457 609 In general, however, coloration is similar among animals from different parts of the range. The skin cover has a thin dermal layer with loose fascicles of collagen fibers. At the boundary with the subcutaneous musculature, elastin fibers and even small fascicles are intertwined. The usual number of vertebrae is: cervical 7, thoracic 14, lumbar 21, and caudal 31: total 73. Cervical vertebrae are often fused. Ribs 13-15 pairs; their number may differ on the left and right sides. An instance has been reported of rudiments of the posterior limbs being detected along the flanks of the urogenital slit among some females caught in the Yalta region (Sleptsov, 1939). The rudiments are in the form of triangular lobes with a height of 3.4 cm (right) and 1.6 cm (left). Each contains two bony members (rudiments of femur and tibia), three cartilages on the right and two on the left (rudiments of tarsus, metatarsus, and phalangeal digits). The main measurements (as percentage of body length) of adult male common dolphins (six) with a body length of 155 to 190 cm and young females (three) with a body length of 142 to 161 cm (Tomilin, 1957) are respectively: distance from tip of snout to anterior edge of dorsal fin 47.1 and 48.0, up to blowhole 17.9 and 19.6 (two measurements each), from anal opening to notch between caudal flukes 27.7 (three measurements) and 28.6 (two measurements), length of flippers, 16.3 and 17.0, maximum width of flippers 6.2 and 6.1, height of dorsal fin 10.3 and 10.1, length of dorsal fin along the base 15.9 and 14.3, and width of caudal flukes from fork to corner 11.8 and 12.3. The most common body length of the Black Sea dolphin is 160 - 170 cm: males 161.8 - 165.8 cm and females 158 - 158.7 cm; maximum length 210-219 cm. Dolphins with a body length exceeding 200 cm are extremely rare: of 38,000 or more dolphins examined, only 26 were longer than 200 cm (Barabash-Nikiforov, 1940). Atlantic and North Pacific common dolphins are larger than their Black Sea counterparts: the maximum body length of the former reaches 258 and 259 cm respectively. Individual variation in skull dimensions (Fig. 250) is very significant (Kleinenberg, 1956a). The height and width of the occipital foramen, size of the parietotemporal fossa, length of the basilar bone [OCHObHbI KOCTD, basilar process], height of the rostrum, as also the number of teeth are highly variable. The total and base length of the skull and the height of the occipital region are relatively stable. Sexual dimorphism of the skull is poorly manifest (Tryuber, 1937). Among males, the skull length relative to body length is less, the cerebral section and rostrum shorter, while the frontal section on the contrary, is greater than among 610 5 YZ Pw \ . ot мА й «Я 2 457 _ Fig. 250. Skull of the common dolphin, Delphinus delphis (figure by М.М. Kondakov). females. With the jaws closed, the rostral portion of the skull of males resembles a broad obtuse wedge while in females it is more pointed and elongated. The main dimensions of the skull (as percentage of total length of skull) of 23 adult males with an average total skull length of 39 cm and 11 adult females with an average skull length of 38 cm (Black Sea; Kleinenberg, 1956a) are respectively: length of frontal section 72.3 and 71.7, cerebral section 27.7 and 28.3, height of occipital region 31.6 and 31.2, width of cranium 35.7 and 35.5, length of rostrum 60.2 and 60.2, width of rostrum at the base 21.0 and 20.6, height of rostrum at the last tooth 8.0 and 7.7, and length of lower jaw 84.7 and 84.7. Age-related changes in the skull (Tryuber, 1937; Barabash-Nikiforov, 458 1940; Tomilin, 1957) are significant. In the newborn, the cranium does not have crests, the skull bones are not completely fused, and the teeth have not cut through. In the first year, sutures between the bones are closed, crests are formed, and the teeth cut through (45 - 48 in each jaw). On taking to independent feeding, the muscular apparatus strengthens due to the greater dimensions of the crests and projections and the rotundity of the skull disappears. Skull growth is most vigorous in the first two years; increase in total length of the skull mainly occurs in the frontal section. At this time the dental system is fully developed. By three years of age, the skull is finally formed. Crests in the skull attain maximum development by four to five years of age. (V.S.) 460 611 Geographic Distribution These animals inhabit the temperate and warm waters of the Atlantic, Indian, and Pacific oceans in both the hemispheres. Geographic Range in the USSR Black Sea, Sea of Japan, Sea of Okhotsk, Pacific Ocean around the Kuril Islands, Bering Sea, waters of southern Kamchatka, and Commander Islands (Fig. 251). These dolphins may be present in our waters of the Baltic Sea since they transgress (albeit rarely) into the southern part of the sea and even up to the estuary of the Visla (Koval’skii, 1964). Transgression into the Barents Sea is also possible. Geographic Range outside the USSR In the North Atlantic, it reaches the coasts of northern Norway, Iceland, the southern coasts of Greenland, and is encountered in Newfoundland and the Gulf of St. Lawrence (Fig. 252). It is quite common in the Mediterranean Sea and is encountered on the Canary Islands and along the entire African coast right up to the Cape of Good Hope and Tristan da Cunha Island. In the western part of the Atlantic, it is known in the Bahamas, the Gulf of Mexico, the Caribbean Sea, and in the waters of South America up to 45° $ lat. It inhabits the entire Indian Ocean and is known in the south along the coasts of Tasmania and Australia. In the Pacific Ocean, it inhabits from Kamchatka and British Columbia south to the Australian and New Zealand coasts and along the coasts of Chile up to 45° $ lat. (V.A.) Geographic Variation This aspect has not been adequately studied. Based on the main dimen- sions and coloration of the animals, a few subspecies have been described. Of these, three inhabit or transgress into the waters of the Soviet Union. 1. Atlantic common dolphin, D. d. delphis Linnaeus, 1758. Largest of the forms, with an average body length of adults at 225 cm. Color variation is very significant. Its presence in the Barents and Baltic seas is possible. Outside the USSR, it is found in the waters of the Atlantic Ccean in the Northern as well as Southern hemisphere. 612 458 lite И Fig. 251. Range of the common dolphin, Delphinus delphis, in the Pacific Ocean waters of the USSR (V.A. Arsen’ev). 2. Black Sea common dolphin, D. 4. ponticus Barabasch, 1935. Smallest in skull and body dimensions; body length of adults averages 204 cm. Color variation relatively minor. Black Sea. Outside the USSR, it is found in the waters of Turkey, Bulgaria, and Rumania.® 3. Pacific common dolphin, D. d. bairdi Dall, 1873. In body dimensions it occupies an intermediate position between the two preceding forms. The average body length of adults is 208 cm; thus it is larger than the Black Sea but smaller than the Atlantic form. Waters of the Pacific Ocean. $ The characteristics of the Black Sea and Atlantic common dolphins have been analyzed and compared. However, no comparison with the Mediterranean populations has been done, which is essential. (V. H.) SH ООО АИ И 1 at af (ny Milage via Perey lity th | | a | В МИ | ee UI al ПИ | | И {lil и И Mh | a hi! И NAL i | 1 И НИ ih ANIM и м Hea и м yh Hi nt | i i | ] rv С hoe И Pye И i OSS Ss, a Sb 0 ПО ar “il | (ty tlaytl ian {1 ЩИ || ait ТТНТ т] Avs ЩИ uh ТИ И | И | Л 2 a | mn i | | || и IP (ih i it т sill ute an eight | Г i НН | ИИ nite a) I ee it | И И НИ и т с И | nH Fig. 252. Range of the common dolphin, Delphinus delphis (V.A. Arsen’ev). 46 — 614 Outside the USSR, it is found throughout the North Pacific Ocean. (V.A.) Biology Population. The total population of the Atlantic and Pacific subspecies has not been established. The population of the Black Sea common dolphin, in spite of the fact that this subspecies inhabits a comparatively small landlocked water body, apparently reached 1.5 to 2 million, with the catch in some years in the Black Sea exceeding 100,000. At present, however, its population is considerably reduced, and probably does not exceed 200,000 to 250,000 animals. Food. The food of the Black Sea common dolphin is fish, crus- taceans, and mollusks. The common dolphin feeds on anchovy (Engraulis encrosicholus), pelagic pipefish of the family Syngnathidae (evidently, Syngnathus schmidti and Sighonostomus typhle), sprat (Sprotella spretus phalaerica), haddock (Gadus euxinus), scad (Trachurus trachurus), mullet (Mullus barbatus), and bluefish (Tomnodon saltator) [Pomatomus salta- trix]. From among crustaceans, large isopods (Idothea algirica) and the shrimp (Crangon crangon) (sometimes stray specimens) form the food of this subspecies; from among mollusks, Nassa recticulata, Mactra subtrun- cata, Guoldia minima, Venus gallina, Colyptera hinensis, Modiola adriatica, Mytilaster sp., and Fellina fabula, constitute the food of this subspecies. Three types of fish form the food base. These are anchovy, pelagic pipefish, and sprat, i.e., small schooling fish often forming large concen- trations. All of the other species are rarely encountered and, in small numbers, in the stomach of this dolphin, constituting a few tenths of a percent of the total food. Dolphins swallow mollusks incidentally during the intake of fish, mainly anchovy. The food of the Black Sea dolphin changes, depending on the region of habitation and the season of the year (Table 35). In winter and early spring months, in the coastal zone, dolphins feed exclusively (or almost 50) оп anchovy. In summer and autumn, pelagic pipefish predominate in terms of frequency of encounter but are of minor significance in the entire food. Sprat in this respect plays a major role, although in terms of frequency of encounter, it is slightly inferior to the pelagic pipefish (Kleinenberg, 1940). In the summer months, the common dolphin forms massive schools in the open sea and the food composition there differs sharply from that established for the coastal zone (Table 36). In the coastal zone, this dolphin feeds on many species of fish but almost exclusively on sprat in the open sea. However, from May through 615 (рзиоэ) с ysyanig € yooppeH р vayjop] ЕТ 12145 8 Aaoyouy ysijodid OL 218 1эа 007 Aacyouy 001 Аоцму — — = = — ЕЕ I yOOoppeHH © Уоррен I вор С vayjop] ysijedid ysyodid ysiedid = == 6S asefad bh ева = SZ 218124 Zz Aaoysuy = — — if yeids ZI yeids SZ 1214$ TZ 1е14$ ysijedid 86 osejag = — — 85 Алоцоцу — 0 Kasoyouy — 06 Алоцоцу — 81 Алоцоиу = $Х$185010^0М @ vaylop] = = == 0OT — Aaoyouy 86 Алоцоиу 00 — Алоцоиу uoyy лом С payjiop] = = 86 Asoysuy = QOT_=— Aaoyouy 001 Алоцочу = goT.—Ss Ano youy rune g Sa a a ee ao a a А % wot % шэи 9% wo}jl % Wo}jl % two % mol poof poo poo poo poo poo ounr Key judy Че Areniqa.J Атепчег UOIBdYy (egc61 ‘ЗлэчиэщэГУ) % ‘vag Чо эцу Jo эпох [е}5е0э ay) UY ищаюр поинио» ay} Jo poog ‘sE эает, Z9p % 616 TINA yooppeH vaylop] ysijadid 218 за yooppeH yeids Aaoyouy ysyadid o18e[ag ша poo 12490120 58 % paylop] 1214$ ysijadid 18 |э4 Ч$9эщ& yooppeH] Aaoyouy ysijodid 218 |эа wo! poo Jaquiaidag яя + 98 % yooppeH paylopy yeids Aaoyouy ysiedid 218 1э4 Ч$3211Я peos Aaoyouy ystjadid 218 |э4 шэи poo isnsny NAA 85 % Ч$3эЯ yooppeH вом yeids Aaoyouy ysijodid o18e[9g yooppeH Ppeos yeids AAoyouy’ ysijedid 218124 э poo Aqne eer YSIISSOIOAON ‘pluoo 5$ этавт, 617 462 Table 36. Food of the common dolphin in the coastal zone of northern Caucasus and away from coasts (Tsalkin, 1938a) Percentage of total number of items detected In coastal zone Away from coasts (S.E. Kleinenberg) Pelagic pipefish 72.0 — Anchovy 15.0 0.1 Sprat =— 99.6 Haddock 1.0 0.3 Idothea (slater) LES — Scad 8.0 — Mullet — — Bluefish 2.0 — August, sprat serves as the main food not only in the open sea, but also in the coastal zone, constituting, on average, over 90% of the food for the season as a whole. In various regions, the age composition of sprat varies sharply in different months. In May, the stomach of the dolphins in the coastal zone as well as in the open sea contained both adult and young sprat, while in June in the open sea, the preference was for young sprat and in the coastal zone of Crimea (Alushta) and the Caucasus (Gelendzhik), mainly the adult fish. In July and August, the stomach of all the dolphins from all the regions contained mainly young sprat. Thus young sprat constituted the maximum specific proportion in the food of these dolphins in the summer months (Tarasevich, 1958b). The seasonal change in the food of the Black Sea common dolphin is explained by the biological features of the fish consumed by it. Sprat is a cryophilic fish and its spawning continues throughout the winter and spring months. In the period of spawning, sprat is scattered over immense pelagic regions of the sea and does not form significant concentrations. In summer, however, young sprat as well as adult fish remain sufficiently concentrated and hence serve as the main food of these dolphins at this time. Anchovy, on the other hand, is a thermophilic fish and spawns in the summer months. At this time it is also scattered over a large expanse in the coastal zone as well as in the open sea and nowhere forms notice- able concentrations. In the winter months, anchovy gathers at certain places, 1.е., in regions of steep precipitous drops of coastal waters. In December, massive collections of anchovy form almost simultaneously along the southern coasts of Crimea (Balaklava-Sarych region) and the Caucasus (Poti-Batumi region). It is in this period that anchovy serves as the main food of these dolphins (Freiman, 1950). 463 618 Thus the main food items of the common dolphin lead a pelagic way of life, are relatively small in size, and form dense concentrations. Information on the food of Atlantic and Pacific subspecies of the common dolphin is very scant. Fish apparently serve as their main suste- nance. Sardines, flying fish, anchovy, herring, mackerel, and gray mullets have been indicated as food items. Sometimes crustaceans and mollusks (bivalves and cephalopods) are encountered. In a dolphin caught close to Corsica, the stomach contained four specimens of Enoploteuthis mar- garitifera and three of Chirteuthis veranyi in addition to Loligo vulgaris, Todarodes sagittatus, Onychoteuthis lichtensteini, Heteroteuthis dispar (?), and three squids identified as Ctenopteryx cyprinoides (Tomilin, 1957). The stomach of a dolphin caught on May 22, 1957 in Newfoundland contained the remains of 20 Atlantic shortfin squids, Шех illecebrosus, which are quite numerous in these waters in summer (Sergeant, 1959b). Daily activity and behavior. No changes have been observed in the daily activity of dolphins. Fast and impetuous movements are charac- teristic of the common dolphin, as in the case of many other species. Feeding mainly on pelagic fish, the common dolphin does not go very deep and short respiratory intervals are characteristic. It usually emerges onto the water surface every 0.5 -1 min, less often every 1.5-3 min. It can remain submerged in water for a maximum of 5 min. The duration on the surface is reckoned at a few tenths of a second. The common dolphin does not produce blows and splashes water sideways only slightly. Foraging dolphins move slowly and submerge steeply in the water. When feeding on fish schools confined roughly at one place, the dolphins dive almost vertically. They cannot perhaps dive deeper than 70 m since they never descend below the lower seine ropes of 70-m high fishing nets. A “traveling” dolphin swims rapidly, easily overtaking ships sailing at 15-18 miles an hour. It swims right on the surface of the water and at short intervals (at the time of inhalation/exhalation) leaps powerfully out of the water, often over a span of up to 10 m (Fig. 253). The habit of dolphins running behind ships is interesting. Small schools very often accompany a ship for a few hours, sometimes on one side, sometimes on the other, straying off and again rapidly closing in. Dolphins usually love to play before the very stem, leaping out of the water into the surf formed by the ship. When moving in schools, the dolphins do not leap simultaneously; thus at any given moment a few animals can be seen on the surface of the sea (Kleinenberg, 1956a). The females are greatly attached to their calves. Two females swam for a long time around a sweep net enclosing a water body in which their calves were held. 463 464 619 Fig. 253. Leaping common dolphins (photograph by S.E. Kleinenberg). The auditory faculty is well developed among dolphins. Common dolphins are frightened of sharp sounds, which in fact is taken advantage of when trapping them in sweep nets. A school of dolphins is chased away from the wall of the net using a “telephone”, 1.е., striking stones one against the other in the water (Tomilin, 1957). Vision is not as well developed as hearing. Dolphins can probably see in water no more than a few tens of meters. They produce extremely diverse sounds—whistling, squeaking, and crackling. Some sounds are produced at high frequencies, beyond the range of human perception. Such sounds are used by the dolphin for echolocation, enabling excellent self-orientation in an aquatic environment. The age and sex composition of schools of dolphins vary at different times of the year. In winter, two types of schools have been noticed. The first type, called the winter female groups, consists of 40 to 50% mature females, some gestating, some barren, and those that have only recently attained maturity (lactating females are very few) and 50 to 60% immature animals of both sexes. Immature females in these schools represent all age groups while immature males are usually young. Much larger males, approaching maturity, are found in small numbers. Schools of the second type are called male groups and contain predominantly males, to the extent of 90% of the total number of animals in a group in January-February and about 80% in March-April. The females in these schools, at best, account for 10-20% and comprise mainly the barren with only 4-5% being immature. Most of the males in the school are 620 mature animals while the immature are mainly represented by older age . groups. х The following types of schools have been established for the spring- summer months: with the approach of the en masse calving season (March-April), the gestating females begin to separate from the female schools and gradually form individual precalving schools. The maximum number of such schools is formed in May and June but the precalving schools invariably contain immature animals too, which sometimes constitute up to 30% and even 45% of the school’s strength. As the gestating females leave the winter female schools, the remain- ing dolphins form independent groups. In some cases, they consist of mainly the immature animals of both sexes and may be called schools of immature dolphins. Such schools formed in March represent long-time formations and are seen later throughout the year. If, however, the win- ter female schools comprise mainly young and barren females, with only a few gestating ones, after the separation of the latter, residual female groups are formed with predominantly barren and just recently matured females. These groups remain active for a short duration, i.e., only in early spring. Females of the Black Sea common dolphin in the period preceding calving concentrate in regions with more favorable meteorological con- ditions, in the so-called calm zone. The animals find such conditions in the open sea far away from the coasts. It is here that precalving groups remain and are gradually transformed into schools of calves. The period of parturition is very prolonged in this dolphin and hence the number of gestating females in the precalving groups diminishes gradually while the number of lactating females concomitantly increases. Later, the lactating females begin to predominate and such groups can be called nurseries. Nevertheless, they invariably contain immature dolphins too and their relative proportion is even slightly more than in the precalving groups. Schools of calves are encountered mainly in the second half of June and in July. In August, and more so in September, they become quite rare. Females begin to gather with the winter male schools in spring and by April-May such groups are transformed into breeding schools consisting of 60-80% mature males and females in roughly an equal proportion. Such groups are most often seen in the period of en masse mating (in July and August) and hence are called breeding schools. In the early spring months, they are formed by the merger of male and female groups (as soon as the gestating females have left the latter). Therefore, at this time the breeding schools contain only barren and recently matured females. In the summer months, lactating females join them in large numbers 465 621 and in July and August, form the bulk of the females in the school. The growing juveniles form the majority of immature animals present in the breeding schools. : The process of reformation of summer into autumn schools and the order of formation of winter types of common dolphin schools of the Black Sea population has not been traced (Tarasevich, 1951). The above extremely schematic description reflects the process of formation of schools of different types and their transformation from one type into another. In nature, this process is far more complex. Such a diversity of schools and the numerous transformations which they undergo in the course of a year with respect to age and sex com- position, compel us to estimate very carefully the percentage ratio of different age and sex groups in a school of dolphins using the data of hunters. Not only the time, but also the region of catch should be taken into consideration. The common dolphin has been observed amidst groups of other species of dolphins, such as the bottlenose dolphin, the shorthead dol- phin, and even the pilot whale. The common dolphin does not take very kindly to captivity compared to the other species and usually does not survive for more than two or three months in ocean aquariums. Its reproduction in captivity is not known (Tomilin, 1962). Seasonal migrations and transgressions. The nature of seasonal migrations among Atlantic and Pacific subspecies has not been established. This aspect has been studied somewhat better for the Black Sea subspecies and it has been assumed that the common dolphin in general does not perform seasonal migrations, instead its movements should be regarded as migrations in search of food (Tomilin, 1957). These assumptions give rise to serious doubts, however. The common dolphin is a typical pelagic animal avoiding sections with fresh and turbid waters and evidently, therefore, does not enter the Azov Sea (Freiman, 1950). It does not dive very deep in search of food but is satisfied with what is available in the upper horizons of the sea. It inhabits almost the whole of the Black Sea, in the coastal zone of Crimea, the Caucasus, Turkey, Bulgaria, and Rumania and also far away from the coasts. But its distribution in the equatorial seas is very uneven. In winter months, in the period of spawning, sprat is scattered over a large water body and is almost not utilized as food by this dolphin. At this time the Black Sea anchovy concentrations are formed at the wintering sites in the coastal waters of Georgia (Poti-Batumi) and on the southern coast of Crimea (Balaklava). The presence of anchovy determines the wintering regions of the common dolphin, most of which concentrate on the coasts of Georgia and a small percentage south of the Crimean 466 622 peninsula. In spring, as the water begins to warm up, anchovy begin to spawn and are scattered over a large expanse without forming concentra- tions. Simultaneously, adult sprat and their young begin to concentrate in the coastal waters of Crimea and northern Caucasus and in the open northeastern waters of the Black Sea. Dolphins in search of food at this time move northwest where they encounter sufficient collections of sprat on which they feed in summer. Depending on the collections of food objects, the common dolphin forms concentrations of different types: (1) encountered very rarely—only a few animals or small schools separated from each other; (2) encountered rarely—some animals live everywhere while schools are small and sparse; (3) encountered very frequently—numerous small- and medium-size schools (up to 100 animals each) predominate, large schools are very rarely seen; (4) encountered very often—large number of predominantly large schools, of the order of hundreds or thousands of animals in each, forming dense congregations in an extremely small water body (Tsalkin, 1938). These dolphins usually form congregations in the Tuapse-Sochi region along the coasts of the Caucasus; they may be closer or farther away from the coasts (20 to 60 miles) in different years. These are not long-time schools, surviving for just one to three months. Simultaneous with them, schools of different strengths and some individual dolphins are very widely distributed along the Black Sea in the coastal zone as well as in the open sea, depending on the availability of food. In some years, the distribution of the groups and much smaller formations changes considerably according to seasonal conditions (Tarasevich, 1958a). As the dispersal of sprat concentrations increases, schools of dol- phins begin to abandon the regions of summer habitation and gradu- ally gather at the wintering sites where they live throughout the win- ter months. The migrations of the Black Sea common dolphin are very small but they bear a distinctly manifest seasonal character and are quite constant in time and direction. There is every reason to consider them seasonal migrations, therefore, at least in the eastern part of the Black Sea. The periods and nature of migrations of the common dolphin in the western part of the sea have yet to be established. In the Bay of Bosporus, schools of dolphins swimming in different directions are often noticed. It is possible that the common dolphin sometimes leaves the Black Sea for the Sea of Marmara or, contrarily, transgresses from the Sea of Marmara into the Black Sea, but there is no recorded proof of this (Freiman, 1950). It has been assumed that a more intense exchange prevails between the Black Sea and the Mediterranean 623 Sea dolphins than is usually acknowledged, which might be reflected to some extent in the population dynamics of the dolphins of the Black Sea. Reproduction. Mating and calving in the common dolphin are highly prolonged but there is no one single view on their duration. Vari- ous sources indicate that mating extends from June through October (Sleptsov, 1941), from July to December (Tomilin, 1957), and even from August to January (Maiorova and Danilevskii, 1934). But assumably most of these dolphins mate over a much shorter duration—in July, August, and the first half of September (Sleptsov, 1941); from August to October with peak activity in September (Tomilin, 1957); or in July (Kleinenberg, 1956a). The period of calving, according to various authors, extends from May through November, peaking in June and July (Sleptsov, 1941); from May to October or from June to November, the majority of females calve from June to August, with maximum reached in July (Tomilin, 1957); or from May through September with the peak in May and June (Kleinen- berg, 1956a). In any case, in most females parturition takes place in the summer months. Mating of the common dolphin has been observed time and again from airplanes and hunting craft. A group of mating dolphins consists of one female and six to eight males which pursue her at high speed, chase each other, and hold the competitor behind the fins with the teeth. When the fastest male catches up with the female, she turns her abdomen upward, mates with him, and the two then disappear into the water (Golenchenko, 1949b). The duration of gestation has also been variously determined: about 10 months (Tomilin, 1957), 10-11 months (Kleinenberg, 1956a), and 11 months (Sleptsov, 1941). The question of the periodicity of calving, 1.е., the rate of replen- ishment of the population, has not yet been conclusively established. и was first thought that dolphins produced calves once every four years or twice in three years (Mal’m, 1936). According to M.M. Sleptsov (1941), dolphins tend to calve annually but since fertilization can occur only 1.5 to 2 months after birth, mating advances in time every year and the female becomes ready for fertilization afresh only after the males cease to mate. From this, it may be assumed that the female dolphins calve three years in a row but remain barren in the fourth. If this is so, then one out of four of the eligible females should remain barren every year (Kleinenberg, 1956a). According to V.E. Sokolov (1954), a calved female becomes capable of mating again within three to five weeks after par- turition, 1.е., even in the period of lactation, which would indicate the absence of advancement in the period of mating in subsequent years. Thus this dolphin usually reproduces every year and the percentage of 467 467 624 barren females in a group is not high (У.Е. Sokolov, 1954). Оп this basis, it has been assumed that the sex cycle of the dolphin fits in the annual cycle, with a gestation period of 10 months (Tomilin, 1957). More reliable data are required before the question of the reproduction cycle of the Black Sea common dolphin can be conclusively resolved. Growth and development. The uterine horns and ovaries are usually asymmetrical in the common dolphin; in 82.3% of cases, the left horn was wider and longer than the right one while the left ovary was larger; in 17.7% of cases, the right horn and the right ovary were larger. Probably, in an overwhelming majority of cases, the embryo (Fig. 254) grows in the left uterine horn (Sleptsov, 1941). The newborn calf of the common dolphin reaches, on average, half the body length of the mother (sometimes more), which apparently is far more than the corresponding dimensions of newborns among other whales. The average body length of the newborn is 82-90 cm (Kleinen- berg, 1956a). As a rule, one calf is born and twins are rare. The calf is born not head first, as thought earlier, but tail first. The dorsal fin of the embryo is convoluted and compactly pressed to the body and the caudal flukes are balled into a “fist” (Kleinenberg, 1956a). Lactation extends for about four months (Tomilin, 1957) or five to six months (Sleptsov, 1941; Kleinenberg, 1956a). At the end of lactation, the stomach of calves contained fish remains along with milk. Milk pro- duction in the mammary glands of females is very little; thus a 168-cm long female could yield only 100 ml (Tomilin, 1957). Apparently the calf feeds often but in small portions. The composition of the milk of the common dolphin is shown in Table 37. Fig. 254. Embryo of the common dolphin (figure by N.N. Kondakov). 468 625 Table 37. Chemical composition of the milk of the Black Sea common dolphin (Ural’skaya, 1957) Composition of milk, % Author and i Se a ht te I yeaniog Fat Protein Sugar Dry Ash Water Та analysis matter 43.71 5.62 1.45 7.53 0.45 48.76 100 Mal’m, 1932 41.56 48 1.49 6.82 0.45 51.62 100 Ural’skaya, 1954 The growth of the common dolphin in different age groups has not been studied well. By autumn, the calf reaches a length of 100-110 cm, 1.е., in the period of lactation, it adds 20-30 cm. With cessation of feeding on mother’s milk (winter), growth slows down and by the next summer (toward the end of the first year) the dolphins reach a length of 120-130 cm. The average dimensions of two-year-olds is 140-150 cm and three-year-olds 165 - 170 cm. The length of most adult dolphins varies around 170 cm and their average weight 50-55 kg. Sometimes animals up to 200 cm long and 100 kg weight are encountered (Freiman, 1950). The common dolphin is characterized by sexual dimorphism: males are slightly larger than females right from the embryonic stage. There is no unanimity of opinion regarding the time required to attain sexual maturity. According to some data, female common dolphins attain maturity in the third year and males in the fourth year (Sleptsov, 1941). According to other authors, females become mature at the end of the third year and males in the third or the fourth year (Kleinenberg, 1956a). According to still others, the common dolphin is capable of mating for the first time in the second year of life (Tomilin, 1957). The body length of these dolphins at the time of attaining maturity varies considerably. Mature females 140-150 cm long are encountered while some 161-170 cm long are still immature (Kleinenberg, 1956a). With a high degree of probability, it may be said that females with a body length of less than 155 cm should be regarded as immature and those longer than 155 cm as mature. This limit for males is put at a body length of 165 cm. The maximum longevity of the common dolphin has been roughly put at 20-25 years (Sleptsov, 1941; Freiman, 1950). Enemies, diseases, mortality, parasites, and competitors. The Black Sea common dolphin has no enemies. Subspecies of the Atlantic and Pacific oceans may become prey to the killer whale. There has been almost no study of the diseases afflicting this‘dolphin. Some instances of stones in the ureter (Kleinenberg, 1956a) and of skeletal diseases (Tomilin, 1957) 469 626 are known. From among skin parasites, crustaceans (Xenobalanus glo- bicitis, Penella pustulosa, and Larnaeonema nodicornis) have been found. Nineteen species of endoparasites are known: these include trema- todes (six), cestodes (three), nematodes (six), and acanthocephalans (four). The trematode Brauniina cordiformis Wolf, 1903 is a parasite of the stomach and intestine of the common dolphin, the bottlenose dolphin and some species of spotted dolphins. Three species of trema- todes of the genus Campula have been found in the bile duct and hepatic ducts of just the common dolphin: C. delphini Peirier, in individuals from the European waters of the Atlantic; C. palliata Looss, in the Atlantic subspecies (Europe) and Black Sea residents, and C. rochebruni Peirier, in specimens from the Atlantic waters of Europe. The trematode Dis- tomum philocholum Creplin, which localizes in the liver, was detected in the common dolphin from European waters. The trematode Galac- tosomum erinaceum Peirier was found in the intestine of the common dolphin also only from European waters. The cestode Tetrabothrium fosteri is a parasite of the intestine of the common dolphin and beaked whales. It has been found in the Mediterranean Sea and in the southern part of the Pacific Ocean. Monorygma grimaldii Monier localizes in the abdominal cavity, mesentery, and diaphragm of the common dolphin and four other species of dolphins of the Mediterranean Sea and the Atlantic Ocean. The cestode Phyllobothrium delphini Bosc is quite extensively distributed among marine mammals. In addition to the common dolphin, it has been detected in six species of toothed whales, the bowhead whale, and Weddell’s seal. It localizes in the skin and hypodermic tissue and has been reported from the Atlantic Ocean, Mediterranean Sea, northern and southern parts of the Pacific Ocean, and in the Antarctic. One of the most widely distributed species of nematode parasites, Anisakis (Anisakis) simplex Rudolphi, infects the gullet, stomach, and intestine. Apart from the common dolphin, it has also been detected in ten species of toothed whales, two species of baleen whales, and in Steller’s sea lion. It has been reported in the North Sea and in different parts of the Pacific Ocean (Kamchatka, Japan, and New Zealand). Among the common and three other species of dolphins, Anisakis (Anisakis) typica Diesing is a parasite of the stomach while Anisakis dussmeerii Beneden has been found in the stomach and large intestine of only the common dolphin. The nematode Halocercus (Halocercus) delphini Baylis and Daubney was detected in the bronchi of the common dolphin only from the Atlantic Ocean while another representative of this genus, Halocercus (Posthalocercus) kleinenbergi Delamure, was found in the lungs of only the Black Sea common dolphin. Yet another species 627 of nematode, Skrjabinalius cryptocephalus Delamure,-was found in the lungs of only the Black Sea common dolphin. Of the four species of acanthocephalans, Bolbosoma vasculosum Rudolphi is a parasite of the intestine of common dolphins and beaked whales from the Mediterranean Sea and the Atlantic Ocean. Bolbosoma bellicidum Leuckart has been reported in the common dolphin. Coryno- soma cetaceum Johnston, a parasite of the intestine of common and bot- tlenose dolphins, was detected in Australian waters. The specific affinity of the fourth representative of acanthocephalans, Corynosoma sp., has not been established (Delamure, 1955; Tomilin, 1962). Natural mortality has not been studied. It appears possible that the common dolphin could die from an invasion of the nematode Skrjabi- nalius cryptocephalus, which has infected the lungs of some animals to such a degree that they were totally damaged (Kleinenberg, 1956a). There are almost no competitors of the Black Sea common dolphin. The Atlantic and Pacific subspecies may be food competitors of other species of dolphins. Population dynamics. In the prewar years, when the total catch of the common dolphin in the Black Sea sometimes exceeded 100,000 animals per year, a distinct reduction in population was noticed. During the years of the Great Patriotic War [WW II], hunting activity perforce declined and the population recouped slightly. Subsequently hunting resumed and the population decimation was such that hunting these dolphins was banned. Information on the population of the common dolphin in other parts of its range is not available. Field characteristics. This is a rather small (body length of adults 150-170 cm) agile dolphin having a high pointed dorsal fin with a crescent-shaped notch in the posterior margin [falcate]. The beak is long and sharply demarcated from the convex corpus adiposum [“melon’]. The dorsum, forehead, and upper caudal crest are black with two long gray fields stretching along the flanks below these black fields; the ven- tral side is white. This animal does not blow fountains [1е. its blows are thin] and exhibits much of its body in normal movements. It often leaps completely clear of the water. In rapid movement, white surf forms along the body flanks. Schools of different strengths are common. Some- times the herds merge into congregations of many thousands of animals. (V.A.) Economic Importance Only the Black Sea common dolphin has been of commercial importance and in times past was caught in large numbers. In all the remaining 470 628 sections of its range, this species has never attracted special interest and was caught in very small numbers. Hunting the Black Sea dolphin is centuries old. For many years Turkish hunters used an extremely primitive method in killing them. Hunting activity intensified only after the October Revolution. The first cartel for this purpose was organized in 1929 in the Crimea. At this time, along with the then-prevailing method of hunting with firearms, dolphins were also caught in sweep nets. Hunting with firearms was carried Out from ships using a smooth-barreled gun and buckshot. Since in summer the killed dolphins sank rapidly, expert divers were always onboard who jumped into the water simultaneous with the shot and held the killed dolphin afloat. Nevertheless, many killed dolphins sank and others who were injured probably died subsequently. Therefore hunting with firearms was ultimately banned and further development proceeded by perfecting the use of sweep nets. After several improvements in catching dolphins in sweep nets in the Black Sea, the following method was adopted. The net, made of fabric, was up to 500 m long (sometimes up to 1,500 m), wall height 60-80 m, and mesh size (diagonally) about 15 cm. Special rings were provided on the lower seine rope of the net through which a rope was run for tighten- ing the bottom of the net (like a purse). A brigade of catchers comprised 30-40 men equipped with one seiner, two or three motorboats, and up to 10 feluccas. The net was held on the stern of the seiner, cast by the mov- ing ship, and drawn up from the water mechanically. The feluccas going out for the catch and returning to the coast, were towed by motorboats. Each felucca had two oarsmen with a sufficient stock of stones. The seiner and another boat approached a school of dolphins from two sides with a comparatively long distance between them, leaving the feluccas with the oarsmen at equal intervals on the trail. Having swept the school from three sides in the form of a horseshoe, they blocked the fourth side with a small sweep net. Striking stones against each other (“telephoning’’) and shouting from the feluccas left along the spread out net, the catchers prevented the dolphins from leaving it until the ends of the net could be drawn together. To prevent the net from sagging or twisting in the water current, the hunters on the feluccas positioned along the entire course of the net held onto its upper edge and flexed it in different directions. By drawing the rope running through the rings on the lower side, a “purse” was formed from which the dolphins caught were gathered in by the feluccas and later loaded on the seiner for trans- porting to the coast. Hundreds of dolphins were often caught in one casting of the net. If another school of dolphins happened to be sighted in the proximity, the brigade began casting a new net without giving up 471 629 Table 38. Catch of dolphins in the Black Sea (Bodrov, Grigor’ev, and Tver’yanovich, 1958) Year Number of Year Number of animals caught animals caught 1931 36,490 1945 3,464 1932 53,858 1946 15,872 1933 67,469 1947 19,400 1934 67,065 1948 20,863 1935 70,448 1949 32,200 1936 62,933 1950 42,535 1937 103,814 1951 20,250 1938 147,653 1952 50,618 1939 81,206 1953 42,757 1940 71,097 1954 82,809 1941 40,250 1955 18,354 1942 1,157 1956 16,082 1943 2,896 1957 29,800 1944 441 the first catch. Thus the brigades sometimes returned to the coast with catches in several nets. Dolphin hunting was successfully developed in Bulgaria later. The results of erstwhile dolphin hunts in the Black Sea (in the Soviet Union) are shown in Table 38. When the catch rapidly diminished in subsequent years, hunting of the Black Sea dolphin was discontinued. Statistics of the catch of the Black Sea dolphin provide no specieswise classification and hence the proportion of the common dolphin is not ascertainable. The ratio of the different species of dolphins in the Black Sea hunts varied considerably in different years but the following figures could be regarded as averages: number of common dolphin, common porpoise (Phocoena), and bottlenose dolphin (Tursiops) = 200:10:1. Thus the common dolphin invariably represented the bulk of the animals caught (Tsalkin, 1937). The average weight of a whole dolphin according to long-term data is 51 kg; thus the quantum of products recovered from a dolphin is relatively small. Weight of body portions and organs of a small female Black sea common dolphin, kg (Tomilin, 1957) Total weight Sy Subcutaneous fat 10.98 Hump flesh 3.85 630 Caudal musculature 2.50 Backbone 2.55 Ribs with musculature in-between 1.85 Adipose body [“‘melon’’] 0.52 Dorsal fin 0.25 Flippers (two) 0.47 Caudal flukes 0.44 Lower jaw 0.48 Tongue Ой Brain 0.67 Intestine 0.97 Gullet 0.23 Heart 0.17 Liver 0.60 Lungs (two with larynx) 1.00 Kidneys (two) 0.19 Stomach 0.20 Skull, blood, etc. 3.91 Dimensions and weight of common dolphins (18 complete and 12 partial weighings) (Dragunov and Kasinova, 1954) Body length, cm 119-190 x (159) Total weight, kg 22.0-90.5 @:=9950:0) % of total body weight Blubber 26.0 - 44.8 (< = 34.4) Carcass without head and 30.0 - 46.8 (x = 37.4) viscera Head 8.1-14.8 (x = 10.6) Tail and fins 25-52 (9 Vicera (entire) 9.2 - 16.2 (х = 12.5) Flesh with carcass 18.6 - 24.8 (x = 23.4) Bones of carcass (not cleaned) 10.3 - 17.0 (x = 14.0) Liver 1.4-3.3 12 Heart 0.39 - 0.86 (= 055) Lungs 1.9-4.1 = Stomach (without contents) O7 1:3 C= 10 Intestine (with contents) Zale @="'3'8) Kidneys 0.40 - 1.02 (= 065) Spleen 0.05 - 0.18 (> = 0.10) ‘Testes 0.5-1.8 G@ = 14) 472 631 Brain 1120 (GE E15) Jaw fat 0.15 - 0.34 (=2N0:25) Blood and unaccounted losses 0105-53 C= 91.2) The amount of subcutaneous fat varies significantly depending on the well-being of the animal. In the period of its maximum fattiness (in April or March), the killed dolphin does not sink. The fat in the animal is minimal in August. The fat recovered from the blubber is used in the paint industry, etc. The better quality fat is used as a substitute for medicinal cod liver oil. The hollow lower jaw of the common dolphin contains a small amount of very valuable fat of a different chemical composition. The oil processed from this fat does not solidify in the cold and is used for oiling fine mechanisms. The carcass of the dolphin is used as raw material for producing meat-bone meal for feeding farm animals. High-quality glue is also produced from this raw material. The skin of this dolphin, after appropriate processing, can be used in the shoe industry for making leather boots. From the viewpoint of the most rational use of the raw material, dolphin hunting is best carried out in the spring and winter months, in the period of maximum fattiness of the animals. With the same number of animals caught then, a greater quantity of products can be recovered. Commencing from 1965, every type of hunting of the Black Sea dolphins has been banned and all the three species inhabiting this water body have been brought under conservation. Similar measures have been adopted in Bulgaria and Rumania. (V.A.) Genus of Bottlenose Dolphins Genus Tursiops Gervais, 1855 1843. Tursio. Gray. List Mamm. Brit. Mus., p.p. XXIII, 105. Delphinus truncatus Montagu, 1821, nom. praeocc. 1855. Tursiops. Gervais. Hist. Nat. Mamm., 2, p. 323. substituted for Tursio Gray, 1843. 1873. Hemisyntrachelus. Brandt. Мет. Ac. Imp. Sc., Pétersbourg. Sub- stituted for Tursio Gray, 1843. (V.H.) Medium-size dolphins with a maximum body length of 3.9 m. The body build is somewhat heavy, with a “beak” of medium length. The high dorsal fin has a fairly deep notch along the posterior margin. The flippers are relatively broad. The lower jaw is slightly longer than the upper one. The upper portion of the body is dark gray (sometimes light gray), the flanks gray, and the abdomen white (sometimes gray). 473 632 The rostrum of the skull is medium in length, longer and narrower in females than in males. The premaxillae and nasal bones are adjacent on the right side of the skull but not on the left. Broad processes join the pterygoid bones which have an oblique notch posteriorly. The teeth are fairly large (up to 10 mm in diameter) and number 15—28 = 76-106. They are often completely worn down. The sternum consists of three fused sections. Vertebrae 63-65. Ribs 12-13 pairs. Phalangeal formula: I,-2, U7-9, Шб-в› [\>-5, and V,_>. The 2nd, sometimes the 3rd digit, is the longest. These dolphins are bentho-ichthyophagous. Periods of mating and parturition are prolonged. Gestation extends for 11 months while lacta- tion apparently continues for 4-6 months. These dolphins are distributed in the Atlantic Ocean from the North and Norwegian seas to the Mediterranean and Black seas and South Africa in the east and from southern Greenland to Patagonia in the west; in the Pacific Ocean, from California in the east and Japan in the west to Chile and Australia and New Zealand; and, in the Indian Ocean, from Australia to Africa. Fossils have been traced to the Upper Pliocene of Europe. The genus comprises a single species, the bottlenose dolphin, Т. truncatus Montagu, 1821. Sometimes, one other species, Т. gilli Dall, 1873, or even six more species are recognized. Apart from T. gilli, the five others are Т. nuuanu Andrews, 1911; Т. aduncas Ehrenberg, 1833; T. parvimanus Reinhardt, 1888; Т. gephyreus Lahille, 1908; and Т. abusalam Ruppell, 1842. (V.S.) BOTTLENOSE DOLPHIN Tursiops truncatus Montagu, 1821 1780. Delphinus tursio. Fabricius. Fauna groenlandica, p. 49, Greenland. The name has no nomenclatural importance since its applicability to this species could not be established. 1804. Delphinus nesarnak. Lacépéde. Hist. Nat. Cétacées, р. XLII, 307. Northern Atlantic (nom. ргаеосс). 1821. Delphinus truncatus. Montagu. Mém. Wernerian Nat. Hist. Soc. 3, p. 75. Devonshire, England. 1832. Delphinus aduncus. Ehrenberg. In: Hemprich et Ehrenberg. Sym- bolae physicae, Mammalia, 2. Red Sea. 1873. Tursiops gillii. Dall. Proc. California Ac. Sc., 5, p. 13. Monterey, California. 1911. Tursiops nuuanu. Andrews. Bull. Amer. Mus. Nat. Hist., 30, p. 233. Gulf of California. 633 1940. Tursiops truncatus ponticus. Barabash-Nikiforov. Cetacean fauna. Black Sea, p. 56. Novorossiisk, Black Sea. (V.H.) Diagnosis Only species of the genus. Description Two color groups are recognized among bottlenose dolphins of the Black Sea (Barabash-Nikiforov, 1940, 1960). Type A is characterized by a fairly distinct boundary between the dark color of the dorsum and the white color of the abdomen with a light-colored angular patch in the dark field midbody; the apex of the patch is turned toward the dorsal fin (Fig. 255). In type B, the boundary between the pigmented upper and lower surfaces is not distinct and appears as vague straight, wavy, or broken line, without the light-colored angle near the dorsal fin. The quantitative ratio of animals of the two types varies in different years. Among 50% of the bottlenose dolphins of the two types, a frontopectoral line joins the eyes and gradually extends from their outer corners toward the flippers. Among the bottlenose dolphins of the Atlantic, the light- colored patch characteristic of type A and the frontopectoral line are invariably absent. The latter can, however, be seen in the dolphins of the Mediterranean Sea. Cervical vertebrae 7, thoracic 12-14, lumbar 17, and caudal 26-27. The cervical vertebrae may be fused in various combinations. The main body dimensions (as percentage of body length) of bottle- nose dolphins averaged for 52 adult animals with a body length of 270-310 cm (Barabash-Nikiforov, 1940) are: distance from tip of lower jaw to base of flippers 18.5, from tip of upper jaw to blowhole 14.0, from 473 Fig. 255. Bottlenose dolphin, Tursiops truncatus (figure by N.N. Kondakov). 474 634 Ир of upper jaw to commencement of dorsal fin 42.7; length of flippers 14.6; length of dorsal fin 15.6; and width of the left caudal fluke 9.6. The average length of males caught in the Black Sea was 228 cm, of females 222 cm; the largest male caught was 310 cm long. Bottlenose dolphins from other regions are larger than their Black Sea counter- parts: in the waters of Japan, the males average a length of 2.7 m and females 2.8 m; bottlenose dolphins in the Atlantic Ocean attain a length of 3.1-3.3 m and in the Mediterranean Sea 3.2 m. With advancing age, a relative reduction (as percentage of body length) is noticed in the length of the oral slit, length of flippers, and distance from tip of snout to eye and up to the flippers (Barabash-Nikiforov, 1940). The average main skull measurements of 10 male bottlenose dolphins from the Black Sea with a body length of 180-310 cm and five females with a body length of 214-234 cm (Tomilin, 1957) are. respectively: condylobasal length 44 and 44, zygomatic width 23 and 22, length of rostrum 24 and 24, width of rostrum at base 12 and 12, and length of lower jaw 36 (three measurements) and 40 (two measurements). The skull of the Black Sea bottlenose dolphin is smaller compared to bottlenose dolphins residing in other seas (Fig. 257). Vertebral sections comprise (as percentage of length): cervical 3, thoracic 23, lumbar 30, and caudal 44 (Slijper, 1936). (V.S.) Geographic Distribution Seas of the temperate and warm belts in the Northern and Southern hemispheres. Geographic Range in the USSR Baltic Sea from Gulfs of Riga and Finland, Barents Sea east to Novaya Zemlya, and the Black Sea; Pacific Ocean Basin, the Sea of Japan, south- ern part of the Sea of Okhotsk, and waters of the southern half of the Kuril range (Fig. 258). Geographic Range outside the USSR In the Atlantic Ocean, in the zone of influence of the Gulf Stream, the range extends north up to the Lofoten Islands on the coasts of Nor- way, coasts of Iceland, Jan Mayen, and south of Greenland (Fig. 259). In the south, it extends along the African coasts right up to South Africa and into the Mediterranean Sea. Along the west coast of the Atlantic Ocean, this dolphin probably inhabits the waters from New- foundland to Florida and the Gulf of Mexico and beyond, along the 635 Spe At OY, ИИ, и, й А? = = > Gh, Licey =F Ё.-—- „= \\ Fig. 256. Embryo of a partly albino bottlenose dolphin, Tursiops truncatus (figure 474 by N.N. Kondakov). 2 UE РИА 2 i i. Ss Миле”, 2 Z Se 474 Fig. 257. Skull of the bottlenose dolphin, Tursiops truncatus (figure by М.М. Kondakov). coasts of Argentina, Uruguay, and Brazil. The bottlenose dolphin resides in the waters of the Indian Ocean (Arabian and Red seas, Bay of Ben- 477 gal, and Seychelles Islands), south up to the coasts of Tasmania and Australia. In the Pacific Ocean, it is encountered on the coasts of the 636 Е a 8 a 5 3 с © 8 < Fig. 258. Range of the bottlenose dolphin Tursiops truncatus т the USSR (М.А. Arsen’ev). 475 4! | Wing i] Ш " | nt wih | 1 cS, il Say" ui ia ae aaa! thy pia IY ° О ПАН й iy une 1 ba pie Quen — ОНИ 0 Nh J 5 ae я НН ани 600 0 1000 2000 3000 3000 5000 km Fig. 259. Species range of the bottlenose dolphin, Tursiops truncatus (V.A. Arsen’ev). 476 638 Korean Peninsula and Japan, on the coasts of North America (Oregon and California states), and Mexico. In the south, it reaches Australia | and New Zealand. No information is available for the Pacific Ocean coast of South America (Sergeant and Fisher, 1957; Tomilin, 1962). (V.A.) Geographic Variation Geographic variation has not been adequately studied. Apparently three subspecies are encountered in the waters of the Soviet Union.’ These are: 1. Atlantic bottlenose dolphin, T. t. truncatus Montagu, 1821. This is the largest form, with body length reaching 390 cm and skull length 550-579 mm. The depression on the lower surface of the frontal bone is inconspicuous. The number of teeth is more than in the other subspecies, usually 21-26 pairs in the upper jaw and 20-25 pairs in the lower. This subspecies is found in the Baltic and Barents seas and in the waters of the North Atlantic Ocean. 2. Black Sea bottlenose dolphin, 7: Е. ponticus Barabash, 1940. In body and skull dimensions, it is the smallest. Its body length does not exceed 310 cm while the skull length may reach 503 mm. The teeth vary in number from 19-22 pairs in the upper jaw and 18-21 pairs in the lower. The rostrum is shortened and broadened at the base. Broadening and dilation in the midportion of the premaxillae are faintly manifest or altogether absent. This subspecies is found in the Black Sea. Outside the USSR, it has been reported in waters south and west of the Black Sea. 3. Far-eastern bottlenose dolphin, T. г. gillii Dall, 1873. In dimensions, it probably occupies an intermediate position with the skull length reaching 540 mm. The articulation for the mandibular condyle is very large. The lower surface of the frontal bone is deeply concave anterior to the optic canal. The rostrum is relatively long and narrow at the base. This subspecies is found in the seas of Japan and Okhotsk and waters of the southern part of the Kuril range. Outside the USSR, it has been reported in the Pacific Ocean, at least in the northern half. (V.A.) ’Diagnosis of the subspecies after А.С. Tomilin (1957) requires verification and supplementation. 478 639 Biology Population. This dolphin is quite rare in the North Atlantic and Pacific oceans. It is comparatively less in number in the Black Sea, especially compared to the large schools of common dolphins. Food. Information on food in the Atlantic and Pacific oceans is frag- mentary. Fishes serve as the main food, including eels, gray mullets, and small sharks. Bottlenose dolphins examined from the Mediterranean Sea contained the remnants of cuttlefish. More complete information is avail- able about the food objects of the Black Sea bottlenose dolphin. The diet includes many species of fish, crustaceans, and mollusks: haddock, floun- der, skate, umbrinas, scorpionfish, anchovy, mullets, gray mullet, Black Sea shad, and bonito; shrimps and isopods; Nassa reticulata, Cardium sp., Modiola phaseolina, Syndesmia sp., Cardium simile, and Mytilus sp. Mollusks are probably not consumed by the bottlenose dolphin but enter the stomach together with sand and gravel, invariably present in large quantity. It is also quite possible that the crustaceans found in the stom- ach of these dolphins had been priorly ingested by the fish the animals consumed. Haddock, followed by flounder, is the most abundant fish found in the stomach of bottlenose dolphins. Anchovies, during the period of their en masse concentration, also play a significant role in the dolphin’s diet, sometimes even occupying a favored position. All other species of fish play only a secondary role. Thus the bottlenose dolphin is, for the most part, benthophagous since pelagic fish, which even form massive concentrations (anchovy), are consumed more rarely (Kleinenberg, 1936, 1938, 1956a). Data on the quantum of food intake of the bottlenose dolphin (under natural conditions) are not available. A bottlenose dolphin held captive in a Florida aquarium consumed 20 kg of fresh fish every day while adults in a New York aquarium could manage with 32 kg of fish each (Tomilin, 1957). Daily activity and behavior. It was earlier assumed that bottlenose dolphins feed only at night but the stomach of those caught during the day contained partly digested food. This led to the conclusion that there are no obvious changes in their diurnal activity (Kleinenberg, 1956a). Bottlenose dolphins do not form large schools (Fig. 260). They are usually found in groups of ten or a few tens, and only rarely in hundreds. Sometimes the animals produce blows rising to a height of 1-1.5 m. The respiratory pause in an animal moving undisturbed aver- ages 15-17 sec. Feeding dolphins rise to the water surface at irregular intervals, ranging from 5 sec to 2 min, while inhalation/exhalation extend 640 478 Fig. 260. Bottlenose dolphin at sea (photograph by О.А. Morozov). for 1-2 sec. Sometimes the bottlenose dolphin remains under water for 4 or even 6-7 min. It can dive to a depth of 50-90 m (possibly even to 150 m) at speeds varying from 3-5 to 11-13 km/hr, with a maximum of 30-40 km/hr. Sometimes the bottlenose dolphin breaches the water, “rising” to heights of over 3-4 m (Tomilin, 1957, 1962). Cases of active cooperation have been observed among these dol- phins. On October 30, 1954, an adult dolphin in a herd of some 25 animals was stunned by a dynamite explosion off the coast of Florida. Immediately, two large dolphins came to its rescue, lifted it from under the flippers, and supported it on the water surface. The rescuers peri- odically dived and surfaced and throughout this period the entire group moved in a wide arc. The operation gave the impression that the pair of supporting animals interchanged since the rest of the animals remained in their proximity. After a few minutes, the affected dolphin began to recover and the entire school then quickly moved away from the ship. Every 10-20 sec, the dolphins breached the water, flying in the air for 7-9 м. 479 In another incident, recorded оп November 23, 1954, also in Floridan waters, some dolphins that had been caught were being transferred from a ship into a specially constructed tank. Three adult animals had already acclimatized well while the fourth on being lifted from the deck by the tail struggled fiercely and hit its head against the wall of the tank. The 641 dolphin was stunned and sank to a depth of 2 т. Two of the already transferred dolphins quickly rushed, as in the previous case, to hold and support it on the surface of the water until the affected animal’s respiration normalized. Thereafter the two rescuers returned frequently to the bruised animal, abandoning it only after it had begun to swim freely without their assistance. The third dolphin had meanwhile remained in the proximity of the others (Siebenaler and Caldwell, 1956). The bottlenose dolphin can survive in captivity better than other species, live long in large aquariums, is easily trained, and often raises offspring in captivity. Sometimes a conditioned reflex can be developed in a dolphin within a week’s time by repeating the stimulus no more than two or three times, after which a new stimulus can be introduced. The bottlenose dolphin learns quickly to jump through a hoop, tug a boat or a raft, play with a ball, etc. The dolphins are usually held in tanks in groups, eat well, and sleep well. They often sleep at night and in the morning, occasionally enjoying a siesta after a meal. They produce different types of sounds in the frequency range of 7,000 to 170,000 Hz: whistling, howling, barking, clapping of jaws, etc. Each sound has a particular meaning, relevant to feeding, excitation, mating, fright, etc. (Tomilin, 1962). Seasonal migrations and transgressions. On the European and Amer- ican coasts of the North Atlantic Ocean, bottlenose dolphins proba- bly perform regular migrations but no direct observations have been reported. In the Black Sea, they inhabit only the coastal shallow water zone and are never seen in the open sea. They are most often seen in the northeastern part of the Black Sea along the coasts of southern Crimea and northern Caucasus but have been sighted in small numbers in other parts of the sea as well. During the migration of Azov anchovy, bot- tlenose dolphins form schools that are larger than at any other time and concentrate in the fore-channel zone of the Black Sea and are seen even in Kerch Strait. However, they do not enter the Azov Sea. Apparently, the Black Sea bottlenose dolphin does not undertake regular migrations (Kleinenberg, 1956a). Reproduction, growth, and development. The period of reproduction is highly protracted although most calves are born in the warm season of the year. However, in a Florida aquarium, four calves were born in February, one in March, one in April, and four in May (Tomilin, 1962). Thirty-four females caught in November were accompanied by suck- iing calves and the mammary glands of the mothers held milk. Gestation extends for 12 months. In 1953, in a marine aquarium, a female in the company of a male from February 23 through March 9 underwent partu- rition on March 4, 1954. Some other observations confirm the 12-month 642 480 Fig. 261. Parturition in the bottlenose dolphin, Tursiops truncatus (figure by N.N. Kondakov). cycle of gestation (Tavolga and Essapian, 1957). Large embryos (Fig. 261) measured 90 to 130 cm in length while two newborn calves were 122 and 124 cm long (Barabash-Nikiforov, 1940). According to observations on captive dolphins, calves were born under water tail first, with formed dorsal fin and caudal flukes. Immediately after whelping, the mother swirled about in the tank, thereby readily separating the umbilical cord. The newborn swam slowly along the waterline, inclined to the surface, and took its first breath 10 min after birth. After 1 hr and 45 min, it suck- led milk for the first time. According to other observations, subsequent feedings proceed day and night at intervals of 15-30 min. The females in an advanced stage of pregnancy segregate from the herd and form a separate group of pregnant animals. After parturition, the mother as well as the other females living in the proximity protect the calf by turns from. possible attack by males. For a few weeks, the mother does not allow her calf to swim for more than two-three meters 480 out of her sight. The duration of lactation apparently varies markedly. In one case, the calf was milk-fed for 3.5 months while another calf was found to partake of a fish by itself just seven months after birth. Yet a case is known of a young dolphin continuing to suckle its mother for 18 months (Tavolga and Essapian, 1961*; Tomilin, 1962). . 481 643 Growth rate has not been established. Apparently the newborn grows rapidly on the high fat content of the mother’s milk, which in composi- tion contains: fat 46.1%, protein 11.55%, sugar 1.57%, dry residue 13.5%, ash 0.38%, and water 40.5% (Ural’skaya, 1957). A female born in a marine aquarium was fertilized for the first time at six years of age and the first calf born a year later. Considering the duration of gestation and lactation, it may be assumed that the bot- tlenose dolphin has a two-year reproductive cycle. A female born in a Florida aquarium gave birth to four calves over a 15-year period. A male when caught was approximately 1.5 - 2 years of age with a body length of 183 cm; he lived in the marine aquarium for six years and ten months and grew in this period to 231 cm, 1.е., added 48 cm in almost seven years (Sergeant, 1959b). Enemies, diseases, parasites, mortality, and competitors. The killer whale can be an enemy of the bottlenose dolphin, as also of other dolphins. The diseases affecting the bottlenose have not been investigated. In a Florida aquarium, two dolphins died of infectious erysipeloid (pathogen: Erysipelothrix rhusiopathiae). In a New York aquarium, instances of pneumonia were recorded. Skeletal diseases were also reported (Tomilin, 1957). A female bottlenose dolphin measuring 266 cm long and weighing 180 kg was caught in the waters of Japan in July, 1954 and placed in a tank. In December of the same year, many nodular formations were noticed on her skin, the animal became debilitated, and a month later died. The affliction turned out to be a bacterial fungal infection caused by a parasite of the genus Trichophytum Malm. Acute pneumonia was also reported in this dolphin (Hoshina and Sugiura, 1956). Eleven species of helminths have been reported in the bottlenose dolphin: three trematodes, three cestodes, four nematodes, and one species of acanthocephalan. The trematode Braunina cordiformis Wolf, a parasite of the stomach and intestine of the bottlenose dolphin, was also found in the common dolphin and one species of spotted dolphins in the Adriatic Sea, in the region of Rio de Janeiro, and in the waters of California and Panama. Another species of trematode, Synthesium tursionis Marchi, was detected in the intestine of only the bottlenose dolphine in European waters. In a Californian aquarium, the bile ducts of the liver of a bottlenose dolphin revealed Zalophotrema hepaticum Stunkard and Alvey, known earlier from the Californian sea lion. The cestode Phyllobothrium delphini Bosc, parasitizing the skin and subcutaneous tissue, is widely distributed among marine mammals. Besides the bottlenose dolphin, it is found in the bowhead whale, 644 six species of toothed whales, and one species of seals. It has been detected in the Atlantic and Pacific oceans, Mediterranean Sea, and in the waters of Australia and Antarctica. In the bottlenose dolphin and four other species of dolphins, the cestode Monorygma grimaldii Monier parasitizes the abdominal cavity, mesentery, and diaphragm; it has been detected in the Atlantic Ocean and the Mediterranean Sea. The third species of cestode belongs to the genus Diphyllobothrium (species not established). Of the four species of nematodes, Anisakis tursionis Gruz was detected in the bottlenose dolphin from the Indian Ocean. Crassicauda crassicauda Creplin, parasite of the urogenital system, was found in beaked whales and six species of baleen whales, in addition to the bottlenose dolphin; it was detected in the Atlantic Ocean in the Northern and Southern hemispheres. The nematode Halocercus lagenorhynchi Baylis and Daubney is a parasite of the lungs (bronchi) of the bottlenose dolphin and the white-beaked dolphin; it was detected on the European coast of the Atlantic Ocean and in the waters of Australia. Stenurus ovatus Linstow was detected in the blowhole, bronchi, and blood vessels of only the bottlenose dolphin in the Mediterranean and Black seas. Forty-one larvae of the nematode genus Anisakis were found in a female bottlenose dolphin caught in the waters of Japan, probably the larvae of Anisakis catodonis Baylis (Hoshina and Sugiura, 1956). The only species of acanthocephalans localizes in the intestine of the bottlenose dolphin Corynosoma cetaceum Johnston, and has also been found in the common dolphin in the waters of Australia (Delamure, 1955). 1 The natural mortality of the bottlenose has not been studied. Some other species of dolphin feed on the same diet and constitute food competitors. Field characteristics. Adults usually measure 220 to 250 cm. They are dorsally and ventrally light in color; the transition in coloration is gentle, with no sharp demarcation between the two. Body movements are placid. It neither splashes water while swimming nor does it breach high. (V.A.) Economic Importance In the recent past, several hundreds of bottlenose dolphins were caught every year in the Black Sea. In other regions, where many other species of dolphins were caught, the bottlenose constituted a negligible per- centage of the total catch. Various types of nets were used to catch it 482 along with other dolphins. The weight of the animals ranged from 90 to 645 almost 200 kg, of which the skin with the fat accounted for 30% and the flesh 33%. An analysis of the seasonal catch of the Black Sea bottlenose dolphin showed that the average weight of the dolphin was 175 kg, of which the subcutaneous fat was 50.8 kg (29%) and the skin 7 kg (4%) (Dragunov and Kasinova, 1951). The total quantum of products obtained from bottlenose dolphins caught all over the range was extremely meagre. Hunting of the bot- tlenose dolphin, and of other species of dolphins as well, has been banned in the Black Sea. (V.A.) Genus of Right Whale Dolphins Genus Lissodelphis Gloger, 1841 1830. Tursio. Wagler. Nat. Syst. Amphibien, р. 34. Delphinus ретопи Cuvier. Nom. ргаеосс. 1841. Lissodelphis. Gloger. Gemein. Naturgesch, 1, р. 169. Delphinus per- onii Lacépéde, 1804. 1846. Delphinapterus. Gray. Zoology. Voyage Erebus and Terror, 1, p. 36. Delphinus peronii Lacépede. Nom. ргаеосс. (V.H.) Small dolphins, with a body length of up to 2.4 m. Table 39. Weight of body parts and organs of bottlenose dolphin from the Black Sea (in March), kg (percentage of total weight shown in parentheses) (Okuneva, 1934) Total weight 143.53 (100.0) Head 12.0 (8.36) Flippers and dorsal fins 2.06 (1.43) Caudal flukes 2.09 (1.45) Skin 6.0 (4.11) Carcass fat 46.0 (32.11) Flesh (muscles) 39.9 (27.17) Bones (trunk skeleton) 17.0 (11.83) Brain 1.46 (1.01) Tongue 0.37 (0.25) Lower jaw 0.79 (0.55) Upper jaw 1.09 (0.76) Viscera, including: 173 (12.05) Liver 2.1 (1.88) Heart 0.67 (0.46) Lungs 4.5 (3.13) Stomach 1.55 (1.08) Intestine 7.2 (5.01) Kidneys 0.68 (0.47) 646 The body is highly elongated and well proportioned. The head, with a low and inclined forehead gradually passes into the relatively short “beak”. The “beak” is set off from the adipose body of the forehead by lateral furrows. The flippers are crescent-shaped. A dorsal fin is lacking. Caudal flukes are relatively small. The lower jaw is slightly longer than the upper. The body is black above and whitish below and the boundary between the upper and lower body colors is very sharply defined. The rostrum is broad and slightly longer than the cranium. The ptery- goid bones are not adjacent. The premaxillae are flattened anteriorly. The mandibular symphysis is short, less than one-fifth the jaw length. The teeth are small and number 73-4. The scapula, almost semicir- cular in form, has prominent coracoid and acromion processes. Cervi- cal vertebrae 7, thoracic 14-17, lumbar 29-33, and caudal 35 - 40; total 88 - 92. The biology of this genus is hardly known but apparently these dol- phin species feed mainly on fish and cephalopods. Information on the distribution of this genus is very inadequate and sketchy: Atlantic Ocean around South Africa and south of southern Brazil in South America, Pacific Ocean along the coasts of Chile, north up to 37° М lat., and from Tasmania and New Zealand (evidently Aus- tralia too) north up to New Guinea, and in the North Pacific Ocean from the Bering Sea south to California and Japan. The genus comprises two species: the southern right whale dolphin (Peron’s dolphin), Lissodelphis peroni Lacépéde, 1804, and the northern right whale dolphin L. borealis Peale, 1848. Only the latter has been established in USSR waters but the appearance of the southern species is quite likely. (V. S.) NORTHERN RIGHT WHALE DOLPHIN Lissodelphis borealis Peale, 1848 1848. Delphinapterus borealis. Peale. U. S. Explor. Exped. 8, p. 35, 10° west of Astoria, Oregon, USA; 46°6’50” М lat. and 134°5' Е long. (V. H.) Diagnosis The body length, up to 2.4 m, is maximum for the genus. For a descrip- tion of the external features, see under the characteristics of the genus. Typically, the body is almost entirely black. A narrow light-colored band runs only on the ventral side from the throat to the tail and forms a 647 483 rhomboid patch between the flippers. The upper side of the beak, part of the forehead, and tip of the lower jaw may also be white (Fig. 262). For skull characteristics, see under the description of the genus. Phalangeal formula: I, -5, Пз, Шо, [Уз, and V,-3. (V.S.) Description The upper side of the caudal flukes is black but partly white or gray on the underside and dark along the margins. The flippers are black. Teeth 2-77. Cervical vertebrae 7, thoracic 14-17, lumbar 29-33, and caudal 35-40; total 88-92. The first two cervical vertebrae are fused. The body dimensions of four males and seven females caught on the coasts of Japan (Tobayama, Uchida, and Nishiwaki, 1969) are (in cm): body length 145 - 292 (x 204); distance from tip of snout to forehead 3-6 (x 4.6), to center of blowhole 22 - 34 (x 30), and to flippers 39 -64 (x 53); distance from anal opening to notch between caudal flukes 40-70 (х 53); length of flippers 24-31 (х 28), maximum width of flippers 7.3-9.7 (х 8.6); distance between caudal flukes from apex to apex 18-41 (х 33); and maximum height of body 20-37 (x 30). The skull (Fig. 263) dimensions of three male northern right whale dolphins (Tomilin, 1957) of 213, 208, and 246 cm, body length are respec- tively: condylobasal length 45, 42, and 44 cm; length of rostrum 26, 23, and 24 cm; width of rostrum at base 11, 11, and 11 cm; and length of lower jaw 38, 37, and 38 cm. (V.S.) Geographic Distribution North Pacific Ocean. 483 Fig. 262. Northern right whale dolphin, Lissodelphis borealis (figure by М.М. Kondakov). 648 arn ras SSS Sanaa { SESS 483 Fig. 263. Skull of the northern nght whale dolphin, Lissodelphis borealis (figure by N.N. Kondakov). Geographic Range in the USSR Waters of the Kuril Islands (from Shpanberg to Paramushir) but 484 encountered more often from the Pacific Ocean side. Apparently also occurs in the eastern parts of the seas of Okhotsk and Japan (Fig. 264). Geographic Range outside the USSR (Fig. 265) Widely distributed in the Pacific Ocean along the American and Asian coasts, in the waters of Honshu and Hokkaido islands, on the coasts of North America from California and Washington states to the Gulf of Alaska and the eastern part of the Bering Sea; residence in the western part of the sea has not been established. (V.A.) Geographic Variation Not established. Quite likely the accepted species of this genus are only subspecies of a single species (L. peroni Lac.). (V.A.) Biology The population of this species is very small over much of its range. Махипит concentration in all probability occurs in the waters of Japan. 484 486 649 Fig. 264. Range of the northern right whale dolphin, Lissodelphis borealis in the USSR (V.A. Arsen’ev). 250 0 250 500 750 1000km = —————— | 150 160 170 Fish represent the main food while cephalopods are consumed in small quantities. A female caught on March 28, 1959 east of Honshu Island contained an embryo 43 cm long. These dolphins live predominantly in small schools far away from the coasts. They leap clear of the water. They exhibit a strong mutual affinity, never abandoning an injured animal and remaining with it for a long time. In the waters of Japan, the northern right whale dolphin is caught regularly along with other species. At least a few hundred of them are hunted every year. Thus, in May and June of 1949 alone, in the central part of the coastal Pacific Ocean waters of Honshu Island, a whaling com- pany caught 465 of them. Hunting is usually donw with rifles, followed by harpooning the injured animal manually. The shooter and the harpoon striker operate on a special wooden platform fitted on the bow of the 650 (лэмэзту “W'A) 51122404 siydjapossr] ‘uydjop этеча з48ы way OU эц1 Jo э8иет $э0э4$ ‘695 “31 9 486 651 Fig. 266. Northern right whale dolphin, Lissodelphis borealis, on the deck of a ship. Pacific Ocean east of Honshu Island, 1959 (photograph by G.M. Kosygin). ship. Ships with a water displacement of 20 to 30 tons and capable of log- ging 7 to 10 miles per hour are used. The killed dolphins are processed on the deck of the ship (Fig. 266). A ship enjoying a successful hunting expe- dition returns with a catch of 200 or more dolphins of different species. An experimental catch of these dolphins using woven nets was саг- ried out near Iturup in the Kuril Islands in September, 1955. The nets were cast over a herd of 25 animals but only 5 were actually caught. The products of dolphin hunting are marketed in Japan where the liver, heart, and kidneys are consumed. Bones and various remnants are ground into fertilizer meal. Oil is obtained from the blubber. The skin is used in making leather goods (Wilke, Taniwaki, and Kuroda, 1953; Sleptsov, 1955, 1961; Klumov, 1959). (V.A.) SOUTHERN RIGHT WHALE DOLPHIN (PERON’S DOLPHIN) Lissodelphis peroni Lacépéde, 1804 1804. Delphinus peronii Lacépéde. Hist. Nat. Cétacées, p.p. XLIII, 316. Tasmanian waters. (V.H.) These dolphins are smaller than the northern right whale dolphin. The body length of a male caught in waters of Japan was 227 cm (Tobayama et al., 1969). In external features and structure and skull proportions, they in no way differ from their northern counterparts. The typical coloration of the southern species is: black dorsally and on most of the lateral surface, and white on the abdomen and the lower parts of the body flanks (Fig. 267). 487 487 652 The tip of the snout and small sections of the vertical surface of the caudal flukes adjoining the caudal crest are black. The white field covers the level of articulation of the black flippers. The edge of the upper jaw (apart from its apex) and the zone posterior to and below the eyes are white. Teeth 4%. Cervical vertebrae 7, thoracic 17, lumbar 29, and caudal 37; total 90. The body dimensions of a male caught on the coasts of Japan (Tobayama et al., 1969) are (in cm): body length 227; distance from tip of snout to forehead 4, to center of blowhole 36, and to flippers 59; distance from anal opening to slit between caudal flukes 61; length of flipper 30, maximum width of flipper 9.5; distance between caudal flukes from tip to tip 38; maximum height of body 27. The skull (Fig. 268) dimensions of the same specimen are (in cm): condylobasal length 47, length of rostrum 26, width of rostrum at base 12, interorbital width 18, length of lower jaw 40, and length of mandibular symphysis—[48]. Geographic Distribution Habitation in the waters of the USSR has not been established but res- idence in or transgressions into the Sea of Japan or waters of the Kuril Islands are possible. Outside the USSR, it is encountered in the southern part of the range (see under the characteristics of the genus); indicated for the waters of Japan. (V.H.) Fig. 267. Southern right whale dolphin, Lissodelphis peroni (figure by N.N. Kondakov). 653 Biology Not known. Japanese fishermen from Otsu, Ibaraki Prefecture caught a group of dolphins swimming together on April 1, 1969, which included the southern right whale dolphin, the northern right whale dolphin, and the Pacific common dolphin (Tobayama, Uchida, and Nishiwaki, 1969). (V.A.) Genus of Shorthead Dolphins Genus Lagenorhynchus Gray, 1846 1846. Lagenorhynchus. Gray. Ann. Mag. Nat. Hist., 17, p. 84. Delphinus albirostris Gray, 1846. 1866. Electra. Gray. Cat. Seals and Whales Brit. Mus., p. 268. Lagenorhynchus electra Gray, 1846. Nom. ргаеосс. 1866. Leucopleurus. Gray. Proc. Zool. Soc. London, p. 216. Delphinus leucopleurus Rash, 1843. Sr a ey, SLE! Са ^^ В tA 77° Fig. 268. Skuil of the southern right whale dolphin, Lissodelphis peroni (figure , by N.N. Kondakov). 489 654 1966. Peponocephala. Nishiwaki and Norris. Ocean Res. Inst., Univ. Tokyo, 5, 139. Electra electra Gray, 1846. (V.H.) Small dolphins, with a well-proportioned body ranging in length from 1.5 to 3 м. The “ЪеаК” is short, not more than 7 cm; in most of these animals, it is well demarcated from the forehead by a furrow. The high dorsal fin is crescent-shaped and located at the center of the dorsum or slightly anterior to it. A crescent shape is also characteristic of the flippers. Dermal keels run along the dorsal and ventral sides of the caudal stem. The coloration of most of 'thé species is a combination of black, gray, and white fields extending along the body. The rostrum of the skull is wide at the base and is as long as the cranium or slightly longer. The anterior section of the ргетахШае is flattened or slightly concave. The large pterygoid bones adjoin each other or are slightly separated. The length of the mandibular symphysis is less than one-fifth that of the lower jaw. The teeth are small and number 2 43. The number of vertebrae varies from 71 to 94 and their centrum is highly flattened. The vertebrae of the lumbar section have very long, thin spinous and transverse processes. The first two or three cervical vertebrae are fused. Very little is known about the biology of this genus. These dolphins feed mainly on fish and cephalopods. Mating and parturition occur in the summer months. Shorthead dolphins are distributed in all the oceans in the north up to Greenland and the Barents Sea; in the south they reach the edge of the Antarctic ice (Fig. 269). The genus apparently comprises six species, although some authors recognize up to ten (Nishiwaki and Norris, 1967): 1) L. acutus Gray, 1828; 2) L. albirostris Gray, 1846; 3) L. cruciger Quoy and Graimard, 1824; 4) Г. electra Gray, 1846; 5) Г. obliquidens Gill, 1865; and 6) Г. thicolea Gray, 1846. L. electra is sometimes considered as an indepen- dent genus, Peponocephala Nishiwaki and Norris, 1966; this aspect calls for further study but it would appear to deserve only a subgeneric rank. The following species inhabit the waters of the USSR: (1) Atlantic white-sided dolphin, L. acutus Gray, 1828; (2) white-beaked dolphin, L. albirostris Gray, 1846; and (3) Pacific white-sided dolphin, L. obliquidens Gill, 1865. Residence of the broadsnout dolphin, L. electra Gray, 1846, is possible. These animals are not hunted in our waters. (V.S.) "(лэ.иэзту А) snyoudysouasvT ‘зищаюр реэчзлоцз Jo $пиэ8 ay] Jo э8ие “697 “314 Ss wy 000$ 000% 0006 000Z 0001 0 0001 490 656 ATLANTIC WHITE-SIDED DOLPHIN Lagenorhynchus (Lagenorhynchus) acutus Gray, 1828 1828. Delphinus (Grampus) acutus. Gray. Spicil. Zoologica, I, p. 2. Faroe Islands. 1843. Delphinus leucopleurus. Rash. Nytt. Mag. Natur., vol. 4, p. 100. Kristiana Bay (Oslo), Norway. (V.H.) Diagnosis Body length up to 275 cm. The body is mainly black dorsally and on the flanks and whitish ventrally. A broad white band extends along the flanks, roughly from the level of the dorsal fin and almost up to the caudal stem. A narrow black band runs from the base of the flippers anteriorly and upward toward the section between the eye and the corner of the mouth. The dark-colored flippers are surrounded by a white field. The pterygoid bones are often adjacent in the skull. Teeth Vertebrae 77 - 82. 30-37 30/5 3iir Description The head is relatively small. The “beak” is not well demarcated from the inclined forehead. The caudal flukes are ventrally light-colored. The margin of the lower jaw is dark-colored. A black ring is seen around the eye and anus (Fig. 270). The embryo bears seven or eight hairs, each measuring roughly 1 cm, on each side of the beak. The rostrum is slightly longer than the cranial section. The pre- maxillae are flat and their outer edge sinuate. The temporal fossa is elongated. Cervical vertebrae 7, thoracic 14-15, lumbar 18-22, and caudal 38-41. Phalangeal formula: I, -5, Ij, Шб, [У› -3, and V>. 490 Fig. 270. Atlantic white-sided dolphin, Lagenorhynchus acutus (figure by М.М. Kondakov). 490 ‘657 The body length of an adult female Atlantic white-sided dolphin was 225 cm (Schevill, 1956); distance from tip of snout to commencement of flippers 40 cm, from tip of snout to dorsal fin 87 cm, and from tip of snout to blowhole 29 cm; height of dorsal fin 24 cm, length of dorsal fin along base 34 cm; length of flippers 30 cm; and spread of caudal flukes 65 cm. The skull dimensions (Fig. 271) (Tomilin, 1957) (average of 10 mea- surements) are (in cm): condylobasal length 41, length of rostrum 21, width of rostrum at base 11, length of lower jaw (three measurements) 33, and length of mandibular symphysis (two measurements) 43. The rostrum of the skull is shorter than in the Pacific white-striped dolphin but longer than in the white-beaked dolphin. The teeth are usually more numerous than in the other two species but their dimensions are smaller. (V.S.) Geographic Distribution North Atlantic Ocean. Wi SSSe fae Sl == oa < За. = ме: Ste = мл, Fig. 271. Skull of the Atlantic white-sided dolphin, Lagenorhynchus acutus (figure by N.N. Kondakov). 491 658 Geographic Range т the USSR Barents Sea up to Spitsbergen and Murman coast of the Barents Sea up to Kanin (Fig. 272). Transgressions are possible into our waters of the Baltic Sea. Geographic Range outside the USSR (Fig. 273) In American waters from Davis Strait to the Gulf of Maine (Cape Cod). In the eastern part of the ocean from the Barents Sea (Kanin and Spits- bergen) and southern Greenland to the southern part of the North Sea (England, Belgium, Denmark, and Ireland). In the Baltic Sea, only in the southwest (waters of Denmark, Oslo Fjord, and western coasts to Sweden), in the east not farther than Penemund close to Oder estuary (about 18° W long.) and Koloberg (Poland; Kowalski, 1964). This species is apparently most numerous on the coasts of Norway, especially in the region of Lofoten Islands and Bergen. Encountered Fig. 272. Range of the Atlantic white-sided dolphin, Lagenorhynchus acutus, in the USSR (V.A. Arsen’ev). 492 659 Fig. 273. Species range of the Atlantic white-sided dolphin, Lagenorhynchus acutus (V.A. Arsen’ev). rather often on Faroe and Orkney Islands and on the coasts of Ireland and Iceland (Jonsgard and Nordli, 1952; Tomilin, 1957, 1962). (V.A.) Geographic Variation Not established. Biology Only fragmentary information is available. Pelagic animals are encountered more often than benthic animals in the stomach of these dolphins: salmon, herring, mackerel, and the squid, Шех illecebrosus. On one occasion, bottom-dwelling hermit crabs (Pagurus bernhardtii) and mollusks (Buccinum) were found. It has been suggested that gestation extends for about 10 months and that parturition takes place midsummer since fully mature embryos were found in June. The newborn is 100 cm or slightly more in length, reaching 140 cm by November. A male 180 cm long proved to be three years old while a female 225 cm long was neither pregnant nor lactating. In Kalvag Fjord on the west coast of Norway, a school of these dolphins arrived on March 10, 1952 and 52 were caught. Visually, many were about 2.5 m long and the longest about 3 m. About one-third of the dolphins caught contained embryos 55 to 70 cm long. One female contained two embryos, each about 65 cm long. 493 660 These dolphins are usually confined to groups of 10 to 50 individuals but at places where fishes concentrate they gather into large schools of 1,000 to 1,500 animals. Migrations have not been traced. Beached dolphins have been found on the northeastern coast of England, northwestern coast of Ireland, and the coasts of Holland. Once a school of 30-35 animals was found beached on the shoals of Scotland. Beached animals sometimes include both dolphins and pilot whales. The helminth fauna of the Atlantic white-sided dolphin is rather poor. The nematode Anisakis (Anisakis) simplex Rudolphi, wide-ranging in marine mammals, was detected in the gullet, stomach, and intes- tine. Two species of cestodes were also found: Strobilocephalus triangu- laris Diesing, also parasitizing the intestine of the bottlenose whale and Mesoplodon localizes in the abdominal cavity, the mesentery, and the diaphragm. Monorygma grimaldi Monier has also been found in three other species of dolphins (Jonsgard and Nordli, 1952; Delamure, 1955; Tomilin, 1957, 1962). The most characteristic external feature is the broad white band running along the body from both sides below the dorsal fin up to the caudal stem. This band is distinctly visible since it lies between the dark coloration on top and the yellowish-gray color below, the latter cov- ering the lower flanks of the body. The belly of this dolphin is pure white. This species is of minimal economic importance on the coasts of Norway where up to 1,000 animals are caught in some years. In other regions of the range, it is caught incidentally along with other species. (V.A.) WHITE-BEAKED DOLPHIN Lagenorhynchus (Lagenorhynchus) albirostris Gray, 1846 1846. Delphinus albirostris. Gray. Ann. Mag. Nat. Hist., 17; p. 84. Near Great Yarmouth, England. (V.H.) Diagnosis Body length up to 304 cm. The color of the upper portion of the body is grayish-black and the underside whitish. The base of the forehead and the “beak” are light gray. The premaxillae are broad and flat and their outer margin is curved. The temporal fossa is ellipsoidal. the pterygoid bones are often adjacent. Teeth 227 (of these only 3=% are seen above the gums). Vertebrae 88-92. (V.S.) 661 Description This species differs from the Atlantic white-sided dolphin in having less developed keels on the caudal stem, very large flippers, and a very high dorsal fin. The flippers and caudal flukes are dorsally dark and somewhat lighter ventrally. The dark color extends down along the flanks of the body and covers the point of articulation of the flippers (Fig. 274). The rostrum is almost as long as the cranial section. Usually, the cervical vertebrae number 7, thoracic 14-16, lumbar 24-27, and caudal 43 - 45. Phalangeal formula: [,_ 3, Пб-7, Ш4-5, [1V,->, and Уу-1. The body length usually varies from 270 to 300 cm. The average main measurements of the skull (Fig. 275) (Tomilin, 1957) are (in cm): condylobasal length 44 (eight measurements), length of rostrum 21 (eight), width of rostrum at base 14 (seven), and length of lower jaw 36 (four). (V.S.) Geographic Distribution North Atlantic Ocean. Geographic Range in the USSR Barents Sea, Murman coast, and waters of Rybachii Peninsula; Baltic Sea, including the gulfs of Finland and Riga (Fig. 276). 493 Fig. 274. White-beaked dolphin, Lagenorhynchus albirostris (figure by N.N. Kondakov). 662 493 Fig. 275. Skull of the white-beaked dolphin, Lagenorhynchus albirostris (figure by N.N. Kondakov). 194 Fig. 276. Range of the white-beaked dolphin, Lagenorhynchus albirostris, in the USSR (V.A. Arsen’ev). 495 663 Geographic Range outside the USSR Along the American coast, it is encountered in Davis Strait, waters of western and southern Greenland and Labrador in the north to Mas- sachusetts Bay in the south. In the eastern part of the Atlantic, from the Barents Sea, Iceland, and Greenland to the coasts of France (Vannes), Great Britain, and Ireland. Waters of Denmark and southern Sweden in the Baltic Sea (Fig. 277). Often seen in the western waters of the Baltic Sea. (V.A.) Geographic Variation Not known. Biology Large herds of thousands of animals are seen in the summer months in the northern parts of the range (Barents Sea, waters of Iceland, southern Greenland, and coasts of Labrador and Newfoundland). This species feeds mainly on benthic and bottom-dwelling fish and other animals. The stomach of examined dolphins contained cod, capelin, 494 Fig. 277. Species range of the white-beaked dolphin, Lagenorhynchus albirostris (V.A. Arsen’ev). 664 navaga, whiting, herring, and less frequently hermit crabs, cephalopods, and some mollusks. In summer they usually live in pairs or small groups but some- times form schools of significant strength. They probably perform regular migrations since, for example, they are encountered only in spring and summer but disappear in autumn in Davis Strait; they are more numerous in summer than winter in the waters of Norway and Great Britain. Instances of beached dolphins have been reported, although compar- atively not often, on the coasts of Orkney Islands, Ireland, Sweden, Denmark, Holland, the Federal Republic of Germany, and France. The period of mating is somewhat extended but is confined to the summer months. Most females give birth in midsummer. The embryos measured were 113 to 122 cm long but a female 305 cm long contained an embryo 165 cm long. Yet a suckling calf found on the beach was only 122 cm long. Two species of nematodes are known among the endoparasites. Anisakis (Anisakis) simplex Rudolphi, wide-ranging among marine mammals, was detected in the gullet, stomach, and intestine of the white- beaked dolphin. It has also been recorded in 10 other species of toothed whales, two species of baleen whales, and Steller’s sea lion in the North Sea and in the waters of the Pacific Ocean. Halocercus lagenorhynchi Baylis and Daubney is a parasite of the lungs (bronchi) of the white- beaked dolphin and the bottlenose dolphin; it has been detected in dolphins of the European waters of the Atlantic Ocean and in the waters of Australia (Delamure, 1955). The white-beaked dolphin is regularly hunted on the Norwegian coasts but the-volume of hunting is minimal. Previously hunting was relatively rewarding in Davis Strait but not at present, while the catch on the coasts of Great Britain is incidental. Hence this species is of very little economic importance. (V.A.) PACIFIC WHITE-SIDED DOLPHIN Lagenorhynchus (Lagenorhynchus) obliquidens Gill, 1865 1865. Lagenorhynchus obliquidens. Gill. Proc. Ac. Sc. Philadelphia, 17, p. 177. Near San Francisco, California. 1955. Lagenorhynchus ognevi. Sleptsov (Slepzov). Tr. Instituta Okeano- logii AN SSSR, 18, р. 60, 15-20 km east of Kunashir Island, Kuril range. (V.H.) 496 496 665 Diagnosis Body length up to 230 cm. The body is blackish on top and white under- neath, the flanks grayish with a dark field in the middle. The premaxillae are rounded and their outer edge slightly sinuate. The temporal fossa is large, rounded. The pterygoid bones are often separated. Teeth 3=¥. Vertebrae 74. (V.S.) Description Body coloration is subject to considerable individual variation. Usually the upper side of the body, end of the snout, anterior portion of the dor- sal fin, and the flippers are dark-colored. The flanks are lighter in color. A narrow black band runs on each side from the base of the flippers anteriorly and upward toward the corner of the mouth and posteriorly along the body up to the lower caudal stem (Fig. 278). Sometimes two symmetrical longitudinal narrow white bands run along the dorsum, com- mencing along the flanks near the blowhole. These bands later run onto the flanks anterior to the dorsal fin and continue up to the caudal stem. The integument is characterized by a thin dermal layer with abundant elastin fibers which are found in small bundles at the border of the subcutaneous fatty tissue and subcutaneous musculature. The embryos have whiskers on the snout. The length of the rostrum of the skull (Fig. 279) is roughly equal to or slightly more than the length of the cranial section. The usual dental formula is о. The diameter of the largest tooth does not exceed Fig. 278. Pacific white-sided dolphin, Lagenorhynchus obliquidens (figure by М.М. Kondakov). 496 497 666 О << _3, and V,_>. - The main body measurements of the Pacific white-sided dolphin (Wilke, Taniwaki, and Kuroda, 1953) averaged for four females and ten males are respectively (in cm): body length 172 and 167; distance from tip of snout to dorsal fin 112 and 106, to anterior margin of base of flippers 43 and 42; distance from anus to caudal bifurcation 50 and 49; maximum width of caudal flukes 50 and 44; height of dorsal fin 20 and 18; length of flippers 30 and 28 and maximum width 11 and 10. The length of the largest male was 222.6 cm and of the female 221 cm. Females weigh 67 - 93 kg, average 80 kg, and males 58 - 84 kg, average 66 kg (Japanese Hunting; Wilke, Taniwaki, and Kuroda, 1953). The average skull dimensions (Tomilin, 1957) are (in cm): condy- lobasal length 40 (ten measurements), length of rostrum 22 (eight), width of rostrum at base 10 (eight), zygomatic width 20 (six), length of lower jaw 34 (eight), and length of mandibular symphysis 4 (five). (V.S.) Geographic Distribution North ‘Pacific Ocean. 499 667 Geographic Range in the USSR Coasts of the Sea of Japan, including Peter the Great Gulf and Amur and Vityaz’ Bay. Waters of Kuril Islands—Shikotan, Iturup, Urup, Kunashir, and Paramushir on the Pacific Ocean as well as the Sea of Okhotsk side (Fig. 280). Geographic Range outside the USSR (Fig. 281) Coastal waters of Honshu and Hokkaido Islands, Pacific Ocean coast of North America from the Gulf of California (Mexico) to the Gulf of Alaska and the Aleutian Islands. (V.A.) Geographic Variation Not studied. Biology Population. One of the abundant species of Pacific dolphins. In some parts of the range, it sometimes forms schools of thousands of animals. It is numerous among our Kuril Islands on the Sea of Okhotsk as well as the Pacific side. Food. Small fish and cephalopods which form large concentrations serve as the main food. Lantern fishes—Scopelidae (Myctophidae)—are of utmost importance and represent 77% of the total intake by volume, Japanese anchovy (Engraulis japonica) 9%, Pacific mackerel (Scomber japonicus) 7%, and squids (“beaks” and eye lenses probably of Watasenia scintillans) also 7% (Wilke, Taniwaki, and Kuroda, 1953). In the coastal waters of North America, the stomach of dolphins contained herring, salmon, sardine (Sardinops caerulea), saury, northern anchovy (Engraulis mordax), scad (Decapterus paliaspis), squids including Loligo opalescens, and jellyfish. In Nemuro Strait, the stomach of a female contained squids (Ommatostrephes sloanei-pacificus) and scad [Japanese horse mackerel] (Trachurus japonicus) (Scheffer and Slipp, 1948). Lantern fishes are of major importance in the food intake of these dolphins in the waters of Japan. Squids were found in almost every stom- ach dissected but in small quantities. But their importance is undoubt- edly far greater than revealed by analysis based on the volume ratio in the intake since the stomach contents contained only the remnants of squids, such as “beaks” and eye lenses (100 to 200 “beaks” in one stom- ach). Apparently the small schooling fish and squids constitute the main food of dolphins in this region. 497 668 0 250 500 750 1000 km Fig. 280. Range of the Pacific white-sided dolphin, Lagenorhynchus obliquidens, in the USSR (V.A. Arsen’ev). Daily activity and behavior. These dolphins are usually more active in the day. They are encountered in small groups as well as in large herds. Herds comprising animals of different ages predominate in the summer months but sometimes separate groups of young dolphins are also encountered. These dolphins are more often seen in large bays and gulfs than in the open sea. They quite often follow a ship but on closing in on it, dive to a depth of 1-3 m. Sometimes playful dolphins leaping high above the water land on their side or back with a loud thud and raise 669 ‘(ao,uasty А) suapinbygo snysukysouasyT чиц@ор рэр!з-эицл эцоеа ay) Jo э8ие1 $910э4$ ‘185 “S14 ост Ost 86? 500 499 670 much spray (Fig. 282). These dolphins are encountered in groups with other species also. On one occasion an adult dolphin trying to rescue a harpooned comrade was observed. It dove frequently between the ship and the injured dolphin and each time pushed it farther away from the ship. These animals survive well in tanks, live long, and can be trained well. They quickly learn to play with a ball, jump through a hoop, etc., and to snatch food from the trainer’s hand (Tomilin, 1962). Migration. Seasonal migrations probably do occur but the informa- tion available is highly fragmentary. In the waters of California, these dolphins are noticed in the winter months (November, December, Febru- ary, March, and April); on the coasts of Washington state in March and later in July, August, September, and November; on the coast of British Columbia in July;.and in the waters of the Alaskan peninsula from July to September. Judging from the Japanese hunting experience, these dolphins are encountered in early March on the Pacific Ocean coasts of Honshu at 36° М lat. and in mid-March at 39 to 40° М lat. where they remain until June, partly advancing farther north toward Hokkaido coasts. In summer and autumn, large schools are seen in the coastal waters of the Fig. 282. Pacific white-sided dolphins leaping above the water, East China Sea, March, 1968 (photograph by A.V. Kucheryavenko). 671 Kuril Islands in the Pacific Ocean and in the Sea of .Okhotsk. A large school was found on August 1, 1951, feeding on anchovy in the region of Chetvertyi Kuril Strait; at the end of August, a herd more than 1,000 strong was sighted on the Pacific Ocean side of Iturup Island where saury concentrations were present. At the end of September, a large herd of these dolphins was noticed southeast of Shikotan Island where, too, a school of saury was encountered. Hunting on a small scale is resumed in the autumn months on the coasts of Japan (Klumov, 1959). Reproduction, growth, and development. Mating and parturition occur in summer. An embryo 22 mm long, detected in September, had grown to 370 mm by December and 761 mm by March. A male 124 cm long turned out to be a suckling calf (weight 29.5 kg). The ovaries of a 171-cm long female (weight 62 kg) did not contain corpora lutea but the Graafian follicles were well developed; this female was close to attaining sexual maturity (Houck, 1961). Females 180 cm or longer were already with embryos. In oceanariums, mating of female white-sided dolphins with male bottlenose dolphins has been observed time and again (Tomilin, 1962). The parasites of these dolphins have not been studied. (V.A.) Economic Importance These dolphins are regularly hunted along with other species only in the waters of Japan. Their annual catch does not exceed 1,000 head. The weight of the females varies from 67-93 kg (mean 80 kg) and of males from 58-84 kg (66 kg). They are of little economic importance. Hunting is carried out using the same small ships (20 to 30 tons) and the very same methods as practiced in hunting other species of dol- phins. The dorsal fin, flippers, and caudal flukes are first separated from the trunk of the killed animal. The skin is then peeled off in two layers together with the subcutaneous fat and the flesh is separated in large chunks from the vertebral column. The intestine, heart, and other inter- nal organs are then cut out and the ribs chopped. The flesh, heart, lungs, and liver are used as food. The skin is tanned to make low-quality leather goods. The blubber is melted into oil; the blubber from the head and in the hollow of the jaws can be converted into a high-grade machine oil. The skeletal bones as well as all the remnants are used for making fertil- izer meal (Wilke, Taniwaki, and Kuroda, 1953; Siebenaler and Caldwell, 1956; Klumov, 1959; Tomilin, 1962). Large schools of these white-sided dolphins prevailing in the waters of the Kuril Islands suggest the possibility of their organized hunting here. (V.A.) 501 501 672 BROADSNOUT DOLPHIN Lagenorhynchus (Peponocephala) electra Gray, 1846 1846. Lagenorhynchus electra. Gray. Zoology. Voyage Erebus and Terror, 1, p. 35. Type locality not established. 1848. Delphius pectoralis. Peale. U.S. Explor. Exped., 8, p. 32. Hawaiian Islands. 1868. Electra obtusa. Gray. Synopsis Whales Dolphins. Brit. Mus., 7. Substituted for Lagenorhynchus electra Gray, 1846. (V.H.) The broadsnout dolphin differs significantly from all other species of the genus Lagenorhynchus in several characteristics: it has no beak whatso- ever or such is barely visible and not demarcated from the forehead; a uniform arcuate line of the profile is formed from the blowhole to the edge of the mouth. The preorbital depression is larger than in the other species of the genus. The first three vertebrae, and not the first two as in the other species, are fused. The general body shape is more elongated, with a relatively longer caudal stem. The color is monochromatic, dark, without black, gray, or white fields; a light-colored (white) field is seen only in the cervical zone and on the breast; sometimes, a similar field is seen in the posterior section of the abdomen (Fig. 283). This is a somewhat peripheral form of the genus and reveals fea- tures of similarity and association with the genera Pseudorca (false killer whale) and Feresa (dwarf killer whale). (V.H.) Its range is known only from some fragmentary information. Its presence has not been established in USSR waters but residence in or transgressions into the Sea of Japan and waters of the Kuril Islands are possible. Outside the USSR, it is found in tropical seas up to north of 10° N lat. Its presence has been indicated in the waters of Senegal, the Gulf of Guinea, India, Indonesia, Hawaiian Islands, and Japan (transgression Fig. 283. Broadsnout dolphin, Lagenorhynchus (Peponocephala) electra (figure by N.N. Kondakov). 501 673 SS Va LN RS Е W BS % у) Я] ИИ GR ny Gis il sis Meat , ) ys \ И Fig. 284. Skull of the broadsnout dolphin, Lagenorhynchus (Peponocephala) electra (figure by N.N. Kondakov). of a herd of 500 animals in 1965 into Suruga Bay on the ocean side of Honshu; Nishiwaki and Norris, 1967). (V.H.) Biology is almost not known. Very rare. In January, 1963, fourteen dolphins were caught in a net in one of the bays on the Pacific Ocean side of Honshu Island. All of them showed their heads above the water most of the time, thus exhibiting near-total fearlessness of man. Held in an Oceanarium they exhibited no restlessness and were at peace with man (Nishiwaki, 1965). (V.A.) Genus of False Killer Whales Genus Pseudorca Reinhardt, 1862 1846. Phocaena crassidens. Owen. Hist. Brit. Foss. Mamm. p. 516. 1862. Pseudorca. Reinhardt. Overs. Danske Vidensk. Selsk. Forh., p. 151. (V.H.) Medium-size dolphins, with body length up to 6 m. The body is elongated. The small rounded head has a fairly large adi- pose body [melon]. The obtuse “beak” is barely perceptible. The upper jaws are slightly longer than the lower. The relatively small dorsal fin 502 674 lies midbody and is deeply notched posteriorly. The о are narrow and pointed at the tip. The body is black but the ventral side somewhat lighter. The rostrum is short, not longer than the cranium, but broad. The premaxillae are very broad and identical throughout their length. Their anterior margin is rounded. The reduced pterygoid bones are adjacent. Teeth a. Vertebrae 50-51. Phalangeal formula: I, 5, Пб_з› III;_¢, IV,_,, and V,_>. The sternum consists of four sections. Ribs 10 ‘pairs, of which four articulate with the sternum. False killer whales feed mainly on schooling fish and cephalopods. Mating and parturition are protracted. These whales are distributed in the warm and temperate waters of the Pacific, Indian, and Atlantic oceans. They are not caught in USSR waters but might possibly be found along the Kuril Islands and in the Sea of Japan and the Baltic Sea. Fossils of false killer whales have been detected in the Upper Pliocene of Europe and Japan. The genus comprises a single species: the false killer whale, P. cras- sidens Owen, 1816. (V.S.) FALSE KILLER WHALE Pseudorca crassidens Owen, 1846 1846. Phocoena crassidens. Owen. British Fossil Mamm. and Birds, p. 516. Lincolnshire, England (subfossil specimen). 1882. Pseudorca mediterranea. Giglioli. Zool. Anzeiger, 5, р. 268. Mediterranean Sea. (V.H.) Diagnosis Only species of the genus. Description The body is elongated, spindle-shaped, and the head relatively small. It somewhat resembles the killer whale but the body build is less compact, the head more elongated, and the dorsal fin considerably shorter. The flippers are not rounded but narrow and pointed (Fig. 285). Very light-colored star-shaped scars are often seen on the black sur- face of the body. A gray longitudinal band sometimes occurs on the midabdomen and in the region of the urogenital opening. Teeth usually number 5. Unlike those of the killer whale, their cross section is not oval but circular. 675 503 503 Fig. 285. False killer whale, Pseudorca crassidens (figure by М.М. Kondakov). The width of the rostrum increases with age and can be corre- lated with the intense growth of the adipose body [melon]. Cervical vertebrae 7, thoracic 10, lumbar 9-11, and caudal 22-24. The anterior and sometimes all the cervical vertebrae are fused. The indices of the main body measurements of the false killer whales as percentage of the total length of males (three specimens) with a body length of 279-465 cm and females (two) with a body length of 251 and 267 cm (Tomilin, 1957) are respectively: distance from tip of snout to blowhole 9.6-12.4 and 11.7-12.3, up to base of flippers 14.5 - 19.6 and 15.3-21.4, up to posterior margin of dorsal fin 51.2 -62.0 and 63.2 - 71.7; length of base of dorsal fin 15.2- 15.7 and 13.8 - 14.7, height of dorsal fin 8.3-8.6 and 7.4-8.3; length of flippers 11.8-13.2 and 11.5-12.6, maxi- mum width of flippers 4.7 -5.0 and 4.6-5.0; width of caudal flukes (from tip to tip) 24.0-24.4 and 20.9-21.4. The largest of the false killer whales caught measured 596 cm in length. Males are 0.6-1.0 m longer than females. False killer whales weigh up to 1.5 tons. The main dimensions of the skull (eight measurements) of adult false killer whales (Tomilin, 1957) are (in cm): condylobasal length 54-62, length of rostrum 27 - 30, width of rostrum at base 18-21, and length of lower jaw 46-51 (Fig. 286). The vertebral sections constitute (as percentage of length of the ver- tebral column): cervical 3, thoracic 17, lumbar 31, and caudal 49 (Slijper, 1936). (V.S.) Geographic Distribution Encountered in temperate and warm waters of the Northern and South- ern hemispheres (Fig. 287). 676 Geographic Range т the USSR Waters of the Kuril Islands (apparently the southern islands) and probably the Sea of Japan. Transgressions into our waters of the Baltic Sea are quite possible. Geographic Range outside the USSR Encountered in the Atlantic Ocean everywhere from the North Sea and North Carolina® to the South African and South American coasts (Tierra del Fuego). It has been reported on the eastern (Baltic) coasts of Den- mark, western and partly eastern (evidently the southernmost) coasts of Sweden, and on the Baltic coasts of the Federal Republic of Germany (Kiel Bay). Not reported on the coasts of Poland. It has been reported in the Pacific Ocean from Washington state in the north to the coasts of Peru, probably Chile, New Zealand, Australia, 222 Li 0222222 7 2 2042 == > метки 2222 WZ, TES APE LO A JAN EB ася о ЕЕТА ree 2... Ма 7 a » Ау \\ am See Meas Saheb ns м, * a ко т w & | wv А Te ` yf 503 Fig. 286. Skull of the false killer whale, Pseudorca crassidens (figure by М.М. Kondakov). 8 References to habitation farther north (Davis Strait and Aleutian Islands; Tomilin, 1957, 1962) have either not been confirmed (Murie, 1959; Hall and Kelson, 1959; Manvill and Joung, 1965), or have been questioned, or treated as erroneous (Hershkovitz, 1966). (V.H.) OE ROMO ily iy TAH “Teall | [lay fil ryt 9 и 1 | ПА Ни СЫН 1 || Wy Ils hill ‘ $5 We реет 4 ИН и 1000 0 1000 2000 3000 4000 5000 km Fig. 287. Species range of the false killer whale, Pseudorca crassidens (У.А. Arsen’ev). 504 505 506 678 and Tasmania in the south; it has been reported in the Indian Ocean in. the warm and temperate water zones. Numerous beached animals have been found on the coasts of Great Britain, Denmark, Holland, the Baltic and Mediterranean seas in the eastern part of the North Atlantic; in the western part, on the coasts of North Carolina and Florida states, Cuba, Venezuela, Brazil, and Argentina. In the Pacific Ocean, beached animals have been found on the Kuril Islands, the coasts of Japan, Galapagos Islands, California, and Mexico; and in the Indian Ocean, on the coasts of Sri Lanka, India, South Africa, and Australia (Tomilin, 1962; Mitchell, 1965; Fiscus and Niggol, 1965; Nishiwaki, 1966). (V.A.) Geographic Variation Not established. Biology As the range of the false killer whale is very large and instances of beaching (sometimes in large herds) are well known, this species could correctly be classed as a relatively abundant one. Food. Fish (e.g., haddock, cod, and salmon) and some species of cephalopods serve as the food of the false killer whale; these animals quite often come close to the coast in pursuit of food objects. It is at such times that they become stranded on the shoals. Behavior. False killer whales more often live far from the coasts in the open sea in small groups although sometimes they do form large herds (Fig. 288). Instances of large herds of these dolphins being cast on the beach are known. For example, in Mar del Plata in Buenos Aires province of Argentina, 835 false killer whales were stranded on a sandy beach on October 10, 1946 and perished soon thereafter. They suffered from extreme fatigue and had slowly sunk into the sand (Obruchev, 1949). Migrations of false killer whales have not been studied. Reproduction. Judging from the fact that embryos of different sizes are encountered simultaneously, mating and parturition are undoubt- edly protracted among these dolphins. An embryo 117 cm long was well pigmented while another embryo 160 cm long was almost mature. The largest of the embryos examined was 183 cm long. It has been reck- oned that females attain sexual maturity at a body length of 366 to 427 cm. An examination of many groups of false killer whales that per- ished through beaching established a male:female ratio of almost one in each herd. 505 Fig. 288. False killer whales at sea (figure by М.М. Kondakov). Enemies, diseases, parasites, mortality, and competitors. Skin parasites have not been reported but two species of helminths have. The nematode Anisakis (Anisakis) simplex Rudolphi, parasitizing the gullet, stomach, and intestine, has been found in ten other species of toothed whales, two species of baleen whales, and in Steller’s sea lion, as well as in the false killer whale. It has also been detected in various species of marine mammals in the North Sea, on the coasts of Kamchatka, Japan, and New Zealand. The acanthocephalan Bolbosoma capitatum Linstow parasitizes the intestine of the false killer whale, the pilot whale, and the sperm whale in the Atlantic Ocean and the Mediterranean Sea. (V.A.) Economic Importance There is no special hunting for the false killer whale; it is caught inci- dentally while hunting for other species. In some cases, the carcasses of beached animals are utilized. The overall weight of a false killer whale may reach 1.5 tons (Delamure, 1955; Sleptsov, 1955; Tomilin, 1957, 1962). (V.A.) Genus of Killer Whales Genus Orcinus Fitzinger, 1860 1846. Orca. Gray. Zoology. Voyage Erebus and Terror, I, p. 33. Orca gladiator Gray = Delphinus orca Linnaeus, 1758. Мот. ргаеосс. —Orca Wagler, 1830 = Hyperoodon Lacépede. 1860. Orcinus. Fitzinger. Wiss. Popul. Naturg. Saugeth., 6, p. 204. 507 680 1870. Gladiator. Gray. Proc. Zool. Soc. London, р. 71. Delphinus orca Linnaeus, 1758. 1933.? Grampus. Iredale and Troughton. Rec. Austr. Mus., 19 (1), p. 28. Delphinus orca Linnaeus, 1758. (V.H.) Dolphins of large proportions, being the largest members of the family. Body length up to 10 m. The head is somewhat flattened dorsoventrally. There is no “beak”. The broad flippers are oval in shape. The dorsal fin is very high, especially among males (up to 1.7 m). The overall color of the body along the upper side and flanks is black while the belly and neck are white. A white patch occurs on each side posterior to the dorsal fin and on the temple above the eye (sometimes these patches are lacking). The broad and flattened rostrum is slightly shorter or as long as the cranium. The maxillae are well developed and very broad. The occipital crest is absent in the young but high among the adults, especially males. The temporal fossa is large. The teeth are large, number Я, com- pressed front to back and have extremely well-developed roots. Verte- brae 50-52. The sternum consists of three or four sections.!° The length of the phalanges of the digits and the metacarpals is less than their width. Phalangeal formula: 15, Il,_7, Ш4_5, [V3_4, and V>_ 3. The killer whale is sarcophagous and attacks even large whales. The periods of mating and parturition are protracted. Gestation probably requires one year. These whales are distributed in the World Ocean except in glacial regions of the Arctic and the Antarctic. Fossil remains have been detected in the Middle Pliocene of Europe. The genus comprises a single species: the killer whale O. orca Lin- naeus, 1758. There is no special hunting of the killer whale. (V.S.) KILLER WHALE Orcinus orca Linnaeus, 1758 1758. Delphinus orca. Linnaeus. Syst. Nat., ed. X, I, 77. Eastern part of the northern half of the Atlantic Ocean (“Oceano Europaeo”). 3 The introduction of the name Grampus to denote the genus of killer whales is asso- ciated with so many inaccuracies that it must be considered irrational from the viewpoint of nomenclature. Yet the name (Grampus orca) did prevail in the literature for sometime. Today, as earlier, it is used only in reference to Risso’s dolphin (Grampus griseus). (V.H.) 10 According to other data, the sternum is not segmented (Sleptsov, 1955). 508 681 1789. Delphinus gladiator. Bonnaterre. Tabl. Encycl. Méth. Cétologie, p. 23. Spitsbergen, Davis Strait, New England. 1869. Orca ater. Cope. Proc. Ac. Nat. Hist. Philadelphia, 21, p. 22. North- ern Pacific Ocean (“Northwestern coasts from Oregon to Aleutian Islands’’). 1869. Orca rectipinna. Cope. I[bid., 21, р. 22. Californian coast. 1874. Orca ater var. fusca. Dall. In: Scammon. Marine Mammals of the N. W. coast of N. America, p. 298. Coasts of California and Oregon. (У.Н.) Diagnosis Only species of the genus. Description This whale has a strongly built body, a fairly large head, a large mouth, and is a powerful predator. Large rounded flippers (Fig. 289) are a characteristic feature. Another distinctive feature of this whale is the high dorsal fin, which is straight along the posterior margin. This whale exhibits distinct sexual dimorphism in the dimensions of the dorsal fin, which is considerably larger in the males. The size and relative dimensions of the flippers in both sexes and the caudal flukes among males increase with age. Unlike in other whales, the head in the killer whale (Fig. 290) becomes relatively shorter with advancing age while the caudal section becomes elongated, i.e., the elongation of the head is relatively less compared to the caudal section (Ivanova, 1959). The white coloration on the belly is divided posteriorly into three tongues, of which the two lateral ones (right and left) terminate behind the anal opening while the middle one may terminate at the same point or run along the right up to the caudal flukes (which then merge with the white coloration), or be altogether absent. Sometimes the right and left white tongues fuse posterior to the dorsal fin. Wholly dark-colored killer whales with only a white spot under the eye are encountered in the Far East. Unlike most other dolphins, the integument is characterized by a well-developed dermal layer with a dense network of fascicles of col- lagen fibers. Small processes project into the dermal papillae from the epidermal septa. Up to seven hairs are seen on the upper jaw of embryos. Sex- and age-related variations are distinctly seen in the skull. The temporal fossa and occipital crest enlarge with age. Among adult males, indices of the width of the rostrum are slightly higher than in the young. The lower jaw in males is relatively fonger than in females. The occipital crest of adult males is larger than that of females. 682 508 ‘Fig. 289. Killer whale, Orcinus orca (figure by М.М. Kondakov), A—male; B—female. 508 Fig. 290. Head of the killer whale, Orcinus orca (figure by N.N. Kondakov). 509 683 The teeth are covered with enamel and are very strong. When the mouth is closed, the teeth on the upper jaw fall in the gaps between the lower teeth, thus forming a powerful gripping apparatus. The teeth of the killer whale, compressed front to back, have apparently adapted to feeding on large quarry from which the killer whale can tear off large chunks of flesh. The foreteeth, inclined slightly forward and outward, are capable of withstanding extremely powerful jerks of the quarry while the middle and hind teeth perform the function of holding it firmly (Tomilin, 1957). The surface of the teeth is sometimes worn down to the pulp cavity. Vertebrae: cervical 7, thoracic 11-12, lumbar 10, and caudal 21-24. The first two to three vertebrae are usually fused. The percentages of the different sections of the vertebral column to its total length constitute: cervical 3.2, thoracic 22.9, lumbar 28.0, and caudal 45.9. Ribs 11-12 pairs, of which 5-7 anterior pairs articulate with the third or fourth section of the sternum (Tomilin, 1957). The maximum length of 320 male killer whales caught in the waters of Japan during 1948 through 1957 was 9.4 m, of 247 females 8.2 m. Among these killer whales, males with a body length of 6.3 m and females 6.0 were more common. Males of maximum proportions attain a body length of 10 m and females 8.2 m (Nishiwaki and Handa, 1958). The main body dimensions of males (based on measurements of two to four animals) and females (one to three animals) vary as follows (in cm): body length 579-830 and 495-670; distance from tip of snout to commencement of dorsal fin 245-400 and 335, up to blowhole 80-90 and 75, up to anal opening 221-273 and 225; length of base of dorsal fin 77-91 and 62, height of dorsal fin 121-170 and 52-95; and width of caudal flukes 195-279 and 99 - 139. The main measurements of the skull (Fig. 291) of five adult males and one adult female killer whale (Tomilin, 1957) are respectively (in cm): condylobasal length 100-112 and 91, zygomatic width 68-78 and 57, length of rostrum 51-57 and 48; width of rostrum at base 34-37 and 27, and length of lower jaw 85-95 and 74. (V.S.) Geographic Distribution The killer whale represents a cosmopolitan species. It inhabits coastal and oceanic waters of the world’s oceans, including the Arctic and Antarctic seas. Geographic Range in the USSR Barents Sea, northwestern and western parts of the Kara Sea (does not transgress into its eastern part nor into the Laptev Sea), coastal waters of 509 684 SSS SSS Fig. 291. Skull of the killer whale, Orcinus orca (figure by N.N. Kondakov). Novaya Zemlya, Murman coast, Kil’din Island, Rybachii Peninsula, and the White Sea (Fig. 292). Transgressions into our waters of the Baltic Sea are highly possible (transgression into bay near Gdansk is known). In the Pacific Ocean waters of the Soviet Union, this species is dis- tributed everywhere in the seas of Japan and Okhotsk, around the Kuril Islands, in the Bering Sea, near the Commander Islands, in the Bering Strait, and in the southern part of the Chukchi Sea (probably north of 70°N lat.). Geographic Range outside the USSR From the coasts of Spitsbergen, Greenland, and Baffin Bay in the north to the southern waters of the Atlantic Ocean. It is quite common in much of the eastern and western halves of the Atlantic (coastal regions of Great Britain, Norway, Holland, France, western Africa, Atlantic coast of North and South America, and the Mediterranean Sea). It inhabits the westernmost, southwestern, and southern parts of the Baltic Sea up to Gdansk (Danzig), including the bay (Fig. 293). In the Pacific Ocean, it is encountered off the coasts of the Korean peninsula, Japan, Aleutian Islands, and Alaska, including its northern half (noticed off Cape Barrow, 71°24’N lat.), off the Pacific coast of North America and Mexico, and along the coasts of South America from Panama to Tierra del Fuego. It lives in the Indian Ocean and along ‘(лэмэзту У) USSN Э41 Ul ‘9240 5ии1240 ‘ЭТеЧм ло BY1 JO suey “767 “Bly отб от 691 ах | Wy 0001 OSL 005 055 O 052 ‘(Ad цэзту УД) 0240 5пи1240 ‘TRUM ты эЧ1 Jo э8иел $э0э4$ ‘©6291 119 wy 000$ 006 0006 0005 COOL 0 0001 512 687 the coasts of Australia and New Zealand. It is widely distributed in the Antarctic where it reaches the high southern latitudes, right up to the zone of permanent ice. (V.A.) Geographic Variation In spite of the extensive distribution of this species in the World Ocean, no geographic variation has been established. Sometimes the Pacific killer whale is regarded as a species (Subspecies), Orcinus rectipinna Cope, 1869, but the characteristics of this form have not been established, albeit mention has been made that its dorsal fin is not as high as in the Atlantic species (Hall and Kelson, 1959). (V.A.) Biology Population. One of the abundant species of dolphins. It is regularly encountered in the World Ocean (usually in small groups) and is fairly uniformly distributed over its immense range. Food. The killer whale is the only true carnivore in the order of cetaceans and even consumes warm-blooded animals. Its food is quite diverse: chum salmon, chinook salmon, coho salmon, cod, capelin, hal- ibut, skate, shark, herring, smelt, etc. Cephalopods are consumed quite often and sometimes various types of marine birds. The killer whale is known to consume seals and small dolphins and to attack fur seals, Steller’s sea lion, and even walruses (an instance of a group of killer whales attacking a herd of walruses was observed from an airplane; Zenkovich, 1938b). There are frequent references in the literature to the killer whale lying in wait around the coastal rookeries of fur seals and Steller’s sea lion but the many observations reported in recent years do not confirm this information (V.A. Arsen’ev). Reports of the killer whale attacking balsen whales and tearing out their tongues and large chunks of blubber, and even killing large animals, have been copied from book to book from the middle of the last century without crit- ical review. Even a recent and thorough publication (Tomilin, 1957) describes the methods of killer whale groups attacking large whales, pre- senting information copied from various sources. Such reports give rise to considerable doubt. A study of the stomach contents of several hundreds of killer whales caught in a ten-year period in the waters of Japan revealed that squids (octopuses to a very small extent) and fish (including sharks) represent the main food while marine mammals play a secondary role. The number of stomachs with remnants of dolphins was more than double the num- ber of stomachs with remnants of seals. Among fish, cod predominated, 513 688 followed by (using the source terminology) flatfish (probably flounder, halibut, etc.). Sardines, salmon, and tuna were of lesser importance. Some stomachs revealed mackerel, greenling, sea perch, and bonito, which have no practical importance as food for the killer whale. Dolphins of three species exclusively represent the cetaceans found in the stomach rem- nants: Dall porpoise (Phocoenoides dalli dalli and Р. 4. truet) found in the stomach of killer whales on Hokkaido coasts and blue-white dolphin (Stenella coeruleoalba) in killer whales from the Pacific waters off Hon- shu. There were some finds of pilot whales, beaked whales, and two cases of sei whales (chunks of flesh). Young killer whales up to 4.3 m long feed exclusively on squids and fish. They begin attacking seals and dol- phins as their body proportions increase and thus remnants of seals and dolphins are found in the stomach of only older killer whales (Nishiwaki and Handa, 1958). In July and August, 10 stomachs of killer whales were found filled with remnants of fish and squids (11 other stomachs in the same months were empty). No remnants whatsoever of marine mammals were found (Ivanova, 1961c). In the eastern half of the North Pacific Ocean, 10 stomachs of killer whales were investigated. Of these, one was caught close to Kodiak Island in the Gulf of Alaska, five near San Francisco, and four in Californian waters. Two of these stomachs were empty while the remaining eight con- tained the remnants of marine mammals (California sea lion, Steller’s sea lion, elephant seals, two species of dolphins, and Minke whale), hal- ibut, ocean sunfish, shark, and squids. The frequency of encounter of the remnants of these food objects is shown below. Quantitative characteristics of the food of killer whales (number of finds) (Rice, 1968) California sea lion, Zalophus californianus Steller’s sea lion, Eumetopias jubatus Elephant seal, Mirounga angustirostris Harbor porpoise, Phocoena phocoena Dall porpoise, Phocoenoides dalli Minke whale, Balaenoptera acutirostrata Opah, Lampris regius Pacific halibut, Hippoglossus stenolepis Carcharinid, ? Prionace glauca Squids BPNFPNPNN IA W The above eight killer whales with remnants of marine mammals in their stomachs included six adult males, one adult female. (California), 689 and one immature male (Gulf of Alaska). The stomach of five adult males contained only the remnants of marine mammals while the sixth addi- tionally had remnants of squids. The stomachs of the adult female and the immature male had only the remnants of fish, including sharks. It may be assumed that male killer whales feed predominantly on marine mammals. The mammals were represented mostly by the remnants of one, less frequently two, and only in one case (elephant seal) four animals in the same stomach. All of them were thoroughly digested, since mostly the teeth, nails, and sometimes ribs or skin remnants were found. Minke whale was represented by baleen plates and chunks of blubber. Layer formations on the teeth helped to establish the age of some of the pin- nipeds consumed by killer whales. Thus the stomach of a male killer whale 7.6 m long contained the remnants of a nine-year-old sea lion, of another male killer whale 6.86 m long the remnants of a one-year- old Steller’s sea lion (probable weight of the animal 90 kg), and of yet another male 7.24 m long a two-year-old (probably) Steller’s sea lion (Rice, 1968). The hunting of sea lions by killer whales has been described as follows: a group of five killer whales chased a large male sea lion on May 25, 1965 in the San Francisco region. The adult killer whale in the group (it also contained calves) continuously lay under the sea lion, or above it when the sea lion dived. From time to time, the killer whales struck the quarry with their bodies or attempted to crush it. The sea lion jumped from side to side, sometimes leaped out of the water, and eventually became exhausted. At this point one killer whale swam to its right, another underneath it, while a third grabbed the quarry with its teeth and towed it under water. That was the last sighting of the sea lion on the water surface. This happened very close to a ship following the attacking killer whales; the latter paid no attention to the ship behind them. In another case, one of the six killer whales (the group contained two calves and one adult male) appeared on the water surface holding a small sea lion in its mouth. The nature of mammal remnants seen in the stomach of killer whales suggests that they dismember large animals before ingestion but swallow smaller prey entire (Rice, 1968). Reports of killer whales attacking large whales require documentary proof. In the Antarctic, near a whaling base, groups of killer whales were noticed floating alongside fin whales, humpback whales, and other species of baleen whales time and again. Yet no attempts by the killer whales to attack the others were ever recorded. Concomitantly, the other whales (even when alone) never exhibited signs of restlessness on the approach of a herd of killer whales. However, killer whales regularly tore out the tongues of dead and air-filled whales with mouths wide open 514 514 690 (Fig. 294). The carnivores were bold enough to tear out the tongues of killed whales tied to the board of a whale boat towing them to the base. Often the killer whales almost wholly consumed the tongues of whale carcasses stocked as feed for the whale base. Attempts to drive away the killer whales by throwing various objects at them from the whale boat or even by opening fire were almost of no avail. The tongue of whales weighs two to three tons and contains a large amount of fat; thus killer whales quite often cause serious losses to whale hunters by snatching away the valuable tongues. Yet there are no reports of killer whales attempting to tear off chunks of blubber from a whale carcass or to snatch its fins (V.A. Arsen’ev). Apparently, there is no justification for ascribing the various damages caused to dead whales to the activity of killer whales. The killer whale is generally described as an unusually greedy car- nivore and fantastic figures are cited about the quantum of food found in its stomach. The stomach of a 6.5-m long killer whale purportedly contained “five or six seals and semidigested remains of another six or seven seals and thirteen harbor porpoises; in addition, there were two nearly whole seals in the gullet” (Tomilin, 1957). If the average weight of these animals is taken as even 30 kg, the stomach contents of this killer whale should have weighed at least 600 kg, while the contents of Fig. 294. Killer whales tearing out the tongues of dead whales (photograph by V.A. Zemskii). 515 515 691 а closely packed stomach of a 20-m long fin whale would weigh about 1,000 kilograms. The killer whale is no doubt a powerful and dangerous carnivore but its food habits have yet to be properly studied. Behavior. Killer whales usually live in groups of a few to 100 or more animals. They are encountered in the open sea as well as close to coasts. Of the 567 killer whales caught in the waters of Japan, 391 (69%) were killed within the 30-mile zone, 101 (17.8%) at a distance of 31 to 60 miles, and 36 (6.3%) beyond 60 miles from the coasts. The distance at which the remaining 39 killer whales were killed could not be ascertained. However, the spread of the party hunting for killer whales (along with other species of cetaceans) is also of some importance in such cases (Nishiwaki and Handa, 1958). Some instances are known of the transgression of killer whales into a river at a distance of a few tens of kilometers from the estuary in which the animals were confined to a freshwater zone, sometimes for a few weeks. While feeding, killer whales move at 5-7 miles per hour and during migrations at 15-17 miles per hour. It has been noticed quite often that large groups of killer whales move in a broad front or even in columns (Fig. 295). They usually spend 2 to 5-6 min under the water and perform five to six brief spells of exhalation/inhalation in the interval between Fig. 295. Killer whales at sea. Pacific Ocean, 1948 (photograph by М.М. Sleptsov). 516 692 dives. Sometimes they remain on the water surface for quite a long time, with their high dorsal fins visible above the sea level throughout this period. The blow produced by the killer whale is rather low, not more than 2 m high. The strong affinity of adult animals for each other and their calves is a characteristic feature. It is difficult to break up a herd of killer whales and quite often many animals of a given group can only be caught on just one side of the ship. The animals are more active in warm water (20- 25°C) and become sluggish as the water temperature drops to 10- 15°C. It is easy to catch the killer whale at this time and hence most of those caught in the waters of Japan come from the northern part of the country. This fact notwithstanding, killer whales do transgress into polar waters and do not shirk from ice, although none have been sighted among compact ice. Migrations. The courses and periods of seasonal migrations of killer whales have not been studied. They are sighted over much of the range throughout the year but in the polar sections of the Northern and Southern hemispheres apparently only in the summer months. The killer whales seen on the coasts of the Chukchi Peninsula in June abandon these waters by November-December (Nikulin, 1946). They inhabit Hudson Bay, Fox Basin, and Davis Strait too only in the summer period. Killer whales, like other species of cetaceans, are encountered in the waters of Antarctica only in the summer months and abandon them altogether in winter. Killer whales may continue to remain in the winter months even at high latitudes only at such places in the oceans which fall under the influence of powerful warm-water currents (for example, the Gulf Stream). In the waters of Japan, the bulk of the killer whales are caught from April through November inclusive while the catch is insignificant from December through April. More killer whales are caught on the coasts of Hokkaido (northern coast, Sea of Okhotsk; southeastern coast, Pacific Ocean), fewer on the northeastern coast of Honshu (38- 40° N lat.), and even fewer on the southeastern coast of this island (33 -35° М lat.). In the Sea of Japan, there is virtually no hunting of the killer whale (Nishiwaki and Handa, 1958). Reproduction. Mating of killer whales was observed in June and July and births in spring and summer (Scheffer and Slipp, 1948) although the view is prevalent that parturition occurs mainly in autumn. Embryos 27 to 222 cm long were found in July- August among killer whales of the Kuril Islands. At this time, simultaneous with gestating females, four lac- tating mothers were found. They had probably calved in July (Ivanova, 1961с). A nearly mature embryo measured 208 cm in length while the 516 "693 Fig. 296. Killer whale on the deck of a whaling base (photograph by V.A. Zemskii). largest was 274 cm long. The smallest of the measured calves were 236, 246, 251, and 274 cm. Such measurements are evidently characteristic of newborn calves. It has been suggested that gestation among the killer whales can extend for 16 months but the maximum number of births has been recorded in May to July (an instance of mating was photographed from an airplane on June 22, 1957) although mating and parturition can occur in the other summer months as well. Additional documented infor- mation is required to resolve the questions of the duration of gestation and periods of mating and parturition. The duration of lactation has not been established. By the time it is one year of age, the killer whale calf has reached a length of about 350 cm. The average length of 320 males caught in the waters of Japan was 6.4 m, with the largest 9.45 m long. The average length of 247 females was 6.1 m, with the maximum length at 8.23 m. Most of the males caught varied in length from 5.5-7.6 m and females 5.5-6.7 m. Of the 567 killer whales caught from 1948 through 1957, 320 were males (56.4%) and 247 females (43.6%). The sex ratio is evidently close to 1:1 (Nishiwaki and Handa, 1958). Enemies, diseases, parasites, mortality, and competitors. The killer whale has no enemies. The diseases known in them are bone tumors 517 694 (detected in skeletons preserved in museums) and dental caries. Ectopar- asites are not known. Eight species of endoparasites have been reported: two species of trematodes, one species of castode, three species of nema- todes, and two species of acanthocephalans. The trematode Fasciola skrjabini Delamure was found in the bile ducts of the liver of the killer whale and the Minke whale in the North Atlantic. The intestine of a killer whale in the fore-Kuril waters revealed the presence of the trematode Leucasiella subtilla A. Skrjabin. The ces- tode Trigonocotyle spasskyi Gubanov was found in the small intestine of whales only from the Sea of Okhotsk (Kuril waters). The widely preva- lent nematode Anisakis (Anisakis) simplex Rudolphi, parasitizing the gul- let, stomach, and intestine, was detected in 10 other species of toothed whales, two species of baleen whales, and in Steller’s sea lion in addition to the killer whale. The nematode Ана pacificus A. Skrjabin was found in the stomach of killer whales (and also sperm whales and fin whales) in the waters of the Kuril Islands. Another species of nematode, Anisakis sp., found in the killer whale, has eluded precise identifica- tion. The acanthocephalan Bolbosoma physeteris Gubanov, parasitizing the intestine was found in killer whales from the Sea of Okhotsk and Bol- bosoma nipponicum Yamaguti from the Kula Gulf [?] (Margolis, 1954; Delamure, 1955; A. Skrjabin, 1958, 1959, 1960). Field characteristics. A very high dorsal fin, rising to a height of over 1 m in males, distinguishes the killer whale from all other cetaceans. At a close distance, bright white oval patches can be seen on the temples and a large white patch posterior to the dorsal fin. Killer whales are usually confined to groups. Quite often, the groups present a broad front or move in a column. (V.A.) Economic Importance In spite of the comparative abundance of killer whales in the World Ocean, their hunting has acquired no economic importance. Killer whales are mostly caught casually during hunts for other cetaceans, such as beaked whales, Minke whales, and various other species of dolphins. A fairly organized hunting of killer whales exists only in Norway and Japan with whaleboats specially designed for small whales (Minke whale, bot- tlenose whale and other beaked whales, pilot whale, and killer whale). Data on killer whale hunting for several years are given in Table 40. As mentioned, Norwegian and Japanese hunting is carried out in specially designed craft equipped with small-bore harpoon guns. In the Soviet Union, the killer whale is hunted from the large whaleboats used in hunts for large whales in the Far Eastern waters as well as in the 518 695 Table 40. Magnitude of killer whale hunting (international whaling statistics) Year Norway Japan USSR Year Norway Japan USSR 1948 27 27 3 1959 69 36 36 1949 34 43 28 1960 82 48 45 1950 12 18 24 1961 111 54 4 1951 24 67 23 1962 124 47 7 1952 13 54 19 1963 90 43 4 1953 9 65 25 1964 77 99 1954 13 109 4 1965 104 169 1955 26 85 15 1966 161 137 9 1956 40 38 67 1967 36 101 4 1957 48 78 30 1968 86 22 1958 39 73 25 Antarctic. In other countries, some stray killer whales are caught inci- dentally. The fat of the killer whale, like that of other toothed whales, is used for commercial purposes. The skin of the killer whale is regarded as superior to that of other toothed whales and can be used for producing leather goods. The rest of the carcass is used in the preparation of feed or in fertilizer meal. (V.A.) Genus of Risso’s Dolphins Genus Grampus Gray, 1828 1828. Grampus. Gray. Spicilegia. Zoologica, I, p. 2. Delphinus griseus G. Cuvier, 1812. 1933. Grampidelphis. Iredale and Troughton. Rec. Austral. Mus., 19, р. 31. Substituted for Grampus Gray, 1846.!! (V.H.) Dolphins of medium proportions, with a body length up to 430 cm. The head is rounded anteriorly and no “beak” is perceptible. The high dorsal fin, located almost midbody, is notched along the posterior margin. The flippers are long and narrow. The body color is basically gray. The rostrum is broad; its length is shorter than the cranial section of the skull. The proximal portions of the broad premaxillae form a prominence. The pterygoid bones are adjacent. The nasal and frontal bones form a crest posterior to the nares. The mandibular symphysis is short. In most of these animals, teeth are present only in the anterior portion of the lower jaw in two to seven pairs (0-2 in the upper jaw). И See footnote on р. 680. 519 696 Vertebrae 68 - 69. Phalangeal formulae: [5, Пу - ;9, Шов, [Уз - $, and У.. Very little is known about the biology of these animals. They feed on cephalopods and live singly or in small groups. They inhabit the warm and moderate waters of both hemispheres but are few in numbers everywhere. They are not caught in the waters of the USSR but might possibly occur in the waters of the Far East. The genus comprises a single species: Risso’s dolphin, Grampus griseus G. Cuvier, 1812. (V.S.) RISSO’S DOLPHIN Grampus griseus G. Cuvier, 1812 1812. Delphinus griseus. G. Cuvier. Ann. Mus. Hist. Nat., Paris, 19, p. 13. Brest, France. 1822. Delphinus rissoanus. Desmarest. Mammalogie, p. 519. Mediter- ranean Sea, Nice. 1866. Globicephalus chinensis. Gray. Cat. Seals and Whales Brit. Mus., р. 323. China Sea. 1873. Grampus stearnsii. Dall. Proc. Calif. Ac. $с., 5, р. 13. Monterey, California. (V.H.) Diagnosis Only species of the genus. Description Body compact. The highly developed adipose body [melon] imparts a circular shape to the head while the frontal portion slightly projects anteriorly. The lower jaw is shorter than the upper. The oral cavity runs obliquely upward from front to back. The flippers are long, constituting 13.9 to 19.6% of the body length, narrow, crescent-shaped, and pointed at the ends (Fig. 297). The body color varies from gray to blackish-gray, gradually turning lighter from the dorsum toward the belly. The head is also lighter in color than the dorsum but all the fins are the same color as the dorsum. Small light-colored spots and bands on the skin are the result of various damages. Hairs (up to eight) are sometimes preserved on the head of young animals. The diameter of the teeth of adult dolphins can reach 1.5 cm and the height almost 4 cm (root and crown). The loss of teeth in the upper 519 697 ЕЕ, Е ке Ty, Lie И, ИДЯ Wi Wilf Mt Wife йе / И И Hl И, И fil yy Lt Whee oy Ц AY ma ge ИИ И и ‘A у (er STEM К и. Ra \ Ч ча He: р ips ii H f Boas" ve re ‘‹ ЕН Tiss Fig. 297. Risso’s dolphin, Grampus griseus (figure by N.N. Kondakov). jaw is a recent phylogenetic phenomenon and apparently the result of these animals feeding on cephalopods (Tomilin, 1957). Cervical vertebrae 7, thoracic 12-13, lumbar 18-19, and caudal 30-31. The sections of the vertebral column constitute the following percentage ratio to its total length: cervical 2.5, thoracic 24.5, lumbar 29, and caudal 44 (Slijper, 1936). The basic body measurements of three adult female Risso’s dolphins (Tomilin, 1957; Pilleri, 1969) are respectively (in cm): body length 330, 320, and 275; distance from tip of snout to blowhole 43 and 48, up to dorsal fin 127, 119, and 95, up to base of flippers 56 and 64; length of flippers 54, 61, and 45; maximum width of flippers 22 and 17; height of dorsal fin 34, 41, and 40; length of base of dorsal fin 56 and 35; and width of caudal flukes 74 and 79. The average skull measurements based on nine skulls (Fig. 298) of adults (Tomilin, 1957) are (in cm): condylobasal length 50, length of ros- trum 25, width of rostrum at base 20, and length of lower jaw 39. (V.S.) Geographic Distribution Warm and temperate waters of the Northern and Southern hemispheres. Geographic Range in the USSR (Fig. 299) Coastal waters of the Kuril range, mainly its southern part (Iturup and Shpanberg Islands)!?, and the Commander Islands. 2 Finds reported for the Kuril Islands (Sleptsov, 1952) have yet to be confirmed with specimens. 520 519 698 Geographic Range outside the USSR (Fig. 300) Gulf of Maine and the coastal waters of New Jersey state along the American coast of the North Atlantic (the range may be more extensive but precise data are not available). In the eastern part of the Atlantic Ocean, the range extends from the North Sea (waters of Great Britain, Schleswig in the Federal Republic of Germany, Denmark, and western coast of southern Sweden in Bohus Bay) and Ireland down to South Africa. Mediterranean Sea in the east including the Adriatic. In the east- ern part of the Pacific Ocean, from British Columbia to Baja California (Mexico) and waters of the Chilean coast; in the western part of this ocean from the Commander Islands (?) and Kuril Islands and Japan to the seas of China. Waters of New Zealand and Australia (New South Wales), the Indian Ocean, and the Red Sea. (V.A.) Geographic Variation Not known. $ & к \ NAY tiga git .% ^ (и of all ie [О } Ry) Fig. 298. Skull of Risso’s dolphin, Статриб griseus (figure by М.М. Kondakov). 522 520 699 Biology In all probability, this species is not very numerous throughout its range. Some species of cephalopods constitute its main food. Remnants of other food items have not been detected in the stomach of Risso’s dolphins. In search of cephalopods, this dolphin makes comparatively long dives and remains on the water surface longer than other species. It mainly inhabits the open seas. In most cases, it swims singly, in pairs or in small groups. These dolphins gather in herds of a few tens of animals only occasionally at places of food concentration. Fig. 299. Range of Risso’s dolphin, Grampus griseus, in the USSR (V.A. Arsen’ev). ‘(Ad UasIy УЛ) 5725148 зпашиоло ‘ичАрюр $,08sty jo э8иел $э10э4$ ‘00$ ‘314 155 SSS SS Wy 0005 000% 0005 0005 0001 0 0001 701 Migrations have not been established. It is known that these dol- phins live year-round in the Mediterranean Sea and in European waters, although some variations in their numerical strength have been recorded in different months. According to some fragmentary information, parturition -occurs in the winter months in comparatively warm waters. The newborn calf aver- age 150 cm in length. Three species of endoparasites are known. The cestode Phylloboth- rium delphini Bosc localizes in the skin of Risso’s dolphin and also six other species of toothed cetaceans, the bowhead whale, and Weddell’s seal. It has been detected in these animals at many points in the Atlantic Ocean, the Mediterranean Sea, waters of the Commander Islands and Australia, the Pacific Ocean, and the Antarctic. The nematode Crassi- cauda grampicola Johnston and Mawson was detected only among Risso’s dolphins in the waters of Australia. Stenurus minor Kuhn parasitizes the bronchi, tympanic cavity, and blood vessels (Delamure, 1955). Risso’s dolphin is of no economic importance. (V.A.) Genus of Pilot Whales Genus Globicephala Lesson, 1828 1828. Globicephala. Lesson. Nat. Hist. Mamm. Diseaux Dépuis 1788, Cétacées, p. 441. Delphinus deductor Scoresby = Delphinus melas Traill. 1828. Globicephalus. Lesson. Férussac Bull. Sci. Nat., 16, p. 116. Substi- tuted for Globicephala Lesson, 1828. 1884. Globiceps. Flower. Proc. Zool. Soc. London, 1883, p. 508. Substi- tuted for Globicephala Lesson, 1828. Nom. ргаеосс. (V.H.) Large dolphins, with a body length of up to 6.5 m. The head is rounded and a beak faintly perceptible. The dorsal fin is low with a curved posterior margin. The flippers are narrow and long. The body is dark, grayish-blue, or black. The skull is broad and slightly flattened dorsoventrally. The rostrum is almost as long as the cranium. The premaxillae are very broad. The pterygoid bones are adjacent. The nasal bones projects into the frontal and lie above the level of all the other bones. Teeth 7-1, present in the anterior half of the jaws. With advancing age, the teeth may wear out and fall out. Vertebrae 58-59. The sternum comprises three to four sec- tions. Five of the 11 pairs of ribs articulate with the sternum. Phalangeal formula: 5-4, Ig-44, HI9-1,, [М ->. These whales feed on large invertebrates (mainly cephalopods) and schooling fish (teutho-ichthyophagous). Periods of mating and 523 702 parturition are protracted. Gestation and lactation continue for about a year each. Members of this genus are confined to large herds of a hundred or more animals. The range of these whales covers all the seas except the polar. In the USSR, they might possibly occur in the Barents Sea and in the waters of the Far East. Fossil remains have been detected in the Pleistocene of North America. The genus comprises a single species: the (common) pilot whale, G. melaena Traill, 1809. Sometimes pilot whales are classified into two species, G. melaena and G. macrorhyncha Gray, 1846 (Rice and Scheffer, 1968*), or into three species, i.e., С. scammoni Cope, 1869, in addition to the two aforesaid. (V.S.) PILOT WHALE Globicephala melaena Traill, 1809 1809. Delphinus melas. Traill. Nicholson’s Journ. Philos., 22, р. 81. Pomona Island, Orkney Islands. 1812. Delphinus globiceps. G. Cuvier. Ann. Mus. Hist. Nat., Paris, 19, р. 14. France. 1820. Delphinus deductor. Scoresby. Account Arctic regions, I, p. 496. North Atlantic. 1824. Delphinus grinda. Lungbye. Kongl. Danske Vedensk. Selsk. Afh. I, р. XI. North Atlantic. 1846. Globicephala sieboldii. Gray. Zoology Voyage Erebus and Terror, I, p. 142. Near Nagasaki, Japan. 1869. Globicephalus scammoni. Cope. Proc. Ac. Sc. Philadelphia, 21, р. 21. California (10°N lIat.). 1871. Globicephalus sibo. Gray. Suppl. Cat. Seals and Whales Brit. Mus., р. 85. Japan. 1898. Globicephala melaene. Thomas. Zoologist, (4), 2, p. 99. Mascu- line form melas for the feminine of the genus, conforming to the generic name. (V.H.) Diagnosis Only species of the genus. Description The body is elongated and slightly thickset in the anterior half. The head is rounded with a projecting frontal part as a result of the highly 524 523 703 developed adipose body [melon]. The oral cavity runs from front to back with an inclination of roughly 30 - 45° to the longitudinal axis of the body from the bottom upward. The upper jaw is longer than the lower one. The blowhole is slightly displaced left of the midline. The long and low dorsal fin is located almost at the boundary of the anterior third of the trunk. The length of the fin at the base is not less than 1.5-2 times its height. Long flippers reach one-fourth of the body length (Fig. 301). The upper portion of the body is darker in color than the under- side. A narrow gray band enlarges into a patch between the flippers midabdomen and on the throat. A newborn pilot whale is gray but soon turns black. With age, high longitudinal ventral and dorsal keels form on the caudal stem of males. The adipose body [melon] of the head and the hump between the head and dorsal fin are larger in males than in females. Embryos sport three to six hairs on each side of the snout. Cervical vertebrae 7, thoracic 11, lumbar 12-14, and caudal 28-29. The first five or six cervical vertebrae are fused. The main body measurements of an adult male pilot whale caught off the coast of Virginia (North Atlantic) (Tomilin, 1957) are (in cm): body length 465; distance from tip of snout to blowhole 54, up to anal opening 318, and up to base of flippers 91; length of flippers 76, maximum width of flippers 25; length of base of dorsal fin 79; height of dorsal fin 35; and width of caudal flukes 117. The largest male measured 6.5 m in length and the female 6.1 m. The main measurements of six skulls (Fig. 302) of adult pilot whales averaged (Tomilin, 1957) (in cm): condylobasal length 63 (six measure- ments), zygomatic width 41 (one), length of rostrum 32 (six), width of rostrum at base 23 (six), and length of lower jaw 49 (two). Fig. 301. Pilot whale, Globicephala melaena (figure by N.N. Kondakov). 524 704 Fig. 302. Skull of pilot whale, Globicephala melaena (figure by М.М. Kondakov). The relative dimensions of the rostrum, the width of the skull at the level of the orbits, and the width of the rostrum at its base and center increase with age (Tomilin, 1957). (V.S.) Geographic Distribution Almost all the seas and oceans from the Arctic to the Antarctic except the frozen seas of high latitudes. Geographic Range in the USSR (Fig. 303) Southwestern part of the Barents Sea, especially the waters of Rybachii peninsula (irregular transgressions; Nemirovich-Danchenko, 1877; Chapskii, 1941), waters of the Kuril range, and possibly southeastern Kamchatka, Commander Islands, and the Sea of Japan (La Perouse Strait). Geographic Range outside the USSR In the North Atlantic it is found along the American coast from Delaware Bay to Newfoundland, Labrador, Davis Strait (up to 70°N lat.) and the southeastern coast of Greenland (67°N lat.). In the eastern part it occurs 705 ‘(лэ.цэзгу У) USSN Э41 Ul 9720]2\ vjoydasigopy ‘ayeyM 1014 до э8иеч ‘50$ ‘8 97S 525 706 from Iceland and northwestern Norway along the entire coast south up to Madeira,!? Mediterranean Sea. In the south, evidently inhabits waters up to 60°S lat. in the Atlantic (Fig. 304). In the North Pacific Ocean, up to the waters of southeastern China to the west and south; in the eastern part of the ocean from the Aleutian Islands and Alaskan peninsula down to Mexico (Baja California). In the tropical zone, the pilot whale is probably distributed over large expanses far away from the coasts. In the Southern hemisphere, it is known on the coasts of India (Bay of Bengal), Java, Guatemala, Ecuador, Peru, Chile, Australia, Tasmania, New Zealand, Kerguelen Island, and South Africa. The range apparently reaches 50 to 60°S lat. (Sergeant and Fisher, 1957; Tomilin, 1957; Brown, 1961; Hershkovitz, 1966; Nishiwaki, 1967). (V.A.) Geographic Variation Some authors recognize up to five or six subspecies but their validity (even of the two subspecies mentioned below, the most sharply mani- fest) is usually questioned or contradicted (Hershkovitz, 1966). Distant migrations characteristic of the species result in a thorough mix-up over the entire extremely extensive but essentially single range. Deviations in the picture of distribution of the various forms, especially of those given below, are highly significant (see Tomilin, 1957, 1962; Hershkovitz, 1966). Two subspecies can be recognized in the waters of the Soviet Union: 1. Common, or Atlantic pilot whale, С. т. melaena Traill, 1809 (syns. melas, globiceps, deductor). Thirteen pairs of teeth in each jaw; the white band on the ventral side of the body is broadened in the region of the flippers, and a large white, anchor-shaped patch occurs on the throat. Southeastern part of the Barents Sea, especially waters of the Rybachii peninsula. 13 References to habitation in the Baltic Sea are very general and neither positive nor reliable. They are mainly communications from one author to another and it is very difficult to identify the actual sources. More positive references are available only for the waters of Denmark (Jutland peninsula) and southwestern Sweden, i.e., the same section of the coast facing Jutland. Such references usually extend to the entire Baltic Sea. For the waters of the German Democratic Republic and Poland, there are no direct references to the habitation of the pilot whale and this species is not included among the cetaceans inhabiting these waters (Gentshel’, 1937; van den Brink, 1958; Koval’skii, 1964; Siivonen, 1968). Transgres- sions into the basin of the Baltic Sea proper might occur very rarely but the appearance of this animal in our waters far into the northeast is highly improbable. (V.H.) 707 “(лэ.чэзгу У’) Duanjau п]244э2190]0 ‘этеЧчм 1014 ay) Jo э8иел $э0э4$ “pog “314 528 708 Outside the USSR, it is encountered in the North Atlantic Ocean up to southern Greenland (almost 70°N lat.).!4 2. Black, or Pacific pilot whale, С. m. sieboldii Gray, 1846 (syns. scam- moni, sibo). Seven or eight pairs of teeth in each jaw. Color black, with no white markings. Waters of Kuril Islands (not known more accurately), probably Kam- chatka, Commander Islands, and the Sea of Japan (La Perouse Strait). Outside the USSR, this subspecies is encountered in the North Pacific Ocean up to the Yangtze estuary in the west and from the Aleu- tian Islands and Alaska to Guatemala in the east. Pilot whales belonging to two or three other subspecies of this species are sometimes listed as inhabitants of the Southern hemisphere (Tomilin, 1962; Nishiwaki, 1966). (V.A.) Biology Population. The pilot whale can be reckoned as a relatively abundant species. In some regions of the range (North Atlantic, waters of Japan, and South Pacific Ocean), this dolphin often forms herds a thousand or more strong. It is difficult to indicate the regions of maximum population of the pilot whale. Food. Squids serve as the main food. In the waters of Newfound- land, the pilot whale consumes the squid Illex illicebrosus almost exclu- sively. This squid is remarkably abundant here. In the summer months, pilot whales approach the island coasts chasing after it. In the absence of squids, pilot whales consume cod (in small quantities). In a Florida oceanarium, a young pilot whale initially refused to eat fish but after sometime was taught to accept it. In the waters in which I. illicebrosus is not available, the stomach of pilot whales contained: the remnants of other species of squids, e.g., Ommatostrephes (Todarodes) sagittatus, and some species of the family Onychoteuthidae. Otoliths of mackerels (Caranx trachurus), small flounders (Pleuronectidae), and herrings have been found from time to time. Young pilot whales feed on much smaller squids than the adults do. The nearly identical degree of digestion of the food in the stomach of pilot whales caught at the same time suggests that the entire herd feeds together. By converting the quantum of the remnants of squids and fishes into the average weight of intake, it was established that the stomach of a pilot whale of medium proportions (length 396 cm) held 12-14 kg 14 The following distribution has been given for this form: “Atlantic, Pacific, and Indian oceans from 70°N lat. to roughly 60°S lat.” (Hershkovitz, 1966). 709 of food. The stomach of a female 247 cm long contained 155 squids weighing in toto 27 kg. In an oceanarium, a male pilot whale 525 cm in length daily consumed an average of 45 kg of squids and mackerel, while a female 396 cm long consumed up to 36 kg; another female (366 cm long) ate 18 kg three times a day. Apparently, under natural conditions, the pilot whale may fill its stomach two or three times a day. Such voracious feeding is only resorted to, however, during the three or four summer months when immense concentrations of squids are available. During the rest of the year the daily ration of the pilot whale is quite modest. At a daily consumption of 27 kg, the annual food requirements are about 10 tons. Taking the weight of a pilot whale of medium proportions (396 cm long) as 830 kg, its annual food intake represents 11.5 times the weight of the animal itself. The daily quantum of food consumed by the dolphin constitutes 3 to 5% of the animal’s body weight. Observations on three pilot whales in a basin showed that they fed only at night and rested during the day. Under natural conditions in Newfoundland, pilot whales were observed feeding during the day also. The stomach ofa pilot whale caught between 7:00 and 11:00 a.m. revealed food recently consumed. It is probable that the long summer day in these latitudes causes the squids to rise to the upper horizon not only at night, but also during the day, which enables successful catches by the pilot whales (Sergeant and Fisher, 1957; Tomilin, 1957). Behavior. Pilot whales usually live in herds of some 20 animals, on average, but sometimes do form herds of up to a thousand or even several thousands of animals. They dive for comparatively longer periods and surface sometimes 8-10 times in a row, producing low (up to 1.5 m high) bushy blows. They never display the caudal flukes and never leap free of the water surface. Large herds of pilot whales are formed from individual groups of 15-25 animals each. The composition of the herds varies. Thus a herd of 16 pilot whales comprised 7 males and 8 females (the sex of one whale could not be ascertained). Of the seven males, three were mature, three immature, and one a newborn calf. The eight females comprised six mature, one on the threshold of maturity, and one newborn. This was a mixed herd. The predominance of mature females over mature males suggests polygamy. The ratio between the adult males (472 cm and above) and adult females (336 cm and above) was close to 1:3. | Another herd studied consisted of 14 pilot whales which comprised 10 males and 4 females. The length of nine males was typical of mature animals and all the four females were large, presumably “old”. Such a herd could be called a “bachelor” group, made up in all probability of S29 710 animals which did not take part in reproduction in that particular year (Sergeant, 1962b). Pilot whales often form mixed herds together with some species of very small dolphins. In such herds, apart from the distinct presence of the shorthead dolphin [Lagenorhynchus electra] and the bottlenose dol- phin, other species of dolphin may also be present. When such a herd approaches a ship standing in the sea, all the small dolphins disappear into the water after some time while the pilot whales invariably continue to remain close to the ship (Brown, 1961). Sometimes some animals surface vertically from the water almost up to the flippers, remain in that position for about 0.5 min, then submerge without altering their upright posture. Some animals strike the water with their fins (Brown, 1961). This ability to assume a vertical position is taken advantage of in oceanariums. The pilot whale is gradually trained to rise higher and higher out of the water to reach food dangled above it. A large dolphin, such as the pilot whale, can thus be trained to leap clear of the water and even to “Пу” above it (Fig. 305). The extraordinary behavior of a herd of pilot whales once came to notice. This mixed herd consisted of about 150 pilot whales and small dolphins. Suddenly, some animals assumed a vertical position but with their head down (under the water) and each struck the water surface five or six times with its caudal flukes. They then assumed a horizontal position and rejoined the herd. Such a head-down position has been reported among pilot whales in other instances too (Brown, 1960). Pilot whales are beached more often than other cetaceans, both indi- vidually as well as in groups. On October 7, 1948, as many as 46 pilot whales were cast ashore simultaneously on a beach in Florida; those found early in the morning, during low tide, were still alive. Four were transported by truck while the others perished on the coast in high tide (Obruchev, 1951). Beached groups usually consist of animals of various age but sometimes young ones exclusively. A pilot whale in a Florida aquarium slept only at night in the first few months of captivity but later continued to sleep during the day also. While asleep, all of its body was submerged and only the dorsal fin and blowhole remained exposed above the water. Pilot whales hear well under water and their aeriai vision is excel- lent. In captivity, they invariably perceive the approach of the food atten- dant and swim immediately to the feeding place. One female pilot whale quickly learned to snatch food from the attendant’s hand; it swam to the feeding site, stood vertical with its head above the water, and opened its mouth for the attendant to throw cephalopods inside. Washing of the 529 530 711 SAN See a ‘Wat ills \ soot es ie 3 И tS nl Ay ae vl ‘ath , eS г. т Tae Fig. 305. Leaping pilot whale (figure by N.N. Kondakov). pail signalled completion of feeding and the pilot whale would immedi- ately swim away. A male loved to be combed with a brush and quickly approached anyone with a brush in his hand (Tomilin, 1957). A pilot whale (body length 366 cm) caught on February 27, 1957 was housed in a tank 30.5 m long, 15 m wide, and 6.7 m deep. For the first few days the animal refused food and the attendant had to enter the tank in a diver’s suit, open her mouth with metal tongs, and push squids inside which the whale gulped rapidly. After 10 days she began to swim to the wooden platform affixed to the side of the tank and to take Ve food from the tongs held by the attendant. The animal consumed more - than 20 kg of squids per day. A large fish and two turtles were also kept in the same tank. During the feeding period sometimes the fish would snatch the squids from the tongs before the pilot whale and sometimes even snatched them from the whale’s open mouth. This angered the pilot whale. She would stop feeding and chase the fish around the tank. The turtles also swam to the feeding platform, which irritated the whale. Once she threw herself on them with such vehemence that one suffered a broken shell. However, the pilot whale soon learned to avoid competition during feeding: she would stand vertically at the feeding platform with a third of her trunk projecting above the water, maintaining balance by slight movements of the caudal ‘flukes, and thus receive the food high above the water surface. This offered a basis for teaching the pilot whale various tricks. First, it was taught to leap clear of the water to reach food dangled high above its head and next to associate feeding with a whistle. This was followed by a “hand shake”. As soon as the pilot whale appeared around the feed- ing platform, the trainer would clasp it around the flippers and whistle before feeding. Very soon the pilot whale began to anticipate the actions of the trainer. On hearing the whistle, it would turn on its side, “extend” the flippers to the trainer, then demand food. By the same method, it was taught to fetch a plastic dumbbell from inside the tank and eventu- ally to “sing”. At the command of the trainer, it would quiver its jaws and simultaneously blow air powerfully through the blowhole. The pilot whale learned these various tricks quite rapidly. It was not at all afraid of any unfamiliar object. When in the course of the experiments a large boat was dropped inside the tank, the whale exhibited no apprehension. To the contrary, after a few minutes it swam to the boat, fed tranquilly, and performed its tricks. | In the first week of February, 1958, the pilot whale lost its appetite. Next, it began to rub the genital slit against a stone or other projecting object at the bottom of the basin, including the diver’s abandoned helmet. This behavior continued for nearly two weeks, after which the animal resumed normal feeding. Perhaps this was the period of its heat. For a long time the pilot whale was friendly with the divers who entered the tank to feed the other animals, even attempting sometimes to snatch the food from their hands. But after nearly 14 months of living in the tank, it began to push the diver (or anyone entering the tank) with its head and to bite his hands while he was feeding the other animals. One diver barely escaped serious injury when, on the water surface, the pilot whale tried to pin him to the wall of the tank. Later, it chased three persons simultaneously, two of whom were photographers. It hit the leg 531 713 of one and zoomed to the second loudly clicking its teeth; all three made a hasty retreat. It then attacked the large photographic camera set up on a tripod at the bottom of the tank, hitting it with such force that the heavy lens was badly damaged. Four days later the pilot whale struck one of the divers so powerfully that he lost consciousness and barely escaped drowning. This change in the behavior of the pilot whale was attributed to its long lonely existence. In July, 1958, it was transferred to a basin with another small pilot whale and two blue-white dolphins. Initially the dolphins were frightened and shied away from the large pilot whale, but soon became emboldened and spent several hours nipping it behind ihe flippers whenever possible. This irritated the pilot whale who chased them but the more agile dolphins easily escaped. After about a month the animals became friends, swimming abreast, and often playing together. From the time the pilot whale began living with the other dolphins, its animosity against the divers ceased (Brown, 1960). Migrations. Pilot whales probably perform seasonal migrations since they are more abundant in the North Atlantic Ocean (as in the Pacific Ocean also) in summer than in winter. Norwegian whalers hunting for small whales in the northwestern Atlantic catch the pilot whales mostly in the second half of summer. Thus in 1958, of the 216 pilot whales caught, 64 were bagged in July and 150 in August (Ostby, 1959). Table 41 provides an idea of the duration of residence of pilot whales in the waters of Newfoundland. In the first half of January, 1953, a large herd of pilot whales was sighted at 42°30’ М lat. and 52° W long.; evidently the effect of the Gulf Stream was more intense in the winter. In the summer months, pilot whales Е visit the large deep- water bays of Newfoundland and, concomitantly, though in small num- bers, are found in the central part of the Labrador Sea. Similarly, pilot whales are regularly sighted in summer in the eastern waters of the North Atlantic, initially off the Faroe Islands and the southwestern coast of Norway, later off northwestern Norway, Iceland, and Jan Mayen Island, and at the end of summer, in the western part of the Barents Sea, in the waters of Medvezhii Island, and even at Spitsbergen. Table 41. Observations on pilot whales in Newfoundland (Sergeant and Fisher, 1957) Year First observation Subsequent observation 1953 July 8 November 4 1954 July 6 November 10 1955 July 15 October 22 532 714 The pilot whale is purportedly encountered in winter east of the Grand Bank south of Newfoundland in the waters of the North Atlantic current. Information on the migrations of the pilot whale in other regions of its range is so scant that no clear picture of migrations can be drawn. Apparently migrations are not very distinctly manifest since these ani- mals are sighted in the northern as well as the southern part of the range throughout the year. From 1926 through 1953 (July to October end), 20 instances of beached pilot whales were recorded on the coast of western Green- land, most of them in September. In the winter of 1931/32, a herd of pilot whales wintered on the coasts of western Greenland (Sergeant and Fisher, 1957). Reproduction, growth, and development. The periods of mating and parturition in the pilot whale evidently last for about six months but most of the animals mate over a brief duration. The maximum number of matings presumably occurs in the spring months with parturition peaking in mid-August, although there have been instances of calves born from May through November. Stray cases of births have been recorded in almost all the months of the year. Gestation apparently continues for 15-15.5 months. As a rule, a single calf is born and twins are rare. There is a solitary record of a female with three embryos. The average length of male calves at birth is 178 cm, of female calves 174 cm. The teeth begin to cut at a body length of about 213 cm. This process ceases when the calf attains a length of about 274 cm. The growth of embryos and newborn pilot whales is shown in Table 42. The calves begin independent feeding on squids at an average body length of 230 cm at the age of six- nine months; however, suckling does not cease and extends for about two years. Lactation of the female contin- ues for 21-22 months. Instances of fresh impregnation during lactation are rare; the females usually ovulate at the end of lactation. Thus the full cycle of reproduction roughly covers 40 months. During the period of reproductive capability, the female produces, on average, nine calves. Male calves record a much greater growth tempo than female calves. A young male living in a Florida aquarium added 30 cm to its length and put on 45 to 90 kg in four months. The average length of adult animals is usually about 396 cm. Females ovulate for the first time (attain sexual maturity) at six years of age at an average body length of 356 cm and males at the age of 12 years at an average body length of 490 cm. After attaining sexual maturity, the growth of the animal slows down. Males, which attain maturity later, are considerably larger than females. Most of the males in the group measured, fell in the 565-cm group and the maximum number of females in the 340-cm group. The maximum size 533 5 Table 42. Length and weight of embryos and calves of pilot whales (Sergeant, 1962b) Embryos Calves Length, cm Weight, kg Length, cm Weight, kg 16.5 0.08 193 113.4 19.0 0.12 195 953 21.0 0.18 205 145.6 23.0 0.22 208 130.6 24.0 0.25 231 188.7 25.0 0.32 236 202.8 26.0 0.36 31.0 0.54 36.0 0.75 41.0 1.16 43.0 1.45 48.0 1.54 59.0 3.74 114.0 21.00 119.0 28.50 165.0 60.00 of the males was 617 cm and of the females 511 ст. The maximum recorded age of the males was roughly 40 years and of the females 50 years (Tomilin, 1957; Sergeant, 1959, 1962b). Table 43 shows the composition of pilot whales caught in Newfound- land according to their sizes: the animals have been grouped into a series at 15-cm intervals. The total number of males measured was 1,275, of females 1,951 and the number falling in each length group is given. The data show that the size of adult males far exceeds that of adult females and that animals of all ages fall victim to the hunter. Table 44 gives data on the age composition of the oldest groups of pilot whales caught and investigated. The age of the animals was determined from the tooth layers. From a sample of 518 pilot whales, males with a body length 549 cm and above (only 11 animals) and females with a body length 457 cm and above (22 animals) were taken. It can be seen from this Table that these animals were quite advanced in age, the females reaching a maximum age of 50 years. Enemies, diseases, parasites, mortality, and competitors. As in the case of all other cetaceans, the killer whale is among the enemies of the pilot whale. Bone tumors were detected among the diseases suffered by the pilot whale. The skin parasite Isocyamus delphini Guerin-Meneville was invari- ably detected around the mouth and in old wounds on the body of the Newfoundland pilot whale. Conchoderma auritum Linn. was detected on 532 533 716 Table 43. Size composition of pilot whales studied (Sergeant, 19625) Average length Number of animals Average length Number of animals of animals in in the group of animals in in the group the group, cm LANG ERED OM SEROMA TONE) ole a the group, cm в hy se Males Females Males Females 168 6 10 396 59 169 183 15 22 411 41 259 198 12 17 427 61 278 213 20 16 442 44 221 229 13 12 457 36 170 244 28 32 472 36 50 259 51 48 488 37 10 274 54 61 503 37 5 290 63 66 518 41 — 305 52 78 533 67 — 320 61 83 549 76 — 335 59 83 564 84 — 351 53 59 579 47 — 366 60 73 594 23 — 381 33 129 610 6 — Table 44. Age of the largest pilot whales from a sample of 518 animals (Sergeant, 1962b) Age, years Number of animals Males Females 21-25 4 2 26-30 2 6 31-35 3 8 36-40 2 3 41-45 = 2 46-50 — 1 Total 11 22 the gums and teeth, and Cyanus globicipitis Lutken, Xenobalanus glo- bicipitis Steenstrup, and Cyrolama globicipitis van Beneden on the body. Eight species of endoparasites have been recorded: trematodes one, ‘cestodes three, nematodes three, and acanthocephalans one. The trema- tode Campula gondo Yamaguti was found only in the bile ducts of the pilot whale from the Pacific Ocean. The cestode Trigonocotyle lintoni Guiart, parasitizing the intestine, is likewise known only in pilot whales from the Atlantic Ocean and the Mediteranean Sea. Phyllobothrium del- phini Bosc, widely distributed among marine mammals, parasitizes the skin. In addition to the pilot whale, it has been detected in six other 534 И 17 species of toothed whales, in right whales, and in Weddell’s seal from the Atlantic and Pacific oceans, the Mediterranean Sea, and Antarctic waters. The cestode Monorygma grimaldi Monier is a parasite in the abdominal cavity, mesentery, and diaphragm of pilot whales and three other species of dolphins of the Atlantic Ocean and the Mediterranean Sea. The nematode Anisakis (Anisakis) typica Diesing localizes in the stomach of the pilot whale and three other species of dolphins; it has been found in the North Sea and on the coasts of southeastern Africa. The nematode Stenurus globicephale Baylis and Dauney, parasitizing in the blowhole, bronchi, and the circulatory system, has been reported only in the pilot whale from the North Atlantic. The nematode Torynurus con- volutus Kuhn parasitizes the bronchi and blood vessels of the lungs of the pilot whale and also common porpoises and one more dolphin (species not established). It has been detected in the waters of Europe, the North Atlantic, Sakhalin, and the Sea of Okhotsk. The only species of acan- thocephalans, Bolbosoma capitatum Linstow, localizes in the intestine of the pilot whale, sperm whales, and false killer whales from the Atlantic Ocean and the Mediterranean Sea (Delamure, 1955). Field characteristics. This species differs from other dolphins in hav- ing a round head which projects high above the water, a dorsal fin with a very broad base and its apex turned posteriorly, and slower movements. (V.A.) Economic Importance The pilot whale is hunted in Newfoundland and the Faroe Islands by Norwegian ships operating close to Norway as well as in the North and Barents seas, in addition to other species of small cetaceans; it is also hunted in the waters of Japan (Table 45). A relatively large number of pilot whales is caught only in Newfound- land and in the Faroe Islands; these animals are of considerable local importance. In all other waters, their catch is too small to be significant. Hunting is done by various methods. In Norway and Japan, in addi- tion to other species of small whales (Minke whale, killer whale, beaked dolphins, etc.), they are shot with small-bore harpoon guns from small boats. In Newfoundland, most of the pilot whales are caught by chasing large herds into shallow bays of islands and harpoons used to a lesser extent. The flesh of the animals caught is used as food (in Japan) and also as feed for fur-bearing animals in farms (Newfoundland). Oil is obtained from blubber. (V.A.) 718 Table 45. World catch of pilot whales (international whaling statistics) Year Newfoundland Faroe Norway Japan Islands 1950 172 10 1951 3,100 8 1952 3,155 2 1953 3,584 1 1954 2,298 1955 6,612 13 61 1956 9,799 1 279 1957 7,797 80 174 1958 789 216° 197 1959 1,725 1,422 224 144 1960 1,957 1,680 331 168 1961 6,262 1,892 295 133 1962 150 1,753 43 80 1963 221 2,194 71 228 1964 2,849 1,386 54 217 1965 1,520 1,599 32 288 1966 887 1,488 339 199 1967 739 1,979 117 237 1968 204 1,749 31 166 Genus of Pygmy Killer Whales Genus Feresa Gray, 1870 1870. Feresa. Gray. Proc. Zool. Soc. London, p. 77. Orca intermedia Gray = Feresa attenuata Gray. (V.H.) Small dolphins, with a body length up to 244 cm. The head is rounded and has no “beak”. The dorsal fin is quite high. The main body color is dark gray. The rostrum occupies nearly one-half of the skull length. The ante- rior part of the premaxillae is flattened; their, inner edges are set off from each other. The lower jaw has a keel in the zone of the symphysis. Teeth оц. Vertebrae 68-71. Biology has not been studied. These dolphins apparently feed on squids and fish. There is very little information on the sites where this rare dolphin is found: in the North Pacific Ocean from Alaska to Guatemala in the east, on the coasts of Japan, in the Yangtze estuary, and on the coasts of China in the west. This dolphin has not been reported in the USSR but it might occur in the waters of Kuril Islands and in the Sea of Japan. (V.S.) 535 719 PYGMY KILLER WHALE Feresa attenuata Gray, 1875 1827. Delphinus intermedius. Gray. Philos. Mag. or Annals, 2, p. 376. Type locality not established. Nom. ргаеосс. 1875. Feresa attenuata. Gray. Journ. Mus. Godeffroy (Hamburg), 8, p. 184. 1856. Feresa occuleta. Jones and Packard. Proc. Biol. Soc., Washington, 69, p. 167. Substituted for Delphinus intermedius Gray. (V.H.) Only species of the genus. Somewhat resembles the false killer whale. The body is elongated and spindle-shaped. The head is relatively small. The highly developed adipose pad [melon] projects anteriorly to impart a circular feature to the head (Fig. 306). The flippers are crescent-shaped, quite broad at the base and pointed at the tips. The height of the dorsal fin is slightly less than its basal length. It is arcuately notched along the posterior margin. The caudal flukes are relatively small. The body is a monochromatic dark gray. The margins of the jaws and the region around the anal opening are white. An anchor-shaped light-colored patch is seen on the breast. Wavy pale-colored bands run on the flanks (Nishiwaki, 1966). The skull (Fig. 307) is highly shortened and measures about 16% of the body length. The short rostrum, broad at the base, gradually, but not very significantly, narrows toward the tip. The upper side of the rostrum is Slightly concave (Yamada, 1954). The teeth are fairly large with crowns about 1 cm high. Cervical vertebrae 7, thoracic 12-13, lumbar 16-17, and caudal 32 - 34. Fig. 306. Pygmy killer whale, Feresa attenuata (figure by N.N. Kondakov). 536 720 SY NINES Fig. 307. Skull of the Pygmy killer whale, Feresa attenuata (figure by N.N. Kondakov). The body length of males (Japan; Nishiwaki, 1966) ranges from 214-244 cm, of females 208-227 cm. The maximum weight recorded was 225 kg for a male measuring 244 cm in length. The body propor- tions of males with a body length of 214-244 cm and of females with a body length of 208-227 cm respectively (as percentage of body length) are: from tip of snout to blowhole 6.8-11.9 and 7.2-11.1, to flippers 16.7 - 21.6 and 18.5 - 22.4; from notch between caudal flukes to anal open- ing 34.2 -37.2 and 34.2 - 38.6; length of flippers 18.3 -22.1 and 14.7 - 22.2, maximum width of flippers 6.1-6.7 and 5.8-7.0; length of base of dorsal fin 12.6- 15.4 and 14.0- 16.6, height of dorsal fin 9.4- 11.6 and 9.6 - 10.9; and spread of caudal flukes from apex to apex 23.8- 27.3 and 23.0- 28.4. The measurements of 14 skulls (Japan; Nishiwaki, 1966) with a condylobasal length of 356 to 390 mm (as percentage of this length) are: length of rostrum 44.7-49.2, basal width of rostrum 27.7-31.8, interorbital width 53.7 - 60.7, length of lower jaw 72.2 - 79.9, and length of mandibular symphysis 8.7 - 10.1. Information on the geographic distribution of this species is very scant, with only a few recorded finds and observations (Fig. 308). It has been noticed in the waters of Senegal and in the North Atlantic. It is known in the Yangtze estuary, along the coasts of Honshu Island, and in the North Pacific Ocean. (V.S.) Biology has not been studied. 721 Fig. 308. Pygmy killer whale, Feresa attenuata (figure by М.М. Kondakov). A—coloration of underside; B—of upper side. Genus of Common Porpoises Genus Phocoena G. Cuvier, 1817 1817. Phocoena. G. Cuvier. Rgne Animal, ed. I, I, p. 279. Delphinus phocoena Linnaeus, 1758. 1828. Phocoena. Gray. Spicil. Zoologica, I, р. 2. Delphinus phocoena Lin- naeus, 1758. (V.H.) Small dolphins, with a maximum body length of 2 m. These dolphins have no “beak”. The head is obtuse and the forehead flattened. The anterior margin of the low dorsal fin, short flippers, and caudal flukes bears small horny tubercles (sometimes these are present only on the dorsal fin). The body is dorsally dark, gradually turning white on the ventral surface. The rostrum of the skull is equal to or slightly shorter than the cranium. The posterior section of the premaxillae anterior to the bony nares forms a prominence. The pterygoid bones are separated by a gap. The boundaries of the interparietal bones are invariably very prominent. Teeth 16-39. They have a slightly flattened crown and are set off from the root by a neck. Vertebrae 62-66. Ribs 12-14 pairs. The sternum is non-segmented. Phalangeal formula: I, _ 3, II; - 49, III; - 3, [У›-в›апа У -3. These dolphins are ichthyophagous, surviving mainly on benthic fish (bentho-ichthyophagous). The females whelp every year. Gestation extends for nine to ten months and lactation for about four months. 538 722 Females are ready for fertilization two to three months after parturition, while still nursing a calf. Common porpoises are distributed in the North Atlantic from the Barents Sea and Davis Strait to New Jersey in the west and the Mediter- ranean, Black, and Azov seas, and Senegal waters; in the South Atlantic from Rio de la Plata (35° S lat.) to Cape Horn and South Georgia; in the North Pacific Ocean, from the Chukchi Sea to Mexico in the east and Japan in the west; in the South Pacific Ocean, from Paita in Peru (5° S lat.) to Cape Horn (Hershkovitz, 1966) (Fig. 309). The members of this genus are thus absent in the Indian Ocean, in the western part of the South Pacific Ocean, and in some tropical and equatorial parts of the Atlantic and Pacific oceans. The genus comprises four species: 1) P. phocoena Linnaeus, 1758; 2) P. dioptrica Lahille, 1893; 3) P. spinnipinnis Burmeister, 1865; and 4) P. sinus Norris and McFarlan, 1958. Sometimes the genus is divided into a larger number of species. Only one species is encountered in the USSR waters: the common porpoise, P. phocoena Linnaeus, 1758. The common porpoises are presently not of commercial importance. (V.S.) COMMON PORPOISE} Phocoena phocoena Linnaeus, 1758 1758. Delphinus phocoena. Linnaeus. Syst. Nat., I, p. 77. Baltic Sea (“Oceano Europeo Balthico’’) 1827. Phocoena communis. Lesson. Man. Mammal, p. 413. Renamed as Delphinus phocoena auct. 1865. Phocoena vomerina. Gill. Proc. Ac. Nat. Sc. Philadelphia, 17, p. 178. Puget Sound, Washington State. 1905. Phocoena relicta. Abel. Jahrb. K.K. Geol. Reicheanstate Wien, 55, p. 387. Crimean waters of the Black Sea. (V.H.) Diagnosis Only species of the genus encountered in the USSR. Description The external appearance of the common porpoise is typical: the body build is much heavier than that of dolphins, somewhat stunted, and the 15 Also locally known as “Azovka,” “Pykhtun,” “Svinka” [porpoise], etc. (Black Sea). | | Hl 1f i | th | 1000 © 1000 2000 3000 4000 5000 km Fig. 309. Range of the genus of common porpoises, Phocoena (V.A. Arsen’ev). 537 724 7 fe L SS MODEL Wi LLL fi \ te И i i o “on af) 539 Fig. 310. Common porpoise, Phocoena phocoena (figure by N.N. Kondakov). trunk thickset. The head is short. The flippers are oval and the dorsal fins triangular (Fig. 310). Horny tubercles 12 - 16, less frequently 18-20, occur along the ante- rior margin of the dorsal fin in common porpoises (adult animals and embryos) of the Black and Azov seas (Tsalkin, 1938b). It has been sug- gested that these tubercles represent derivatives of the thick skin cover (Kukenthal, 1889-1893). Two or three hairs are seen on each side of the snout in embryos. The color of the dorsal surface varies from a dark gray to almost black. The light-colored abdomen is sometimes sharply demarcated from the dark dorsum but sometimes the transition in col- oration is altogether imperceptible (Tsalkin, 1938b). A faintly percepti- ble dark gray band runs from the zone of the ear openings toward the navel on each side of the body, while another almost black band extends from the corner of the mouth to the base of each flipper (Barabash- Nikiforov, 1940). Instances of partial or total albinism are encountered occasionally. The form of the tooth crown is variable—from almost conical to a form with three faintly discernible cusps. The anterior surface of the crown is slightly turned inside the cavity. When the mouth is closed, the teeth on the lower jaw fall inward of the upper teeth. Among the Black and Azov sea common porpoises, the number of teeth in the upper jaw varies from 44 to 60 (more often, 54) and in the lower jaw 38 to 55 (more 539 often, 44-46) (Tsalkin, 1938b). The teeth of this common porpoise are characterized by curvature, distortion of the roots, and thickening in the lower portions (Tsalkin, 1938; Barabash-Nikiforov, 1940). As age advances, the teeth may wear down or even drop out. Cervical vertebrae 7, thoracic 12-14, lumbar 14-17, and caudal 27 -32. The sections of the vertebral column comprise (as percentage of its length): cervical 1.9-3, thoracic 22.1 -24.6, lumbar 30 - 34.2, and caudal 40-44 (Tomilin, 1957). Six to eight pairs of ribs are articulated with the sternum in the embryos and usually five pairs in adults. 725 Females are slightly larger than males. The difference in body length between mature females and males from the Black and Azov seas averages 7 cm (Tsalkin, 1938). The main body measurements of this species of common porpoises (eight animals) from the Black Sea average (Tsalkin, 1938) (in cm): body length 163; distance from tip of snout to base of flippers 32, to anterior margin of dorsal fin 73; and length of flippers 22. The length of the largest common porpoises from the Black and Azov seas is (in cm): male 167 and female 180; from the North Pacific Ocean: female 178.5; and from the North Atlantic: male 186. The average main skull measurements (measurements of five males and three females; Tomilin, 1957) (Fig. 311) are respectively (in cm): condylobasal length 25 and 27, length of rostrum 11 and 12, width of rostrum at base 7 and 8, and length of lower jaw 19 and 21. (V.S.) Geographic Distribution - Temperate and partly cold waters of the North Atlantic and the North Pacific oceans; predominantly the coastal zone. Geographic Range in the USSR In the basin of the Atlantic Ocean, this porpoise is commonly encountered in the Barents Sea where it reaches the coasts of Novaya Zemlya and Yugorsk Shar and is abundant on the Murman coast; it is also common in the White Sea (waters of Kanin, Letn’aya and Tersk 539 Fig. 311. Skull of the common porpoise, Phocoena phocoena (figure by М.М. Kondakov). 542 726 coasts, and Gulf of Kandalaksh), Kara Sea (transgresses into its western section), Baltic Sea, and almost and the entire coastal water body of the Black and Azov seas (Fig. 312). In our waters of the Pacific Basin, it is common in the Sea of Japan and the Sea of Okhotsk, on both sides of the Kuril range, along the coasts of Kamchatka, Koryatsk land, and the Gulf of Anadyr, around the Commander Islands in the Bering Sea, in the Bering Strait, and in the Chukchi Sea where it penetrates north of 70° N lat. (east of Point Barrow, 71°24’ N lat.). Its range in the west is not known. Geographic Range outside the USSR In the western part of the North Atlantic, the common porpoise is known from New Jersey (39° N lat.), and possibly more southward, up to Davis Strait, Baffin Bay, and coasts of Greenland in the eastern part of the Atlantic Ocean, from the waters of Senegal and Dakar in northern Africa (about 15° N lat.) to the Barents Sea, Iceland, and east coast of Green- land (Scoresby Sound), i.e., roughly up to 70° N lat. The range covers the entire Baltic Sea. Transgressions into the Mediterranean Sea from the Black Sea as well as from the Atlantic Ocean are known (Fig. 313). In the Pacific Ocean, this species lives along the eastern coasts of the Korean peninsula and in the waters of Japan. It inhabits the entire coastal waters of the American continent down to California and Mexico from the Chukchi Sea (Point Barrow) and the Bering Sea, including the coastal waters of the Aleutian Islands (Kleinenberg, 1956a; Tomilin, 1957; Hall and Kelson, 1959; Nishiwaki, 1966). (V.A.) Geographic Variation In the Northern hemisphere, the species forms three subspecies that are almost indistinguishable although geographically disjointed. All the three subspecies are encountered in the waters of the USSR. 1. Northern Atlantic common porpoise, P. p. phocoena L., 1758 (syn. communis). This is the largest of the subspecies with a body length up to 186 cm, condylobasal length of skull 255-259 mm, width of condyle 65-71 mm, and height of occipital condyle 38-42 mm. The rostrum is hardly shortened. Tubercles are seen on the anterior margins of the flippers, dorsal fin, and caudal flukes. This subspecies is encountered in the Barents, White, Kara, and Baltic seas. Outside the USSR, it has been reported in the waters of the North - Atlantic Ocean. 727 ‘(лэ.иэзту у) USSN э41 Ul 9 и2020ц4 nusos0yq ‘asiodiod иоититоо ay) Jo aduey ‘21$ ‘89 1 OL ov у : №! ic wh se К 1 АИК f =e $ ©. | ' г. A R 2 , ый С 7! a = 5 ЧвеЧ3еЯ ay $) E Г $ ‘ ay а Ч Ч 728 = 3 АНИ i Zar ul И \ | АИ an Wea ah Diet tt male ИКИ tit tin | Г ЧОН ИИ о ОО: HPO © Я Ин ИО И к - ORO а ites ae 72 НОТ ee | т : AU НИ Ва 120 Я Ги oo И i! iy 1 р ИАИИИ НО his | о y Wa к Г хи Nate a UH i a yh 150 150 1000 2000 3000 4000 £000 km 180 1000 0 [ten ЧИ КНИГИ Fig. 313. Species range of the common porpoise, Phocoena phocoena (V.A. Arsen’ev). 541 543 729 2. Black Sea common porpoise, Р. р. relicta Abel, 1905. This form is of medium dimensions, with a body length up to 180 cm, condylobasal length of skull in males 245-255 mm, in females 269-274 mm; width of condyle in males 54-62 mm, in females 58-63 mm; height of occipital condyle in males 32 -36 mm, in females 37-39 mm. The tubercles on the anterior margin of the fins are totally reduced. This subspecies is encountered in the Black and Azov seas. Outside the USSR, transgressions have been reported for the Marmora and Mediterranean seas. 3. Pacific common porpoise, P. p. vomerina Gill, 1865. This is the smallest of the subspecies with a body length up to 178 cm, condylobasal length of skull 258-293 mm, width of condyle 68-70 mm, and height of occipital condyle 40-43 mm. The rostrum is highly elongated. This subspecies is encountered in the Chukchi, Bering, Okhotsk, and Japan seas, and in the waters of Kuril Islands from the Pacific Ocean side. Outside the USSR, it has been reported in the American waters of the Pacific Ocean from the Chukchi Sea to the Mexican coasts and in the waters of Japan. (V.A.) Biology Population. The species can be regarded as relatively abundant. The least populous of the three subspecies is the Azov-Black Sea common por- poise, which is confined to an extremely restricted range. Populations of the other two subspecies occupy extensive water bodies in which they are commonly encountered and are perhaps quite abundant. Food. Information on the food of the Atlantic and Pacific popula- tions is extremely scant while that on the food of the Black Sea popula- tion is more widely available (Table 46). Two species of pelagic fish, viz., smelt (49.5%) and Black Sea anchovy (18.5%), occupy the primary position in terms of the quantum of fish found in the stomach of the Azov-Black Sea common porpoise. All the species of goby (bottom-dwelling fish) comprise only 30.9%. The rest of the species of fish in the food of the common porpoise constitute only a few tenths of one per cent. In terms of the number of fish objects encoun- tered, pelagic fish comprise 68.8% and benthic fish 31.2%. However, in terms of weight (as percentage of total weight of food), rotan goby occu- pies the first position (32.4%), followed by round goby (31.2%), smelt (14.5%), and Black Sea anchovy (11.0%). Thus, the group of benthic fish constitutes 67.9% of the total weight of food intake and the group of pelagic fish 32.1% 730 Table 46. Food objects of the common porpoise Azov-Black Sea (Tsalkin, 1940) Fish Round goby, Gobius melanostomus Rotan goby, G. rotan Mushroom goby, G. cephalarges Syrman goby, G. syrman Toad goby, Mesogobius batrachocephalus Black Sea flounder, Pleuronectes flesus luscus Black Sea sole, Solea nasuta Black Sea anchovy, Engraulis encrasicholus Black Sea silverside-smelt, Atherina pontica Perch, Lucioperca lucioperca Bream, Abramis brama Golden gray mullet, Mugil auratus Black Sea whiting (haddock), Gadus euxinus Black Sea shad (herring), Caspialosa sp. Crustaceans Balanus improvisus Brachinotus lucassi Idothea baltica Leander sp. Mollusks Algae Ulva lactuca North Atlantic (Tomilin, 1957, 1962) Fish Cod, Gadus morhua Capelin, Mallotus villosus Navaga, Eleginus navaga Sand eel, Ammodytes hexapterus Whiting, Odontogadus merlangus Common mackerel, _ Scomber scombrus Pollock, Pollachius virens Herring, Clupea harengus Sardine, Sardina sp. Sole, Solea sp. Eel, Anguilla anguilla Salmon (small) Sprat, Sprattus [= clupea} sprattus Trout Crustaceans Decapods Mollusks Cephalopods (Loligo pealitii) Algae North Pacific (Sleptsov, 1955; Tomilin, 1957) Fish Pacific cod, Gadus morhua macrocephalus Capelin, Mallotus villosus Pacific navaga (saffron cod), Eleginus gracilis Polar cod, Boreogadus saida Herring, Clupea harengus Shad, Alosa sapidissima Shark, Squalus sp. (?) Sablefish (black cod), Anoplopoma fimbria Whitefish (Leucichtys) Crustaceans Shrimps Mollusks Cephalopods Most mollusks probably enter the stomach of porpoises through the fish they consume. Two species of crustaceans (Idothea baltica and Leander sp.) can be regarded as incidental food objects as they were found in only two stomachs. Alga was found in the three groups of porpoise in large quantities and probably serves as food. Thus, fish represent the almost lone source of food to the Azov-Black Sea common porpoise; most of them are benthic and a small percentage 544 731 pelagic. The latter serve as the main food only in the period of massive migrations (Black Sea anchovy) when they form concentrations of large proportions. Seasonal variations in food types have been noticed: benthic fish predominate in winter, while in summer Black Sea anchovy and smelt play an extremely important role in the food of common porpoises during the period of migrations of these fish from the Black Sea into the Azov Sea and back in spring and autumn (Tsalkin, 1940). The North Atlantic porpoises feed mainly on fish (benthic as well as pelagic forms). Cephalopods and other mollusks, crustaceans, and algae play no significant role in their food. The nature of the food of the Pacific porpoises is very similar to that of the other populations. Daily activity and behavior. The porpoises of the Azov-Black Sea pop- ulations (probably the others too) avoid the open seas and are encoun- tered more in the coastal shallow waters (Fig. 314). They are usually confined to small groups of not more than 10 animals. Relatively larger groups are encountered only in the period of massive migrations of fish which serve as their food. Exhalation and inhalation occur at not more than 30-sec intervals although instances of porpoises remaining submerged for 6 min are known. Their movement is comparatively slow. Intense flexure of the body while diving is a characteristic feature. The porpoises of this species, with rare exception, do not trail behind moving ships. Fig. 314. Common porpoises at sea (figure by N.N. Kondakov). 545 32 Instances of the transgression of porpoises into rivers are not rare. They have been noticed not only in the Black Sea rivers (Don and Danube), but also in the Thames, Seine, Rhine, Elba, Neva, and other rivers. Sometimes the animals advance up the river for hundreds of kilo- meters from the estuary (Kleinenberg, 1956a; Tomilin, 1957, 1962). Seasonal migrations and transgressions. The migrations of common porpoises have not been studied although some seasonal migrations undoubtedly do occur. It is quite obvious that these porpoises spend only the summer months in the northern parts of the range of the Atlantic and Pacific oceans but abandon these waters in winter. The southward movement of these porpoises into the Baltic Sea is quite impressive. Short seasonal migrations have been observed in the Black Sea in which the animals spend the winter; in spring, they are seen in the Azov Sea but return again to the Black Sea for winter. The rare departures of common porpoises from the Black Sea into the Sea of Marmora and even into the Mediterranean Sea, their trans- gressions into the Mediterranean Sea from the Atlantic Ocean, and their passage in the easternmost part of the Baltic Sea (Kronshtadt) and through Neva even into Lake Ladoga, can be regarded as unusual transgressions. Reproduction, growth, and development. Data on reproduction biology are available mostly for the Azov-Black Sea populations but these can be extrapolated to the other two populations with sufficient justification. The periods of mating and parturition, although quite protracted, occur predominantly in the summer months. The Black Sea common porpoise mates from the end of June to October with a peak in August. The duration of gestation has been put at 9-10 months. Births occur from April to July. Most females whelp in May-June (Tsalkin, 1940) In the Baltic Sea, mating peaks in July and August while the largest number of calves are born in May and June. The growth rate of the embryo is quite high (Table 47). The embryos of common porpoises (Fig. 315) from the Baltic sea recorded an average weight increment of 5 g per day (total 160 g) in the one-month period from November 15 through December 15, 11 g per day (total 340 g) from December 15 through January 15, and 30 g per day (total 920 g) from January 15 through February 15. Usually a single large calf (twins are very rare) measuring half the length of the mother’s body, sometimes even longer, is delivered. The average length of a newborn calf is 75 cm (63-86 cm) and it weigh 2.8-7.9 kg. Lactation extends for about four months. The composition of the milk of the Azov-Black Sea common porpoise is as follows (7%): fat 33.9, protein 5.22, sugar 1.28, dry residue 7.1, ash 0.6, and water 59.0 (Ural’skaya, 1957). In some cases, the milk may contain up to 45.8% fat. 546 733 Fig. 315. Embryo of the common porpoise (figure by М.М. Kondakov). Table 47. Monthwise dimensions of embryos, mm (Tomilin, 1957) Month Azov-Black Sea basin North Atlantic Ocean and North Pacific Ocean Baltic Sea ; Number Number Number of of of animals Mean Min. Max. animals Mean Min. Max. animals Mean Min. Max. July ау 5 ПУ ое рае ит еб Aug. SOS Oe ee ee ee Sept. 8760 10 0 = Oct. 67 120 50 200 ее See Nov. 33 190 80 240 GT оО ee о Dec. LPO aD 300 ns ee CLS ee ee Jan. Gi 320190} S80 АГ. 429600225.) 380. 212 248.289. 258 Feb. ae en ere 23 E356 250) 450 ВИ ser all March 263 460 210 610 20153552001 550 10909 Ronee April 28 580 370 820 5101650] 1590174011912 14951180510 Мау 35 730 620 840 24) 1675610448740, мили 6б0Зн— В — June 7 820 780 850 eA VE SS ee Total 453 10 850 134 92. 1740058 239 510 The calf is capable of swimming and accompanying the mother immedi- ately after birth. The male to female ratio among newborn calves is close to one. At a body length of 130-145 cm, many females of the Black Sea common porpoise are still immature. More than half the females (59%) with a body length of 145-150 cm and 79-100% of females 150 cm or longer were gestating. The smallest gestating female among the Black Sea common porpoises was 130 cm long and among the Baltic Sea animals 136 cm (weight 50 kg) (Tomilin, 1962). Enemies, diseases, parasites, mortality, and competitors. In oceanic waters the killer whale represents the most dangerous enemy of the com- mon porpoise, which is a food item for the former. It is possible that the 734 porpoise may fall prey to large sharks. Apart from bone tumors, no other diseases have been reported for these porpoises. Very severe infection with helminths could be the cause of death in some cases. Among all the cetaceans, the common porpoise is one of the primary hosts for a large number of helminths. Eighteen species of helminths have been registered in the common porpoise: trematodes four, cestodes three, nematodes nine, and acantho- cephalans two. Two species of trematodes have been reported exclusively in common porpoises. Campula oblonga Cobbold, parasitizing the bile ducts of the liver, was detected in the European and American waters of the North Atlantic. Pholeter gastrophilus Kossack, parasitizing the mucous mem- brane of the pyloric section of the stomach, was found among dolphins caught in Baltiisk port. The trematode Distomum philocholum Creplin, localizing in the liver, has been encountered in white-sided dolphins in European waters as well as in common porpoises. This species para- sitizes the bile ducts of the liver of common porpoises and three species of pinnipeds in the Atlantic and northern Arctic oceans; it is also known in three species of land carnivores. The three species of cestodes belong to the genus Diphyllobothrium. Of these, D. stemmacephalum Cobbold was found in the small intestine of only common porpoises (North Atlantic and Black Sea on the Ruma- nian coasts); D. lanceolatum Krabbe parasitizes the intestine of common porpoises and four species of seals (North Atlantic and Pacific oceans); D. latum L., apart from common porpoise, is found in six species of pin- nipeds, several species of land carnivores, domestic animals, and man; it has been detected among marine animals in the North Atlantic and northern Arctic oceans. Anisakis (Anisakis) simplex Rudolphi, a nematode of common por- poises, is very widely distributed among marine mammals (parasite of the gullet, stomach, and intestine) and has been found in ten species of toothed whales, two species of baleen whales, and in Steller’s sea lion in the North Sea and in the Pacific Ocean (Kamchatka, Japan, and New Zealand). Anisakis (Anisakis) typica Diesing, in addition to common porpoises, is a parasite of the stomach of three species of dolphins from the North Sea and waters of South Africa in the Atlantic Ocean. Terra- nova (Terranova) decipiens Krabbe, apart from common porpoises, has been reported among many species of marine mammals (17 species and subspecies of pinnipeds, 2 species of baleen whales, and 1 species of dol- phins) and localizes in the stomach and intestine; it has been detected in the basin of the northern Arctic, Atlantic, and Pacific oceans, and around Antarctica. The nematode Pseudalius inflexcus Rudolphi has been found 547 735 in the bronchi, blood vessels, and heart only of common porpoises from the North Atlantic coasts of Europe and the Asian coasts of the Pacific Ocean (not recorded in USSR waters). Of the three species of the genus Halocerus, Halocerus (Prohalocerus) invaginatus Quekett has been found in the lungs of only common porpoises from the waters of California, the Baltic Sea, and the Atlantic Ocean (not found in USSR waters). Halo- cerus (Posthalocerus) taurica Delamure and Skrjabin and H. (Posthalo- cerus) ponticus Delamure parasitize the lungs of only the Azov-Black Sea common porpoise. Stenurus minor Kuhn, aside from Azov-Black Sea and Atlantic common porpoises, has been found in belugas in the northern Arctic Ocean, North Atlantic Ocean, North Sea, on the Asian coasts of the Pacific Ocean, and in the Black and Azov seas. It localizes in the bronchi, heart, blood vessels, and auditory organs. It is possible that the large number of the nematode Stenurus minor encountered in all the Azov Sea porpoises may cause deafness in them. Tozynurus convolutus Kuhn, parasitizing the bronchi and blood vessels of the lungs of animals in the European waters of the North Atlantic and coasts of Sakhalin in the Pacific Ocean, has been detected in pilot whales as well as common porpoises. The acanthocephalans Corynosoma semerme Forssell and Coryno- soma strumosum Rudolphi parasitize the intestine. The former is known in common porpoises, 6 species of pinnipeds, and 6 species of sea birds; the latter in common porpoises, 11 species of pinnipeds, belugas, 10 species of birds, and in cats and dogs. These worms have been detected in almost all the northern seas and in the Caspian Sea (Delamure, 1955). Field characteristics. These are rather small, predominantly coastal porpoises. The upper ‘part of the trunk is black and the underside light- colored. The tip of the dorsal fin is almost rectangular. These animals live in small groups. Intense flexure of the body while diving is characteristic. They generally do not breach the water. Economic Importance Common porpoises are of almost no economic importance. Danish fish- ermen catch several hundreds of porpoises a year in the Little Baelt Strait during their migration from the Baltic Sea. They chase them into the narrow strait and block them in nets. In other parts of the range, the catch is only incidental. Regular hunting of common porpoises is carried out only in the Azov and Black seas but here, too, this species is not of great economic importance. In the best years of hunting, the proportions of the differ- ent species of dolphins in the catch from this water body are: common 736 dolphin 200, common porpoise (from Azov Sea) 10, and bottlenose dolphin 1 (Tsalkin, 1937). This is based on a catch of 700-800 Azov Sea common porpoise; in some years up to 4,000 animals are caught and the proportions vary considerably (Bodrov, Grigor’ev, and ‘Tver’yanovich, 1958). The technique of catching the Azov Sea common porpoise, although not different from that used for catching the common dolphin, has cer- tain special features. The much slower Azov Sea common porpoise is usually corifined to small groups, distinctly separated, and in the coastal zone. Therefore, whalers spread the fleet in a broad front, advance for- 548 ward in flanks, and gradually press the animals toward the coast. The net is cast after the dolphins gather in a very dense group close to the coast. Oil, mainly melted from blubber, represents the chief product of hunting. Further, comparatively high-quality leather goods are produced and used satisfactorily in the footwear industry. Despite many success- ful experiments on using the flesh of Black Sea dolphins for human consumption, it has not found wide acceptance. The material remaining after obtaining the oil (musculature, skeleton, fins, viscera, etc.) is made into a flesh-bone meal which can be used as feed for farm animals or as a fertilizer. The hollow lower jaws of dolphins contain a very small amount of so-called jaw oil, from which high-quality oil is produced for lubricating fine mechanisms. In experiments, this oil froze after 10 hr of exposure to —7 to —8°C; the oil was then filtered at this temperature and transferred to a chamber at —25°C where it did not freeze for 24 hr (Okuneva, 1934). The total weight of the Azov-Black Sea common porpoise varies in the range 24 to 55 kg (Okuneva, 1934), or an average of 28 kg (Dragunov and Kasinova, 1951). The weight of the various trunk sections in different seasons is shown in Tables 48 and 49. Three very large female common porpoises from the North Pacific Ocean with a body length of 178.5 cm, 173 cm, and 168 cm weighed respectively: 75.2 kg, 70.3 kg, and 73.4 kg (Scheffer and Slipp, 1948). In the overall balance of useful products produced by hunting marine mammals in our country, common porpoises have no economic impor- tance whatsoever. As a result of the significant reduction in the pop- ulation of dolphins in the Black and Azov seas, hunting dolphins of all species (including the Black Sea common porpoise) has been totally banned since 1965. There is also no hunting in the other areas of the range of this species (in the Atlantic and Pacific oceans). (V.A.) 547 548 737 Genus of Dall Porpoises Genus Phocoenoides Andrews, 1911 1911. Phocoenoides. Andrews. Bull. Amer. Mus. Nat. Hist., 30, p. 31. Phocoenoides truei Andrews, 1911. Small porpoises with a body length up to 2 m. These porpoises lack a “beak”. The low dorsal fin lies slightly ante- rior to midbody. The flippers are relatively small. Table 48. Weight of the body sections of Azov Sea common porpoise caught in October and November (Okuneva, 1934) Item Male Female As % of As % of Weight, kg total weight Weight, kg total weight Total weight 24.0 - 27.0 100 37.0-55.0 100 Head 8123 775) 2.4-3.6 6.5 Skin Iasi 5 7258) 7.3-8.8 2.0-2.5 5.4-5.9 Trunk fat hs) 92 29.3 -35.4 LES = 7.5 25.3-30.6 Fins and tail 1.0. = 1:1 2.5 -4.1 1.5 -1.8 3.3-3.9 Muscles (flesh) 4.8 - 6.5 19.7-24.1 10.0- 15.5 24.5-27.1 Trunk bones 2.9- 43 10.8 -16.3 3.5- 6.5 9.5-11.8 Brain 0.3 - 0.4 1.2-1.4 — — Tongue ONS 02 0.5 - 0.6 0.15 0.40 Blood — — 1.0 2.17 Viscera, inclusive of: 3.4 - 4.2 10.6-15.4 6.0-8.5 13.0 - 17.7 Liver 0.5 - 0.6 1.7-2.3 1.1-1.6 2.0-3.3 Heart 0.15 0.62 0.15 0.40 Lungs 0.6 - 0.8 2.1-2.8 1.0-1.6 1.9-3.3 Stomach and intestine 14- 1.6 5.2-6.2 5.1-5.8 10.4-10.5 Kidney 0.1 - 0.2 0.3-0.6 0.20 0.36 Table 49. Weight of Azov Sea common porpoise in the spring catch (2,611 animals) (Tomilin, 1957) Total weight, Average weight, As % of kg kg total weight Total weight 78,888 30.2 100 Blubber 32,026 12.3 40.6 Muscles (flesh) 24,018 9.2 31.2 Fins 2,956 Л &Ы/ Глуег 1,566 0.6 2.0 Heart 263 0.1 0.3 549 738 The body color is dark (from dark steel to black). A large white patch occurs on each side of the body. The dorsal fin is partly or wholly white. The broad and flattened rostrum is shorter than the cranium. The pterygoid bones are separated. The teeth are very small and number 19-23-28 55-24-58. Vertebrae 92-98. Phalangeal formula: [_›, II¢-7, Ш4- в ГУ, - 3, and У, ->. Ribs 15-18 pairs, of which 3-5 anterior ones articulate with the sternum. Biology of Dall porpoise has not been well studied. They feed on cephalopods and fish. They are distributed in the North Pacific Ocean. The genus comprises a single species, P. dalli True, 1885. There is no hunting of this species. (V.S.) DALL PORPOISE Phocoenoides dalli True, 1885 1885. Phocoena dalli. True. Proc. U. S. Nat. Mus., 8, p. 95. 1911. Phocoenoides truei. Andrews. Bull. Am. Mus. N. H., 30, p. 32. Waters of the eastern coast of Japan; Rikuzen Peninsula, Hondo ‚ Island. (V.H.) Diagnosis Only species of the genus. Description The external appearance is similar to that of other common porpoises. The body is shortened and the anterior half is somewhat thickset. The head is short (Fig. 316). The upper jaw is slightly shorter than the lower. The dorsal fin forms a near-equilateral triangle, slightly concave along the posterior margin. The length of the base of the dorsal fin is 1.7-2.2 times its height. A few small horny tubercles are present on the anterior margin of the dorsal fin. High longitudinal keels are present on the caudal stem, both in the upper and lower parts. These keels increase with age. The flippers are short and have a broad base. The color of the head, dorsum, upper anterior and entire posterior part of the caudal stem is dark, from dark steel gray to black. A large white patch occurs on each flank and extends anteriorly up to the level of the anterior margin of the dorsal fin (P. d. dalli) or almost right up to the eye (P. d. truei). This patch posteriorly extends along the flanks of the caudal stem to the level of the anal opening but does not reach the ‘(aoyepuoy ‘м ‘М Aq ainsi) чеээо эцюеа ‘эпдт, yop 5эрлоизороца ‘[astodiod пе] “AI 971d р и yaad hi 549 550 739 Fig. 316. Dall porpoise, Phocoenoides ааШ (figure by М.М. Kondakov). abdomen. Individuals with a coloration intermediate between these two types have been sighted. The dorsal fin is wholly or partly white (dark only rarely). The flippers are somewhat lighter in color than the trunk, even white. in exceptional cases. Totally black specimens have also been encountered (Nishiwaki, 1966). Young animals are usually more deeply pigmented and newborn calves less pigmented than adult animals. The teeth are small, chisel-shaped, and barely rise above the gums. Tubercles of keratinized epithelium are present on the gums around the teeth. Some teeth may not cut through the gums. Thus, in an adult male, 5 of 15 teeth in the left half of the upper jaw and 9 of 16 in the right half had not emerged; 4 of 22 in the left half of the lower jaw and 4 of 22 in the right half were likewise unexposed (Benson and Grood, 1942). The interalveolar septa are poorly developed. The teeth are surrounded by tubercles of keratinized epithelium of the gums which could rise above the tooth crowns and function as a substitute or them. Cervical vertebrae 7, thoracic 15-18, lumbar 23-27, and caudal 39-49. The cervical section constitutes 3%, thoracic 23%, lumbar 33%, and caudal 41% of the length of the vertebral column. The vertebrae have a flattened centrum and a long thin spinous process reaching maximum height in the lumbar section. All the cervical vertebrae are fused. The minimum and maximum body lengths in 14 adult male and 13 adult female Dall porpoises were respectively (in cm): 172 and 210 (х 186) and 169 and 197 (x 181) (Mizue and Joshida, 1965). The main body measurements of two male Dall porpoises, one caught off the coast of Japan and the other from the Aleutian Islands, and a female caught off the coast of Japan (Tomilin, 1957; Norris, 1966) were respectively (in cm): body length 191, 183, and 182; distance from tip of snout to anterior margin of blowhole 25, 23, and 15; the same up to axilla 41, 36, 740 550 Fig. 317. Dall porpoise, Phocoenoides dalli, on the deck of а ship. Sea of Japan, 1965 (photograph by А.Е. Kuzin). 551 and 24; length of flippers 23, 20, and 23; maximum width of flippers 22, 10, and 10; height of dorsal fin 16, 15, and 16; width of caudal flukes (from tip to tip) 47, 47, and 47; and distance from anal opening to fork of caudal flukes 62, 58, and 51. The largest male caught to date had a body length of 210 cm and the female 198 cm. The skull measurements (Fig. 318) (average of three to five animals; Tomilin, 1957) were (in cm): condylobasal length 32, zygomatic width 19, length or rostrum 13, width of rostrum at base 10, length of lower jaw 25, and length of mandibular symphysis 3.7. (V.S.) Geographic Distribution This species is encountered in the North Pacific Ocean. Geographic Range in the USSR (Fig. 319) Coasts of the Sea of Japan commencing from Peter the Great Gulf, La Perouse Strait, waters of Sakhalin including the Gulf of Sakhalin and Kuril Islands, southeastern part of the Sea of Okhotsk, and western coast of Kamchatka. In the Bering Sea, it is encountered along the eastern coast of Kamchatka, in waters of Koryask land, Gulf of Anadyr and Bering Strait, and the adjoining waters of the Chukchi Sea. 550 553 741 Win 20. ААА а № "ль * (Ам, AMY Ч ty, Woh ee мили Fig. 318. Skull of the Dall porpoise, Phocaenoides ааШ (figure by М.М. Kondakov). Geographic Range outside the USSR (Fig. 320) Waters of the eastern and western coasts of Japan, Aleutian Islands, and eastern part of the Bering Sea, Gulf of Alaska, and along the Amer- ican mainland in the south up to 30° М lat. (McTaggart, 1944; Wilke, Taniwaki, and Kuroda, 1953; Tomilin, 1957; Hall and Kelson, 1959). (V.A) Geographic Variation Two well-distinguished subspecies, also encountered in our waters, are recognized in this species. Some researchers, however, do not recognize these forms while others treat them as independent species. Details of distribution at the points of contact of the ranges, the numerical ratios between the two forms at such places, etc. are not available. Their ranges in Our southern waters overlap. 1. Northern Dall porpoise, Р. d. dalli True, 1885. The white patch on the flanks and ventral side of the body does not extend to the flippers but terminates roughly on the vertical of the anterior margin of the dorsal fin. This subspecies is encountered in the waters south of Ussuri territory (Peter the Great Gulf and Pos’et Bay) and the southern islands of the Kuril range to the southern part of the Chukchi Sea inclusive. Outside the USSR, this subspecies is reported in the waters of the Korean peninsula and Japan in the south up to roughly 38° N lat. and in American waters from the Chukchi Sea up to roughly 34° М lat. 742 2. Southern Dall porpoise, Р. 4. гие Andersen, 1911. The white patch on the flanks runs anteriorly beyond the base of the flippers and almost reaches the eye; on the abdomen, it runs from the base of the caudal stem to the base of the flippers. A small white patch occurs on the throat. This subspecies is reported in the waters of the southernmost parts of Ussuri territory (Primor’e) in the north up to 42 to 43° N lat. (Pos’et Bay, Peter the Great Gulf, and more eastward), and possibly in the southern part of the Kuril range. Outside the USSR, it is reported in the waters of the Korean penin- sula and Japan between 38° and 43° N lat. Along the American coast, it hy НИ 551 Вр. 319. Range of the Dall porpoise, Phocoenoides dalli, т the USSR (V.A. Arsen’ev). 743 (лэмэзту У‘) yjop 52рлоизо2оц4 ‘asiodiod ива эцз Jo э8иел $90245 ‘00$ “314 И ИМ: п Е GSS, 744 has been indicated from 28 -30° М lat. to 45-50° М lat. (Klumov, 1959; Nishiwaki, 1966).'° (V.A.) Biology Population. Probably, one of the relatively abundant species, forming at times large herds of hundreds of animals. It is apparently quite abundant in the western parts of the Pacific Ocean along the coasts of Japan and the Kuril Islands. Food. Pelagic schooling fish and cephalopods constitute the main food. Food composition varies in different parts of the range. The stom- ach of six animals of the northern subspecies caught in the waters of Cal- ifornia and Oregon states contained hake (pike) (Merluccius productus), jack mackerel (Trachurus symmetricus), and squids (Loligo opalescens); the stomach of four animals from British Columbia contained herring, while capelin (Mallotus villosus) was found in the stomach of two ani- mals from the Gulf of Alaska. Lantern fish (few species) are of overriding importance in the waters of Japan and squids and other species of fish are of lesser importance (Wilke and Nicholson, 1958). In the Bering Sea, along the islands of the western part of the Aleutiian range and on the coasts of Kamchatka, squids predominate in the food (Fig. 321) while smaller fish and shrimps are consumed to a lesser extent. In most of the cases examined, various species of animals were found together in the stomach of porpoises (Mizue, Joshida, and Takemura, 1966). The species of cephalopods and fish found in the stomachs have not been - Stated. The food composition of northern and southern Dall porpoises compiled from specimens collected in the waters of Japan from March through June is compared in Table 50. Fishes of the family Sudidae are considered very rare and only rare specimens are found in ichthyological collections; nevertheless common porpoises apparently feed on them quite regularly. The stomach of the porpoises caught in the Sea of Japan contained saury and squids (Sleptsov, 1955). Behavior. Dall porpoises are most often encountered in small groups of 2 to 18 head but herds of 20 to 25 animals have been encountered 16 Hall and Kelson (1959) do not recognize this form and regard Long Beach and Los Angeles at 34° N lat. as the southern point of this range. According to the data of A.G. Tomilin (1957), our northern form covers the Sea of Okhotsk and Bering Sea as also the Sea of Japan. His more recent references correspond to those cited in the text. (V.H.) 555 745 Fig. 321. Squids in the stomach of a Dall porpoise. Pacific Ocean, east of Honshu Island, 1960 (photograph by G.M. Kosygin). in the fore-Kuril waters (Fig. 322). Sometimes herds of these dolphins contain 100 or more animals. While in motion, these porpoises breach the water often and sharply so, flying in the air over quite a long distance. On finding themselves in the vicinity of a ship, they run rapidly ahead of its bow, quickly changing from side to side, or swim along the board, easily overtaking the ship. The speed of these porpoises exceeds 20 km/hr. These porpoises are sighted close to the coasts as also far away from them in deepwater regions. The dolphins studied off the coasts of North America were caught at depths of 180 m and off the coasts of Japan at depths of 2,700 m. Migrations. Information on migrations is extremely fragmentary. In the northern parts of their geographic range, especially at the places of ice cover, these dolphins move from south to north and back, spending the winter season outside the ice-covered regions. Apparently, compar- atively small seasonal migrations are a feature of these porpoises in the other parts of their range also. 554 746 Table 50. Food of Dall porpoises (Wilke and Nicholson, 1958) Food item Southern subspecies Northern subspecies (86 animals) (7 animals) Number of % by _ Number of % by stomachs vol. stomachs vol. Fishes Lantern fishes: Notoscopelus sp. 27 21 2 37 Diaphus sp. 6 2 — — Taletonbeania taylori 5 1 3 11 Lampanychtus sp. 1 Traces — = Unestablished species 44 49 3 35 Sudidae—Paralepis sp. 7 4 2 10 Hake, Laemonema longipes 5 11 — — Mackerel, Scomber japonicus 2 1 — — $4 14$ Ommatostrephes sloanei-pacificus 22 5 1 2 Watasenia scintillans 35 6 3 3 Unestablished species 5 Traces 2 Traces | An \ № < \ № С WS Fig. 322. Dall porpoise, Phocoenoides dalli, at sea (figure by N.N. Kondakov). In the Pacific Ocean waters along the Canadian coast, Dall porpoises were encountered in summer mainly in the bays between the islands north of Vancouver Island. The animals prefer bays which are wide open on both sides with characteristic ebb-tide currents. In the waters of Japan, early in March, Dall porpoises are caught in the southern part of the range; large herds of these animals are seen in mid-March in the waters of Iwata Prefecture (about 40° N lat.). Hunting here continues up to June. Later, most of the porpoises migrate to the 556 747 coasts of Hokkaido and are caught again in autumn along the coasts of Honshu. In the coastal waters of the islands of Japan, hunting is carried out within the 30-mile coastal limit but most of the animals are concentrated in the summer at a distance of 10 to 15 miles from the coasts (Wilke, Taniwaki, and Kuroda, 1953). Reproduction. In May, 1950, embryos weighing over 6 kg were found in the females of the northern subspecies (Wilke er al., 1953). A fully formed embryo was found in a female porpoise in the waters of Queen Charlotte Island in November, 1926. It is possible that the period of parturition is greatly extended among these porpoises. South of Queen Charlotte Island, newborn calves were often sighted in August and not even once encountered before August 7. A large number of animals was studied over several years in the southwestern part of the Bering Sea from the second half of May to early August. Fully mature testes were not seen in the male porpoises during this period. Evidently, their mating occurs later, probably not earlier than August end. This is also supported by the results of studying the female reproductive systems: corpora lutea were not detected in the ovaries of females. The period of parturition has tentatively been placed in the second half of July to mid-August. Gestation among animals of this species thus extends for not less than a year. Among all the pregnant females, without exception, embryos lay exclusively in the left horn of the uterus; the state of the left horn indicated that the embryo had formed in it and the corpus luteum was present in the left ovary. Graafian follicles in the ovaries of all the nonproducing females, without exception, had developed in the left ovary while the right one appeared to be immature. Among the embryos, no difference was detected between the left and right ovaries but in a newborn calf the left ovary was already larger than the right one. Based on a small number of samples, it was tentatively determined that the body length of a calf at birth is 85-90 cm. Most females become mature after two full years and a body length of about 170 cm. Males achieve sexual maturity at the age of three years or even older and a body length roughly of 185 cm. The ratio between the males and females in the population is close to 1:1. Of the 54 animals studied, 26 were females and 28 males (Mizue and Joshida, 1965; Mizue, Joshida, and Takemura, 1966). A slightly different distribution of animals of different sexes has been noticed among the southern subspecies. As established from the results 557 556 748 of spring hunting (from March through May), males remain in the south- ern and females in the northern half of their range (Wilke et al., 1953). Enemies, diseases, parasites, mortality, and competitors. The nema- tode Halocerus (Prohalocerus) kirbyi Dougherty, not known among other marine mammals, has been detected in the lungs of animals from San Franciscan waters. Н. dalli and Irukanema аа are also known. In the stomach of a lone dissected animal from waters of the Kuril Islands, an immature Anisakis sp. was detected (Delamure, 1955; A. Skrjabin, 1960). Field diagnosis: Sharply demarcated large white patches (“‘wings”’) on the sides and a white smear on the dorsal fin are characteristic of this dolphin (Fig. 323). (V.A.) Economic Importance This species is of limited hunting importance only in Japan where up to a thousand or even more porpoises are caught each year. No catches have been reported from other parts of the range. Dall porpoises are hunted along with other species of dolphins and small whales, using small craft (20-30 tons) with a crew of 10-12 per- sons. These vessels are used in spring to catch these porpoises and sharks and swordfish in summer; these vessels are again used in spring to catch Fig. 323. Dall porpoise, Sea of Japan, 1965 (photograph by А.Е. Kuzin). 749 porpoises. These ships cruise at 7-10 miles/hr. Shooting with guns and harpooning the killed animals manually is done from a special wooden platform constructed on the bow. From time to time, when the por- poises approach the vessel quite close, they can be harpooned without using firearms. In most cases, the killed porpoise floats on the water surface for a few minutes in a vertical position with the head above the water. Killed porpoises have even been found floating a day or two days after shooting. Nevertheless, a large number of killed animals escape the hunter’s bag. Thus, in 1950, about one-third of the killed Dall porpoises were lost. With the growing salmon fisheries in the open sea, Japanese fisher- men have begun catching a large number of Dall porpoises every year. The regions of porpoise catching have been identified as regions in which the salmon are caught using gill nets. This catch is practiced in the north- western part of the Pacific Ocean and in the southwestern part of the Bering Sea, including coastal waters of the Aleutian island range and Kamchatka coasts. Porpoises fall into salmon nets in coastal waters more often than in the open sea. It has been assumed that porpoises are also abundant east of this region of salmon catching, where salmon are not fished. Porpoises probably become entangled in the nets when they feed on the fish caught in them. However, an analysis of their stomach con- tents did not reveal a significant number of salmon. In some years, some tens and even more than a thousand Dall porpoises fall into salmon nets, but until recently these porpoises were not utilized and fishermen threw them overboard (Mizue and Joshida, 1965). The output of Japanese porpoise hunting is disposed mainly in the market. The carcass is prepared for sale as follows. The flippers dorsal fin and caudal flukes are dismembered. Two longitudinal sections are made through the skin (along the dorsum and belly) from the tip of the snout to the tail. Later, blubber with the skin is separated from the trunk in two pieces. The flesh is separated from the vertebral column, also in two pieces, the ribs cut out, and the abdominal cavity cleared by removing the intestine, heart, liver, and kidneys, which are sold as edible products. The skin is tanned but the product is of rather poor quality. Oil is rendered from the blubber; fat from the head and jaw regions serves as raw material for producing excellent machine oil. Skeletal bones and all other remnants are processed into a fertilizer meal. The flesh is quite often used as food. Females weigh 91 - 115 kg (М 95 kg) and males 84-118 kg (М 102 kg) (McTaggart, 1944; Wilke, Taniwaki, and Kuroda, 1953; Klumov, 1959). (V.A.) 558 750 Genus of Black Finless Porpoises Genus Neophocaena Palmer, 1899 1846. Neomeris. Gray. Zoology. Voyage Erebus and Terror, I, Mamm., p. 30. Delphinus phocaenoides G. Cuvier. Nom. praeocc. (Neomeris Costa, 1844, from the group Vermes). 1847. Meomeris. Gray. List Osteol. Spec. Brit. Mus., pp. XII, 36. Not important from nomenclatural viewpoint. 1899. Neophocaena. Palmer. Proc. Biol. Soc. Washington, 13, p. 23. Sub- stituted for preoccupied Neomeris Gray, 1846. (V.H.) Maximum body length, up to 190 cm. The head is rounded. No “beak” is evident. A dorsal fin is lacking. A band of fine horny tubercles runs along the middorsal line. The body is lead-black but the abdomen light-colored. The rostrum of the skull is rounded anteriorly, broad, and short (much shorter than the cranium). The premaxillae are broad and not pinched anteriorly. The small pterygoid bones are widely separated. The teeth have broadened and flattened crowns and number #=12. Vertebrae 60-63. Phalangeal formula: 15, II,;-7, III,;_;, [V3, and V>. Ribs, usually in 14 pairs. Almost nothing is known about the biology of these animals. They feed on bottom-dwelling crustaceans, fish, and cephalopods. They inhabit the warm waters of the Indian Ocean and the western part of the Pacific Ocean. They usually live near the coasts and transgress into rivers. They have been reported from the Far Eastern waters of the USSR but none have been caught. The genus comprises a single species, N. phocoenoides G. Cuvier, 1829. (V.S.) BLACK FINLESS PORPOISE Neophocaena phocaenoides G. Cuvier, 1829 1829. Delphinus phocoenoides. G. Cuvier. Régne animale, I, p. 291. Cape of Good Hope, South Africa. (V.H.) Diagnosis Only species of the genus. Description Similar to the other common porpoises in outer appearance. The body build is compact. The trunk is somewhat stunted and thickened in the anterior portion. The head is relatively short (Fig. 324). The adipose 559 ПЭ Fig. 324. Black finless porpoise, Neophocaena phocaenoides (figure by М.М. Kondakov). body (corpus adiposum) [melon] on the head is highly developed and projects forward in the frontal part. The mouth section is small and its corners are upturned. The flippers are quite broad and crescent-shaped. Along the middorsum, from the cervical section to the anus, a band of horny tubercles, 3-6 cm wide, occurs, which broadens slightly anteriorly and narrows posteriorly. Much larger tubercles (up to 0.5 mm high and 2-2.5 mm in diameter) occur along the margins of the band. In the embryos, these tubercles are disposed one each on quadrangular plates closely adjoining each other. The upper lip of the embryo sports eight whiskers (four on each side). The lead-black color on the upper side of the body gradually light- ens notably on the abdomen. A gray patch is visible on the thorax and abdomen. The flippers and caudal flukes sometimes have light-colored sections of varying size, Cervical vertebrae 7, thoracic 13-14, lumbar 11-14, and caudal 28-31. The first five cervical vertebrae are fused. The main body measurements of an adult female black finless por- poise (Tomilin, 1957) are (in cm): body length 124; distance from tip of snout to anal opening 8.5, to base of flippers 23, to posterior margin of blowhole 9; width of caudal flukes from corner to corner 43; maximum length of flippers 26 and maximum width 8. The measurements of the skull (Fig. 325) of a male black finless porpoise with a body length of 116 cm are (Tomilin, 1957) (in cm): condylobasal length 19, zygomatic width 12, length of rostrum 7, width of rostrum at base 6, length of lower jaw 13, and length of mandibular symphysis 1.5. In the skeleton of one male (length of vertebral column 85 cm) there were 7 cervical vertebrae, 12 thoracic, 14 lumbar, and 25 caudal; the cervical section constituted 4.5%, thoracic 27.6%, lumbar 37.2%, and caudal 30.7% of the vertebral length (Tomilin, 1957) (it is possible that some caudal vertebrae were lost). (V.S.) Geographic Distribution Warm waters of the western part of the Pacific Ocean and the Indian Ocean. 752 Fig. 325. Skull of the black finless porpoise, Neophocaena phocaenoides (figure by N.N. Kondakov). Geographic Range in the USSR (Fig. 326) Pacific Ocean waters of the southern Kuril Islands; transgressions are possible into the waters of the Sea of Japan.!’ Geographic Range outside the USSR (Fig. 327) Waters of Japan, Korean peninsula, China (South China Sea), Kaliman- tan and other islands, and also Malaccan peninsula. It is known in the Indian Ocean in the Bay of Bengal and the Persian Gulf, the Arabian Sea, and along the coasts of Africa up to the Cape of Good Hope (Sleptsov, 1961; Tomilin, 1962). (V.A.) Geographic Variation Not established. Biology Information on biology is extremely scant. Crustaceans (macrura group: Penaeus, Palaemon, and especially P. japonicus), cephalopods (Loligo sp. and Sepia sp.), and some species of fish serve as food objects. They consume predominantly benthic animals and hence the black finless por- poise can be regarded as a coastal form. According to observations made 17 Once, in 1951, two porpoises were encountered 20 to 25 miles off Shpanberg Island in the southern Kuril Island range (Sleptsov, 1952). This species has not been caught in our waters. 561 753 Fig. 326. Range of the black finless porpoise, Neophocaena phocaenoides, in the USSR (V.A. Arsen’ev). on the Kuril Islands (Sleptsov, 1961), it is encountered in open waters, mainly in small groups (five or six animals) and singly. Sometimes the animals gather far away from the coasts into very large herds consisting of several tens of animals. Quite often, they transgress into rivers in which they spend much time, ascending hundreds of kilometers upstream (along the Yangtze up to 1,800 km; also sighted in Lake Dongting Hu). Their movements are gentle and they rarely breach the water. Migrations have not been studied. It has been assumed that parturi- tion occurs in October since a female 119 cm long with an embryo 52 cm long was encountered in this month. Seven species of helminths have been registered among black fin- less porposes: trematodes three, cestodes one, and nematodes three. The trematodes Campula folium Ozaki, Orthospianchus elongatus Ozaki, and 560 754 Nasitrema spathulatum Ozaki were detected in the waters of Japan and are known only in the black finless porpoise. The first parasitizes the liver, the second the intestine, and the third the nasal cavity. The ces- tode, Diphyllobothrium fuhrmanni Hsu, detected in waters of Japan and China and parasitizing the small intestine, in addition to the black finless porpoise, has been found in spotted dolphins. All the three species of nematodes were found only in the black finless porpoise from the waters of China. Halocercus pingi Wu localizes in the lungs, Sternus auditivus Hst and Hoeppli in the auditory organs, and Onchocerca fulleborni Hoep- pli and Hsu in the musculature. Most of the species of helminths found in the black finless porpoise have not been detected in other marine mammals. The black finless porpoise has no economic importance whatsoever; it is not hunted except for a few stray animals caught in the rivers of China (Delamure, 1955; Sleptsov, 1955, 1961; Tomilin, 1957, 1962). (V.A.) ’ Fig. 327. Species range of the black finless porpoise, Neophocaena phocaenoides (V.A. Arsen’ev). 562 755 Family of Narwhals Family MONODONTIDAE Gray, 1821 Cetaceans of medium dimensions, with body length up to 610 cm. A dorsal fin is absent although a long low fold occurs on the dorsum. The head is rounded, small, without a beak, and demarcated from the trunk by a distinct neck. The flippers are short and broad. Body coloration is subject to age-related variations: it is dark in young animals, turning light with advancing age. The rostrum is relatively short and broad. The frontal and interpari- etal bones form a crest. The petrous temporal bone grows toward the skull. The broad pterygoids articulate with the squamosals. Teeth ум. Vertebrae 49-55. The first and second cervical vertebrae are not fused whiie the rest may be fused in varying groups. The ulna has.no olecranon process. The geographic range of these animals is restricted to the waters of the northern polar region (Fig. 328). The family comprises two genera: (1) belugas or white whales, Del- phinapterus Lacépéde and (2) narwhals or unicorns, Monodon Linnaeus. Belugas are of economic importance. (V.S.) Genus of Belugas or White Whales Genus Delphinapterus Lacépéde, 1804 1804. Delphinapterus. Lacépéde. Hist. Nat. Cétacées, р. XLI, 243. Del- phinapterus beluga Lacépéde = Delphinus leucas Pallas, 1766. 1815. Beluga. Rafinesque. Anal. Nat., p. 60. Substituted for Delphi- napterus Lacépéde, 1804. (V.H.) Dimensions are the smallest in the subfamily, with a body length up to 6 m. The body is markedly elongated. The relatively small head has a very prominent frontal projection. The monochromatic body is dark in the young, lightening to white (ivory) or yellow in the adult. The epidermis is remarkably well developed. The relatively narrow skull is flattened dorsoventrally. The rostrum, broad at the base, is roughly 1.5 times longer than the cranium. The maxillae dorsally adjoin the bony nares and extend far backward, almost up to the occipital bones. The pterygoids are well developed. Powerful crests are typical. Teeth 5-1, often irregular in form, and sometimes with supplementary cusps. Vertebrae 49-54. Some of the cervical vertebrae are fused. Of the 11-12 pairs of ribs, 5-7 pairs are fused to the sternum. 756 Fig. 328. Range of the narwhal family, Monodontidae (V.A. Arsen’ev). 561 563 ПЭ The sternum is either non-segmented or consists of two or three sections (sometimes even six). Phalangeal formula: I,_4, Пв-э, Шб-в› [V5-7, and V,-.- Digits III and IV are almost equal in length. The number of pha- langes of some digits in embryos is reduced in adult animals. Sometimes the digits are split and resemble claws. Belugas are ichthyophagous; they feed mainly in the coastal zone of the sea. Gestation extends for 11-12 months. Females calve annually. These animals are distributed in the seas of the North Atlantic, Arc- tic, and North Pacific oceans. Fossil remains have been traced in the Pleistocene of North America. The genus comprises a single species: the beluga, D. /eucas Pallas, 1776. These animals are of commercial importance. (V.S.) BELUGA OR WHITE WHALE!’ Delphinapterus leucas Pallas, 1776 1776. Delphinus leucas. Pallas. Reise durch versch. Prov. Russ. Reiches, 3 (I), p. 497. Ob’ estuary. 1776. Delphinus albicans. Muller, Zool. Danicae prodromus, p. 7. Green- land. 1804. Delphinapterus beluga. Lacépéde. Hist. Nat. Cétacées, pp. XLI, 243. Davis Strait. 1935. Delphinapterus dorofeevi. Klumov and Barabasch. Zh. “Rybnoe Khozyaistvo SSSR,” No. 11. Sakhalin Bay, Sea of Okhotsk. 1935. Delphinapterus freimani. Klumov. [bid., no. 7. Dvina Bay, White Sea. 1935.- Delphinapterus leucas maris-albi. Ostroumov. Zh. Za Rybnuyu Industriyu Severa, no. 11. Onega Bay, White Sea. (V.H.) Diagnosis Only species of the genus. Description Age-related changes of body proportions are known among belugas (Arsen’ev, 1936a; Vladykov, 1943, 1944; Kleinenberg et al., 1964). The changes pertain to an increase in ratio of body length to radius of the trunk and relative (to body length) reduction of length of the flippers 18 The local Russian people (coastal population) call the animal “beluga”. “Belukha” represents an artificial (bookish) modification of “beluga” to distinguish the whale from “beluga fish” (giant sturgeon) (Heptner, 1930; Chapskii, 1937). 564 758 as well as narrowing of the caudal flukes. The form of the flippers of belugas is typical: their proximal margin is flexed, the flexure very distinctly manifest in old animals (Vladykov, 1943). Such a form of the flippers (Fig. 329) is possibly important for braking during sharp turns (Kleinenberg et al., 1964). The absence of a dorsal fin is explained by the adaptation of the animal to living amidst ice (the fin could interfere in floating in such conditions; Heptner, 1930) and, further, enables the animal to bend along the longitudinal axis of the body, which is important in catching fish (Yablokov, 1959). A rather low leathery crest replaces the missing dorsal fin in some animals. The epidermis (up to 7-11 mm) and the horny layer [stratum corneum] (up to 1-6 mm) are highly thickened; the dermal layer is very well developed while the subcutaneous adipose tissue is relatively thicker compared to other whales (Bel’kovich, 1959; Kleinenberg et al., 1964). Age-related changes of body color are significant. The light gray color of the newborn changes after a few days or a week into almost black in suckling calves, but lightens gradually thereafter (Bel’kovich, 1959). Whitening is more rapid on the dorsum, body flanks, and abdomen. How- ever, the dark-colored edge of the caudal flukes undergoes no depigmen- tation. The body surface of most of the animals is covered in numerous scars, spots, and stripes. The white color of the belugas is considered a protective adap- tation (Kukenthal, 1900). It has also been suggested that such a col- oration frightens schools of fish, which facilitates their catch by the whale (Yablokov, 1956). Another view (obviously erroneous) holds that the white coloration reduces heat dissipation from the body of the animal (Chapskii, 1941). Four color groups of belugas (Fig. 330), corresponding en Meee typ pple et и, ИЛИ, 77 277: й Fig. 329. Beluga, Delphinapterus leucas (figure Бу М.М. Kondakov). 564 159 to definite age groups (Dorofeev and Klumov, 1936), are usually recog- nized (for more details, see p. 782). However, it has been opined that the variations in coloration of the beluga body should not be taken as an accurate criterion for determining the age of these animals (Bel’kovich, 1959). The simple peg-shaped teeth of belugas are characterized by a typical cutting and supporting form developed as a consequence of the wearing down of the teeth (Yablokov, 1958a, 1959). The maximum length of the teeth in the upper jaw is 5.8 cm and thickness 1.2 cm; in the lower jaw, 5.0 cm and 1.8 cm respectively (Yablokov, 1959). The largest teeth occur in the anterior part of the upper jaw and midpart of the lower jaw. Teeth are usually lacking on the premaxillae. Teeth are cut in year-old calves. Cervical vertebrae 7, thoracic 11-12, lumbar 6-12, and caudal 21-26. The anterior portion of the digestive tract of the beluga, right up to the second chamber of the stomach, is covered with Keratinized stratified epithelium in which (up to the first chamber of the stomach inclusive) a network of tonofibrillae, reinforcing the epithelium, is well developed. Such adaptations help the animal to swallow a large quarry whole without mastication (Kleinenberg, Yablokov and Tarasevich, 1958). The stomach consists of five chambers. The intestine, length varying from 22.5 to 37 m, is not divided distinctly into a small and large intestine due to the absence of a cecum. The duodenum is highly developed; the last third of the rectum is covered by stratified epithelium forming low longitudinal nonexpanding folds, with a powerful muscular cover supplemented by a powerful anal sphincter. Sexual dimorphism is manifest in the dimensions of the animals; males are larger than females. Moreover, the ratio of body length to radius of its cross section is more in males than females, underscoring the excellent hydrodynamic form of the male body; the caudal flukes of males are relatively larger while the flippers, on the contrary, are Fig. 330. Age-related color groups of the beluga, Delphinapterus leucas (figure by N.N. Kondakov). 565 564 760 relatively smaller. In males, the number of teeth is slightly more than in females (Yablokov, 1959). The frontal prominence in most males is more powerfully developed (this is particularly true of aged males). The maximum length of belugas in the Far East is 600 cm among males and 500 cm among females; in the Kara Sea 472 and 413 cm; Gulf ‘of St. Lawrence 447 and 409 cm; western Greenland 572 and 474 ст; Hudson Bay 448 and 400 cm; and the Beaufort Sea 460 and 383 cm (Kleinenberg et al. 1964). The main body measurements of Kara Sea belugas (Tomilin, 1957) (average for 13-14 males and 11 females) are respectively (in cm): body length 410 and 365; length of head 52 and 46; distance from tip of snout to base of flippers 30 and 71; distance from anal opening to fork between caudal flukes 94 and 89; length of flippers 44 and 39, width of flippers 31 and 26; and width of caudal flukes (from tip to tip) 88 and 77. The main measurements of the skull (Fig. 331) of belugas from the White and Kara seas and the Far East (Tomilin, 1957) respectively aver- age (in cm): condylobasal length 52 (10 measurements), 57 (10), and 59 (10); zygomatic width 28 (8), 30 (10), and 32 (4); length of rostrum 25 (10), 28 (10), and 29 (4); width of rostrum at base 17 (9), 18 (10), and 19 (4); and mandibular length 42 (4), 42 (8), and 44 (9). (V.S.) Fig. 331. Skull of the beluga, Delphinapterus leucas (figure by N.N. Kondakov). 761 Geographic Distribution Predominantly the cold waters of the Northern hemisphere where it is very widely distributed almost all around the polar region. Geographic Range in the USSR Apparently no changes have taken place in the range over the historic past. In the Barents Sea, beluga is quite regularly encountered along the Murman coast (mainly in winter); it is sighted almost year-round at Kanin and Kolguyev, and in the Chesha Bay. It is quite common in the Pechora Sea, Vaigach, and along the west coast of Novaya Zemlya. At the end of summer, it is not infrequent around Franz Josef Land. It is most numerous probably in the southeastern corner of the Barents Sea where is is sighted throughout the year. It lives mainly along coasts and has not been sighted away from them. There is no doubt of its absence in the open Barents Sea (Fig. 332). The beluga inhabits the entire White Sea. As an exception, it trans- gresses into the Baltic Sea where it has been sighted in our waters, espe- cially in the Riga (Parnu) and Finland gulfs. In the Kara Sea, it pre- dominantly covers the southwestern and western parts. It is encountered along the east coast of Novaya Zemlya right up to Cape Zhelaniya, at Vaigach, in Vaidarask and Gulf of Ob in Yenisey Gulf and also along the coast east up to Vil’kitskiy Strait. The beluga inhabits the Kara Sea mainly in summer. It can be encountered in autumn in its northern parts. At some places (Gulf of Ob) small herds are sometimes encountered in winter. In the Laptev Sea, the beluga is seen in the summer months along much of the coast from Severnaya Zemlya islands to Novosibirsk. It is most common in the southwestern part of the sea (Pronchishcheva Bay, Begichev Island, etc.), and also in the Lena estuary. From the Novosibirsk islands, the beluga is seen in the waters of the East Siberian Sea. In the winter months it is probably confined to the middle and northern parts of the Laptev Sea. It inhabits the Chukchi Sea predominantly in the summer months. Distribution along the Chukchi peninsula is restricted to the coastal section from Cape Dezhnev to Cape Shmidt but sometimes transgresses into the waters of the East Siberian Sea. Under favorable ice conditions, it moves far north. It is quite regularly found close to Wrangel Island and probably reaches 74 to 75° N lat. It is encountered in Bering Strait. In the Bering Sea, it inhabits the Gulf of Anadyr in which it is most numerous close to the Anadyr estuary. Its distribution southward 762 (2 es у < Ly и (. 2, —' aaa” ES a: OL 09 a x >> © г 5 Se Males ) Wemeg 23 a < 567 763 is limited to Cape Navarin or only slightly farther. Distribution in the open waters of the northern part of the Bering Sea has not been studied. It is found almost all along the coast of the Sea of Okhotsk. Depend- ing on the period of the year, it is common along the Sakhalin coasts (east coast, Terpeniya Bay, La Perouse and Tatar Straits, Amur liman, and the Gulf of Sakhalin). In the southwestern part of the Sea of Okhotsk, it ranges from the Amur estuary to Uda and Ayan bays in the west. It is common along the northern coasts of the Sea of Okhotsk, including Shelikhov Gulf, and the west coast of Kamchatka, mainly its northern part. It is rare along the west coast of the Sea of Okhotsk from Ayan Bay to Okhotsk. It has not been reported in the waters of the Kuril Islands. Geographic Range outside the USSR In the basin of the Atlantic Ocean, it is widely distributed in the Cana- dian Arctic archipelago (Fig. 333). It is encountered in the Gulf of St. Lawrence, along the Labrador coasts, in Ungava Bay, Hudson Bay, Hudson Strait, Foxe Basin, Davis Strait, along the coasts of Greenland and Baffin Island, and in Baffin Bay where it reaches north of Smith Strait. It is encountered in the bays of Lancaster, Jones, and Barrow, and in the Beaufort Sea. It has been sighted in McClure Strait, Amund- sen Gulf, and at many points along the coast in the region of Mackenzie Island. The beluga is also encountered east of the Canadian coast close to Iceland, Jan Mayen Island, Spitsbergen, and Finmark. Its distribution along the east coast of Greenland has not been clearly established. In the Bering Sea, it is known along the coasts of Alaska from Bristol _ Bay to Bering Strait and farther, in the Chukchi Sea up to Cape Bar- 568 row on the northern coast of Alaska. It is also encountered east of this point (Arsen’ev, 1939; Vladykov, 1944; Tomilin, 1957; Sergeant, 1962a; Kleinenberg, Yablokov, Bel’kovich, and Tarasevich, 1964). (V.A.). Geographic Variation The monotypic state of the genus Delphinapterus has been reviewed time and again but the new species described are not usually recognized while the probability of some subspecies very recently described has invited serious dispute. Preliminarily, the existence of three subspecies of belugas may be recognized in our waters: 1. White Sea beluga, D. | maris-albi Ostroumov, 1935. Smallest of the forms; adults 253-376 cm (x 312). This subspecies is found in the waters of the White Sea and the southern part of the Barents Sea. Not reported outside the USSR: ‘(дэ иэзту У ‘A) злэлы ш зе8эа Jo зриц Jo says ay} 1иэзэл4эл $10 502721 smuaidnuiydjag ‘e8njaq эцз JO э8иел $10946 ‘65$ ‘8 195 765 2. Kara Sea beluga, D. /. leucas Pallas, 1776. Occupies an intermediate position among the three subspecies in body dimensions. Adults vary in length from 318 to 464 cm (х 390). This subspecies is found in the Barents, Kara, and Laptev seas. Outside the USSR, it apparently lives in the remaining waters of the North Atlantic. 3. Far Eastern beluga, D. /. dorofeevi Klumov and Barabasch, 1935. Largest of the subspecies. Adults vary in length from 320 to 600 cm (х 424). This subspecies is found in the Okhotsk, Bering, and Chukchi seas. Outside the USSR, it lives in the American waters of the aforesaid seas. A systematic analysis of belugas from the western Atlantic and east- ern Pacific Oceans in relation to the above forms has not been attempted and the classification is tentative. It is possible that the populations in some of our Far Eastern water bodies are not wholly identical from the viewpoint of systematics. Each of the subspecies occupies an immense range and it is not unlikely that many local populations exist within these ranges. Proba- bly, belugas of the Laptev Sea form a distinct local population. Belugas from the Okhotsk and Bering seas definitely represent isolated popula- tions, distinctly separated from each other by a large distance. Within the Okhotsk population per se, two relatively isolated herds have been postulated (see “Seasonal Migrations and Transgressions”’). Belugas inhabiting waters outside the limits of the USSR also probably form several local populations. One such occupies the zone Iceland—Greenland—Spitsbergen. In Canadian waters, three such populations can be recognized: in the Gulf of St. Lawrence, in the eastern Canadian Arctic, and in the western Canadian Arctic or the Beaufort Sea. The first of these, even if it mixes at all with the eastern Arctic population, does so very minimally. The two arctic populations of America are separated by 20° along the longitude in the zone of Melville Strait (Sergeant, 1962a). However, there is as yet no adequate information to resolve this problem. (V.A.) Biology Population. The beluga may be regarded as a moderately populous species. According to approximate calculations, the population of White Sea beluga comprises 8,000 - 10,000 and of the Kara Sea 40,000 - 50,000 animals (Klumov, 1939). No less than several tens of thousands of animals live in the Sea of Okhotsk but the Bering Sea population is 766 probably somewhat less. Information is not available for judging the beluga population in other parts of the range within USSR waters. The commercial utilization of all these populations is so insignificant that hunting has no bearing whatsoever on the natural population dynamics. Food. Fish and crustaceans serve as food objects. The extensive range occupied by the beluga provides quite а large number of species of food objects which differ in various parts of the range. However, fish every- 571 where occupy a predominant position. The large number of organisms found in the stomachs contain some which enter incidentally or represent the food of animals on which the beluga feeds (Table 51). Fish (pelagic and benthic) which form massive schools and bottom- dwelling crustaceans constitute the main food. The least number of empty stomachs among the belugas of the Sea of Okhotsk is seen during the massive arrival of salmon (Arsen’ev, 1939); in the northern seas, the beluga feeds most intensely in the period of arrival of arctic cod (Klumov, 1936a; Tarasevich, 1960a, 1960b). Based on the frequency of encounter and the number of animals found in the stomach of belugas, their main food items are: in the White Sea—herring, capelin, and shrimps; in the Kara Sea—arctic cod, cisco, and other whitefish; in the Sea of Okhotsk—salmon (chum salmon and humpback salmon), navaga, herring, and isopods; in the Gulf of St. Lawrence—capelin, sand eel, cephalopods, and Nereis; and in Hudson Bay—capelin, cephalopods, and Nereis. Arctic char (Salvelinus alpinus; Daan and Douglas, 1953) was additionally encountered in the stomach of belugas of the Canadian Arctic. A seasonal change of food intake, also having a bearing on the age of belugas, has been noticed. For the White Sea beluga inhabiting the waters of Kanin, the main food objects in June are lumpsucker_ and floun- der, changing to navaga, cod, and shrimps in mid-July, and herring and flounder at July end (Klumov, 1936a). In the Gulf of Sakhalin of the Sea of Okhotsk, young belugas at the end of the milk suckling period consume mainly crustaceans, which constitute up to 50% of their ration, and relatively small fish. As the animal grows, the importance of crus- taceans and small fish gives place to large fish (Salmon) whose specific proportion in the intake increases. Large fish serve as the main food of adult animals in which crustaceans account for less than 9% of the intake (Arsen’ev, 1939). Belugas of the Gulf of St. Lawrence also exhibit differences in food intake relative to the regions, seasons, age, and even sex groups (Vladykov, 1947). The food of the beluga has been studied everywhere only in the summer months. No information is available on its winter sustenance. 569 767 Table 51. Food objects of belugas White Sea (Klumov, 1936a) Fish Atlantic herring, Clupea harengus L. West Atlantic capelin, Mallotus villosus Mull. European smelt, Osmerus eperlanus dentex Steind Arctic lamprey, Lampetra japonica tentrionalis Berg Atlantic navaga, Eleginus navaga (Pall.) Lumpsucker, Cyclopterus lumpus (L.) Atlantic cod, Gadus morrhua L. Haddock, Melanogrammus aeglefinus (Hodd.) Flounders: Pleuronectes flesus Р. P. platessa L. Liopsetta glacilis (Pal.) Crustaceans Shrimp, Crangon crangon L. Barents and Kara seas (Klumov, 1936a) Fish Arctic cod, Boreogadus saida (Lepech.) Arctic cisco, Coregonus autumnalis (Pall.) Siberian cisco, Coregonus sardinella Val. Muksun, Coregonus muksun (Pall.) Siberian sturgeon, Acipenser baeri Brandt Nelma, Stenodus leucichthys nelma Pall. Pike, Esox lucius (L.) Siberian powan, Coregonus lavaretus pidschian (Gmel.) Atlantic herring, Clupea harengus L. Smelt, Osmerus sp. Arctic char, Salvelinus alpinus Crustaceans Isopod, Mesidothea entomon Mesidothea sabini Gammaridae Sea of Okhotsk (V.A. Arsen’ev, 1939) Fish Chum, Oncorhynchus keta Walb. Humpback salmon, Oncorhynchus gorbuscha Walb. Navaga, Eleginus navaga gracilis Til. Pacific herring, Clupea harengus pallasi Val. Flounder, Pleuronectidae Rudd, Leuciscus brandti Dyb. Smelt, Hypomesus sp. Goby, Myoxocephalus sp. Pacific eelpout, Zoarces elongatus Kner. Blenny, Blenniidae Arctic lamprey, Lampetra japonica (Martens) Sakhalin char, Salvelinus leacomaenis Pall. Okhotsk whitefish, Coregonus ussuriensis Berg Gulf of St. Lawrence (Vladykov, 1947) Fish Haddock, Melanogrammus aeglefinus (L.) Ocean pout, Zoarces anguillaris (Peck) Blenny, Blenniidae Snailfish, Neoliparis atlanticus (Mull.) Capelin, Mallotus villosus Mull. Sculpin, Myoxocephalus scorpius groen- landicus (Cuvier and Valenciennes) Staghorn sculpin, Gymnocanthus tricuspis (Reinhardt) American smelt, Osmerus mordax (Mitchill) Sturgeon, Acipenser oxyrhynchus Mitch. Atlantic herring, Clupea harengus L. Sea lamprey, Petromyzon marinus L. American sand lance, Ammodytes americanus De Kay Atlantic tomcod, Microgadus tomcod (Walbaum) Contd. 768 Table 51 Continued Sea of Okhotsk (V.A. Arsen’ev, 1939) Gulf of St. Lawrence (Vladykov, 1947) Crustaceans Red hake, Urophycis chuss (Walbaum) Shrimp, Crangon septemspinosa Say Baltic cod, Gadus callaris L. Isopod, Mesidothea entomon orientalis Greenland cod, Gadus ogac Reinhardt Gurjanova Shrenck’s crayfish, Cambaroides Witch flounder, Glyptocephalus cynoglossus schrenckii Kessler (L.) 570 Hermit crab, Pagurus capilatus Smooth flounder, Liopsetta putnami (Gill) Mud shrimp, Upogebia issaeffi Balss. Winter flounder, Pseudopleuronectes americanus (Walbaum) Cumacea Atlantic lumpfish, Cyclopterus lumpus L. Macruran crabs Thorny skate, Raja radiata Donovan Smooth skate, Raja senta Garman Mollusks Skate, Raja sp. Mytilus edulis Linné Atlantic salmon, Salmo salar L. Lamellibranchiata Crustaceans Rhizopoda Shrimp, Pandalus montagui Nemertini Scud, amphipods Argis Copepods Cumacea Schizopodes Cephalopods Bathypolypus obesus (Verrilli) Atlantic shortfin squid, Шех illecebrosus (Le Sueur) Gastropods Whelk, Buccinum undatum Linné Periwinkle, Littorina sp. (?) Bivalves Crenella sp. (?) Cystodaria silgua Spengler Macoma baltica Linné Mesodesma arctata Conrad Mesodesma deaurata Turton Mesodesma sp. (?) Mya sp. (?) Yoldia limatula Say Polychaeta Cistendes gouldii Verrill Nereis virens Sars 572 769 Daily activity and behavior. The beluga is а typical herding animal, found usually in groups of various strengths. Among numerous obser- vations, lone animals were encountered in 16.1% of cases (Bel’kovich, 1960). A herd of belugas irrespective of its strength, invariably con- sists of small groups (Fig. 334) of two to four or five to eight animals (Golenchenko, 1949a). The age and sex composition of the herd can vary while the males within a herd often form independent groups. In other cases, mixed groups are formed in which young ones can also be seen along with mature animals. However, immature animals never form independent groups. In the period of reproduction, groups are noticed within a herd. These groups consist of three or more belugas, among which is a female with a suckling calf and one or two adult animals. The female in such groups probably had given birth only recently and was preparing for fertilization afresh. At the end of the reproduction period, adult males quite often segregate themselves from females and form independent herds. The herd size varies markedly. The most frequently noticed herds comprise tens (54.9% cases) or several tens of animals (25.8%). Herds running into a few hundred animals are rare (3.2%) but herds running into a few thousands of animals can be seen in the period of massive con- centrations of fish (salmon in the Far East). Small herds are invariably formed of animals chasing huge schools of fish. An analysis of hunting data shows that the male to female ratio in a herd is invariably close to 77. 2 CL 4 NOG ААА 72 LEE GLB. Ley a GEL РА, ZZ wy gilli LE gz Z РЕНА =. 2 > у os CELL ee ety bee LFA “Ss 222222, ЕН = 2 Е SS РНР, о р Se Аее. CELA В НК НА <> LL: LL 22% р = A; == ZA Fig. 334. A group of belugas at sea (figure by N.N. Kondakov). 573 572 770 one but differentiation into age- and sex-related groups is noticeable шо the large migrating herds. In such cases, the front of the herd consists exclusively of adult males followed by females with calves and immature animals (see Table 52). A varying composition of animals (age- and sex-related) is noticed in some parts of a large herd not only in the Far Eastern waters, but also in the European north along Novaya Zemlya and in the White Sea (Bel’kovich, 1960). The speed of belugas depends on the state and behavior of the herd. Feeding animals are confined to a relatively small region, dive in different directions, often at the same site, and remain under water for a long time. Such herds move in any direction at a speed of not more than 1 mile/hr. “Migrating” herds follow a distinct direction, are more compact, swim more rapidly, and dive for a very short duration. A migrating herd moving undisturbed travels at 2-5 miles/hr but a frightened herd can gather speed, perhaps up to 10 miles/hr; they, however, cannot sustain this high speed for long. At high speeds the females and calves begin to lag behind since, apparently, they cannot cover more than 7, or in an extreme case, 8 miles/hr. Sleeping belugas have been observed several times. They lie on the sea surface almost immobile, usually close to the coasts, passively drifting with the waves and the current (Arsen’ev, 1939; Bel’kovich, 1960; Kleinenberg et al., 1964). Table 52. Distribution of belugas in a herd (Gulf of Sakhalin) (Arsen’ev, 1939) Front of herd (44 belugas) White Blue Gray Suckling calves Male Female Total Male Female Total Male Female Total Male Female Total 22 6 28 4 4 8 1 3 4 2 1 3 65.1% 18.6% 9.3% 7.0% Total: males 29 (67.4%) and females 14 (32.6%). Rear of herd (23 beluges) White Blue Gray Suckling calves Male Female Total Male Female Total Male Female Total Male Female Total — 6 6 1 6 7 5 4 9 1 — 1 26.1% 30.4% 39.1% 4.4% Total: males 7 (30.4%) and females 16 (69.6%). 71 Belugas feeding predominantly on pelagic life usually do not sub- merge deeper than 8-10 m. In any case, a herd surrounded by a sweep net usually does not escape from the lower edge even when the height of the net does not exceed 5 m and falls short of reaching the bot- tom. It is quite possible, however, that should the need arise, the beluga could submerge to several tens of meters. During feeding, belugas dive for 3-5 min but can remain submerged up to 15 min. After relatively long submergences in search of food, they surface three or four times in a row. Migrating animals surface regularly at 20-40 sec intervals with inhalation/exhalation extending for about a second while the back of the animal can be seen above the water for about four seconds. While surfacing, the first to be seen above the water is the upper part of the head with the blowhole. The head is then submerged as the dorsum arces above the sea surface. This semicircle, moving around its own axis, gradually disappears under water. Caudal flukes are never displayed. The blow is rapid, like a small bush, and is seen like a white flash in sunny weather (V.A. Arsen’ev). The beluga does not avoid ice. In the north and the Far East, at places of its summer habitation, it is seen on days when the ice is broken and the animal remains free for the first time between ice chunks. In the north the beluga is common along the edges of ice and seen quite often in air holes (in the ice) and open water pools among drifting ice up to 9 to 10 points in density. In such cases, small herds are more common. The animals can overcome considerable masses of dense, apparently compact ice, past which they enter the open water in which their food is concen- trated. Such instances are common in the Shantarsk archipelago in the Sea of Okhotsk. Instances are known of encountering small groups of belugas in large air holes among stationary ice in which they sometimes even winter. The beluga does not avoid freshened or even fresh water. In the freshening waters of the estuaries of large rivers, belugas are regularly seen and are hunted. Almost all over the range, dozens of large and small rivers are known in which the animals regularly transgress chasing fish. In most cases, they rise up along the river for several tens of kilometers and are Seen quite often even at a distance of 100-200 km from the estuary. In some large rivers, these animals were sighted at a very long distance away from the estuary: in the Yenisey almost near Podkamennaya Tunguska at over 800 km, in the Amur at Khabarov at about 1,000 km, in the Pechora at 900 km, in the Ob’ at 1,500 km, and in the Yukon almost at 1,500 km. Vision and audition are well developed in the beluga but the former is effective only at a close distance. The animals can orientate themselves 574 UL in many Situations through their sense of hearing. They can perceive various sounds, including those produced in a very wide range by other belugas. These sounds are in the form of clicks and creaks and various types of whistles; they also resemble barking, roaring, and gnashing, and a sound somewhat similar to a trill. The sound frequency varies from a few kHz to 10,000 and even 20,000 kHz. Sounds play the role of var- ious signals and are used for echolocation. Hunters well know that if even one beluga finds an exit from the net surrounding the animals, the entire herd escapes through that very exit in a very short time. It is also known that the animals well perceive all the sounds produced not only in water, but also in air (on the coast or on the ice). Moreover, belugas can very well orientate themselves to sea-level changes (high and low tides) and are almost never cast on the coast (Arsen’ev, 1939; Tomilin, 1962; Kleinenberg et al., 1964). Experiments in raising the beluga in an aquarium have yielded pos- itive results. A male survived in an aquarium for several years and was taught during this time many “acts” of a “dolphin circus”. This, together with the external appeal of this white whale, was greatly appreciated by numerous visitors to the aquarium. Seasonal migrations and transgressions. Large regular migrations rep- resent a characteristic feature of the biology of the beluga. However, the periods and courses of migration of individual populations have been studied very schematically and opinions are often contradictory. White Sea. The beluga is generally seen at May end to early June after the sea becomes free from ice. In the northern portion, mainly along the west coasts of Kanin Peninsula, it has been sighted even earlier, early in April, but such instances are generally rare and the herds few. In summer the beluga is encountered regularly almost all along the entire coast of the sea; it is most numerous in the Dvina and Onega bay’s and along the west coast of Kanin. In August, the number of belugas begins to decrease and by October most of the herd goes into the Bar- ents Sea (Klumov, 1936; Provorov, 1957). Some belugas overwinter in the White Sea, as can be seen mainly in the Kandalaksha and Dvina bays and also in the air holes in Voronka and Gorlo regions. From the air individual animals or small groups were sighted in winter among dense drifting ice in the central part and in Voronka (Khuzin, 1960; Kleinen- berg et al., 1964). Thus the bulk of the White Sea beluga population inhabits the White Sea from early spring to late autumn, then enters the Barents Sea before its total freezing, where, apparently, they winter in its southeastern section. Spring migrations of animals commence as soon as the icy environment permits, sometimes even when a large part 575 773 of the sea is packed with dense drifting ice. In summer belugas are dis- tributed throughout the sea and transgress into the numerous gulfs and bays (Klumov, 1934; Provorov, 1957). Barents Sea.Asmall number of belugas is seen throughout the winter at different places along the Murman coast, which they probably abandon in March. The beluga remains in the region of Kolguev Island and the Chesha Bay throughout the year: in winter, in open water pools and along the edges of ice, moving toward the coasts in summer. In the region of Vaigach Island and along the west coast of the southern island of Novaya Zemlya, belugas are most numerous early in winter but, when the ice recedes, herds of belugas are seen close to Novaya Zemlya throughout winter and even in spring. Early in summer large- scale migrations of animals into the Kara Sea are noticed through the strait as also around the northern extremity of Novaya Zemlya. By mid- July, belugas have almost disappeared from the western coasts of these islands. In autumn large herds of belugas are once again seen here, now moving in westerly and southwesterly directions. At the end of June herds of belugas are seen in the Straits and bays of Franz Josef Land archipelago, where they remain until autumn but disappear again with the formation of an ice cover (Klumov, 1936a; Kleinenberg et al., 1964). Thus belugas arriving from the White and Kara seas winter in the Barents Sea. They live along the edges of drifting ice in the region of Kolguyev Island—Chesha Bay and also along the western coast of Novaya Zemlya depending on ice cover conditions. Small individual herds of belugas move toward the Murman coast. At the beginning of summer, most of the animals leave the Barents Sea for summer feeding in the Kara and White seas. Some small herds remain throughout the year in different regions of the sea (Chesha Bay—Kolguyev Island; southern part of Novaya Zemlya—Vaigach Island; Franz Josef Land archipelago); for wintering, they go beyond the edge of ice or inhabit open water pools. Kara Sea. Herds of belugas are seen in Kara Strait and in Yugorsk Shar Strait from the middle to the end-of May and even in early June and travel into Baydarata Bay. At this time the animals enter the Kara Sea and encircle Cape Zhelaniya (northern tip of Novaya Zemlya). In some years the periods of movement of belugas vary considerably depending on the ice situation; further, in some cases, belogas were sighted even in April. In June, in Amderma region (east of Yugorsk Shar Strait), large herds of belugas move eastward and some are seen along the southeast- ern coast of Novaya Zemlya throughout July. Belugas reach the estuary of the Gulf of Ob early in July. On entering the gulf, the animals gener- ally travel along the right bank since the prevailing northeastern winds 774 press the ice toward the left bank. Belugas return from the gulf mostly along the left bank or along the midsection of the bay. The Gulf of Ob, mainly at the point of its confluence with Tazovsk Inlet, belugas remain until the formation of an ice cover, i.e., until early November (Dukhovnyi, 1933, 1934; Chapskii, 1937; Kleinenberg et al., 1964). In the estuary of Yenisey Gulf, belugas are seen in the first few days of June (waters of Dickson Island) and even earlier in years of little snow. In most cases, herds of belugas arrive there from the east, from Pyasinsk Bay, and sometimes from the side of the open sea. Later, the animals remain there regularly and enter Yenisey Gulf and some- times rise high up along the Yenisey River. In the strait, belugas traverse mainly along the eastern coast. ш’ the September the number of animals decreases but individual herds are seen here throughout this months. In the second half of July and in August, belugas are common along the coasts of Pyasinsk Bay and are sighted quite regularly in August and September in Vil’kitskiy Strait and along the coasts of Severnaya Zemlya where en masse arrivals of arctic cod occur at this time (Heptner, 1930; Zikov, 1934; Klumov, 1936a; Kovalev, 1938; Kleinenberg, Bel’kovich, and Yablokov, 1960). The migrations of belugas in the Kara Sea can be sketched as fol- lows. In spring and early summer herds of belugas enter the Kara Sea through Novaya Zemlya Strait and around its northern extremity. The animals moving through the strait are seen in Baydarata Bay, later move eastward, and reach the Gulf of Ob by mid-July. By this time Matochkin Shar Strait has become ice-free and animals passing through this strait are seen along the eastern coasts of Novaya Zemlya from where, per- haps in part, they move also toward the southern coasts of the Kara Sea. The animals encircling Novaya Zemlya move eastward along the edges of drifting ice or along open water pools. Some move toward .Severnaya Zemlya islands and others reach the mainland coast somewhere north- east of Pyasinsk Bay. These belugas, moving southwest later, are seen in Pyasinsk Bay, around Dickson Island, and in Yenisey Gulf; however, the animals sometimes reach Dickson Island not from the northeast, but from the north. Later, this group of belugas disperses and the animals are sighted along the coasts of Taimyr and in Vil’kitskiy Strait. In August and September and in years of little snow even in October, herds of bel- ugas depart by the same routes (in a reverse direction) to the wintering sites. Therefore, the autumn courses of belugas in the estuary of Yenisey Gulf have a northeasterly direction and in the Gulf of Ob a westerly direction. Small herds of belugas winter in the Kara Sea, probably reg- ularly, in the air holes in the Gulf of Ob and Yenisey Gulf, and in the large air holes among drifting ice in various parts of the Kara Sea. 576 775 The periods of migrations of belugas in the Kara Sea depend ритаг- ily on the period and concentration of the arctic cod, which represents the main food of belugas. Many major rivers enter the Kara Sea and bring with them masses of warm water in spring and summer. These waters, tich in biogenic elements, promote the growth of phyto- and zooplankton and thus provide extremely favorable conditions for the habitation of the arctic cod, in search of which herds of belugas follow. Thus the periods and magnitude of arrival and distribution of belugas depend to some extent on the magnitude and distribution of river waters in the Kara Sea (Tarasevich, 1960a, 1960b). Laptev Sea. The distribution and migrations of belugas in this sea are not quite clear. Herds of belugas (sometimes large ones) have been sighted in Vil’kitskiy Strait, along Cape Chelyuskin, in Pron- chishcheva Bay, in Khatanga Gulf, in the estuaries of the Anabar and Olenek rivers, in the Lena delta, near Tiksi and, what is more, belu- gas have been sighted every day at some of these points. Thus the ani- mals inhabits the entire southwestern coast of the Laptev Sea and some- times herds of belugas are seen in July to September along the west coasts of Lyakhovsk and Kotel’nyi Islands (Klumov, 1936a; Kleinenberg, Bel’kovich, and Yablokov, 1960). In all probability, an isolated but comparatively small population of belugas resides in the Laptev Sea and spends the winter in the large open water pools and air holes of this shallow water basin. In the sum- mer months the animals inhabit the southern and western coasts of the sea including Vil’kitskiy Strait, Severnaya Zemlya coast, and possibly the adjoining Kara Sea waters. During their habitation in the waters of Novosibirsk Islands, belugas are sometimes seen in adjoining regions of the East Siberian Sea. Late in autumn the animals leave the coasts Юг’ the central part of the sea where they winter. © Sea of Okhotsk. The maximum collection of belugas in summer is seen in the western (Gulf of Sakhalin, Amur liman, and Shantarsk archipelago) and northeastern (Shelikhov Gulf with Penzhina and Gizhaga bays) parts of the sea. In the northwestern part of the sea, from Shantarsk Island to the region of Ayan-Okhotsk and along the western coast of Kamchatka, belugas are encountered in small numbers. They have not been noticed in the central deep water portion of the sea. Probably, two local herds of belugas spend the summer in the Sea of Okhotsk and could be called Amur and Penzhina herds (Arsen’ev, 1939). The earliest appearance of large beluga herds in the southwestern part of the Sea of Okhotsk (evidently in April) was recorded in Terpeniya Bay along the southeastern coast of Sakhalin (Polyakov, 1884). At the S77 776 end of April to early May, small numbers of animals arrive in Tatar Strait where they remain on the edges of stationary ice in the northern section of the strait. Early in May many belugas are sighted in the narrow part of Tatar Strait (Pogibi-Lazarev) and the animals approach the coasts as soon as the ice breaks up along Sakhalin. As the strait becomes clear, belugas gradually move northward into the Amur estuary but are con- fined to the narrow part of the strait generally up to mid-July, leaving only after the fish (herring and humpback salmon) disappear there. The animals then move into the Amur estuary and into Sakhalin Strait. Concurrently, at May end to early June, large herds of belugas approach from the southeast to the northern extremity of Sakhalin and, encircling the island from the north, reach the Gulf of Sakhalin, most of which at this time is still covered by compact ice. On the Sakhalin coast of the bay, ice begins to break in the first half of June and herds of belugas are seen immediately thereafter in the Gulf of Sakhalin and later in the Amur estuary. These animals live there until late autumn. Roughly at the same time (or slightly later), belugas are seen in the bays of the Shantarsk Sea while the sea itself and partly the large gulfs are still ice-bound. However, the dense drifting ice is no obstacle to the belugas moving in the upper courses of the bays which are free of ice (Tugursk, Ulbansk, and so оп). The migrations of Amur belugas can be sketched as follows. In April- May, herds of belugas appear in Terpeniya Bay and later a part of this herd moves through La Perouse Strait into Tatar Strait and feeds in its northern section on herring and humpback salmon. By mid-July, belugas ‘leave for the Amur estuary and the Gulf of Sakhalin. Almost concur- rently, at the end of May, most of the herds approach the northern extremity of Sakhalin from the southeast, encircle Sakhalin from the north, enter the Gulf of Sakhalin, and later Amur estuary in the first ten days of June. Here the animals feed throughout the summer. Roughly at the same time, a part of the herd from the northern extremity of Sakhalin turns not into the Gulf of Sakhalin, but moves west and arrives in the large bays of the Shantarsk Sea over large masses of dense ice. There they are encountered throughout July, August, and September. Individ- ual small herds may arrive from the Shantarsk Sea along the coasts in the northeast and are sometimes seen in the region of Ayan Bay. Some animals may even reach the estuaries of Okhota and Kukhtui rivers. The nature of migrations of belugas changes in autumn. Instead of large compact herds, small isolated groups scattered over a large water body are seen. With the formation of a compact ice cover at the end of October - November, belugas abandon points of summer habitation. A small part of the herd arrives October end or early November from the НИ north into Tatar Strait and apparently exits through La Perouse Strait into the Sea of Okhotsk. Most of the herd from the Shantarsk Sea and the Gulf of Sakhalin encircle Sakhalin from the north at October end and in November and head southeasterly. During autumn migrations very large herds are not seen. The periods of migrations cover a long time frame. Probably, with the disappearance of schooling fish, some animals begin to leave the summer feeding waters even before ice formation; formation of dense or stationary ice cover then compels the remaining animals to leave. The wintering sites of belugas have not been established (Dorofeev and Klumov, 1936a; Arsen’ev, 1939). According to the latest observations, groups of belugas have been recorded north of Sakhalin in winter in air holes among dense drifting ice (G.A. Fedoseev). It is possible that some herds spend winter amidst ice in the midpart of the Sea of Okhotsk. In Penzhina and Gizhaga bays of Shelikhov Gulf, belugas are seen close to the coasts at the end of May immediately after the thawing of coastal fast ice in the bays and gulfs. The animals remain there through- out summer feeding on schooling fish (herring and salmon). They per- form regular local migrations, their periods and directions depending on the powerful low and high tides (at some points during low tide, the coast is exposed for a distance of 5 km or more). Belugas inhabiting Shelikhov Gulf reach west up to Tauy Bay and some animals or small groups have been encountered even more westward, right up to Okhotsk. A few animals are encountered in the summer months along the west coast of Kamchatka (predominantly in its northern section). With the appearance of floating anchor ice and the subsequent freezing of coastal waters in October-November, belugas leave the coasts and are no longer seen in these waters in the winter months (Arsen’ev, 1939). The points of wintering of this herd of belugas and the courses of their migrations have not been established. The winter ice cover of Shelikhov Gulf is typical. Thus in Gizhaga Bay coastal shore ice 20-25 miles wide is formed while the midpart of the gulf is covered with drifting ice. In Penzhina Bay only small straits freeze while the entire remaining water body is covered by floating ice of varying density. It is quite possible that herds of belu- gas may winter in the regions of sparse ice without undertaking distant migrations. There are, however, no observations on winter movements. Migrations of belugas in the Bering and Chukchi seas have been very poorly studied. In summer the animals concentrate in the Gulf of Anadyr and the Anadyr estuary. Many schooling fish entering this river serve as food for the belugas and hence beluga arrivals are often en masse. Belugas are seen scattered throughout this bay in July. At about this time 578 also, belugas are seen at many points in the Chukchi Sea: Cape Serdtse 778 Kamen’, Kolyuchin Inlet, Cape Shmidt, and probably somewhat later оп Wrangel Island and north of it (up to 72 to 73° М lat.). Simultaneously they are also seen on the coasts of Alaska where they are known from Bristol Bay to at least Cape Barrow. In Bering Strait, spring (April- May) migrations of belugas occur in the north while autumn migrations (September-October) are observed in the south (Arsen’ev, 1939; Nikulin, 1946). The migrations of belugas in these waters can tentatively be described as follows. The animals winter in the eastern and northeastern parts of the Bering Sea. In April-May, with the disappearance of ice, they begin to move north and northwest. Some belugas exit into the Chukchi Sea and are distributed over an immense water body, from the boundaries of the East Siberian Sea to Beaufort Sea and Wrangel Island, depend- ing on the disposition and density of ice. Other, probably more, animals remain at the inlet into the Gulf of Anadyr and with thawing of ice, are scattered over this water body, where they spend the summer months. In autumn (October-November), with the compaction of ice and the for- mation of young ice, animals from the Chukchi Sea move southward, cross Bering Strait, and leave for their wintering sites. The ice displaces the belugas from the Gulf of Anadyr also, compelling them to move east and southeast. In most cases, the dispersal of migrating marine mammals (whales, belugas, and walruses) in the Chukchi Sea is restricted in the west by the line Cape Shmidt—Wrangel Island since De Long Strait almost year- round is covered by a broad strip of very dense ice (Arsen’ev, 1935). On encountering this massive ice obstacle, the animals change course northward toward Wrangel Island and travel farther west into the waters of the East Siberian Sea only in years of little ice. In winter months belugas are noticed at many points in the Bering and Chukchi seas, where they remain in large air holes and open water pools formed among the floating ice. Wintering belugas are noticed almost every year close to Provideniya Bay (northeastern part of the Gulf of Anadyr); animals are sighted at different points along the coast of the Chukchi Sea: Nunyamo, Lesovsk, Dezhnev, Serdtse-Kamen’, and other capes (Nikulin, 1951; Uspenskii, 1958; Kleinenberg er al., 1964). In the Beaufort Sea, belugas are encountered in summer in Amund- sen Gulf and McClure Strait, to the west of Banks Island, at many points in the MacKenzie delta, and to the west of the estuary of this river. It has been suggested that, in the Beaufort Sea, belugas even winter far from the coasts since they are seen in spring concurrently at many points (Vladykov, 1944; Sergeant, 1962a). 579 dag In the eastern part of the Canadian Arctic, belugas similarly under- take seasonal migrations, which are most distinctly manifest in Davis Strait and Baffin Bay. In summer they sometimes reach 80°N lat. (Hall Basin), descending in winter to the middle of Davis Strait: Cumber- land Sound—Sukkertoppen. Regular migrations in spring and autumn are noticed in Hudson Strait but a small number of belugas winters in the western and northern sections of Hudson Strait, eastern part of Cumberland Sound, and in Lancaster and Jones sounds. In the Gulf of St. Lawrence, as it frees from ice, belugas are seen in the estuary of the river (often transgressing into the river) and at other points on the coast but abandon these regions in autumn. The population living east of Greenland probably overwinters to the north and northwest of Iceland. In spring the animals migrate northward, reaching Spitsbergen, often in large numbers, and return to the wintering sites in autumn (Vladykov, 1944; Daan and Douglas, 1953; Sergeant, 1962a). Transgressions of belugas beyond the limits of their normal range are not very rare. In the Bering Sea, belugas were noticed in the waters of Karagin (58°N lat.) and Commander Islands (54°N lat.) (Tomilin, 1957) although 46° М lat. serves roughly as the southern boundary of the distribution of belugas in the Sea of Okhotsk. From the Chukchi Sea some animals or small groups reach the estuary of the Kolyma and the central part of the East Siberian Sea, although they may reach there later from the Novosibirsk Islands. Animals of the two populations may be encountered there. Transgressions into the Atlantic Ocean usually occur in particularly cold winters. In the western part of the North Atlantic, the southernmost transgression was noticed in Massachusetts Bay (42°N lat.) and in the eastern part along the coasts of Great Britain, Holland, Schleswig coasts in the Baltic Sea, Estonia, Finland, and even France (von den Brin, 1958; Kleinenberg e7 al., 1964). During 1964 to 1966, eleven sightings of belugas were recorded in the waters of Central Europe. The well-known case of the long residence of a beluga in the Rhine is particularly interesting. Close to Nijmegen town, Holland, a beluga was sighted on May 18, 1966. Gradually, ascending against the current, it crossed Deventer, Kampen, Duisburg, Dusseldorf, and Bonn, and on June 13 reached Honnef, which is 400 km from the sea (50° 40/S lat.). Here the beluga turned back and entered the sea on June 18 after moving downstream for 400 km in five days. Attempts to catch it in a net or by roping or using an injector gun were not successful. This adult beluga with a body length (visual) of 5 to 6 m spent a month in fresh water, which is rather unusual (Slijper, 1967). 580 780 Reproduction. ш the Sea of Okhotsk mating of belugas was recorded in April-May (Nikol’skii, 1936); in the Gulf of Ob in July (Zaikov, 1934); in the Barents and Kara seas from May through August, peaking in May- early July (Kleinenberg and Yablokov, 1960); in the Gulf of St. Lawrence from February to August with a large number of animals mating in May and June (Vladykov, 1944); and in Hudson Bay from March through September, peaking in May (Daan and Douglas, 1953). Thus the period of mating extends for about six months but the majority of females are fertilized over a relatively short duration, 1.е., from April end to early May. Only some individual animals mate at other times. In the White Sea, calves with the umbilical cord intact were encoun- tered in mid-July (Provorov, 1957) in the Gulf of Ob (Zaikov, 1934), in Hudson Bay (Daan and Douglas, 1953) on August 15, and in the Sea of Okhotsk from July 18 to 26 (Kleinenberg et al., 1964). However, judging from the dimensions of the suckling calves caught in the summer months, their births may have taken place much earlier (Kleinenberg et al., 1964). In all probability, the period of parturition, like the period of mating, is protracted and births may occur from early spring throughout the sum- mer months. The period of maximum births has not been ascertained. Thus gestation in belugas extends for 11-12 months and, according to one view, might be extended 13-14 months (Sergeant, 1962a). Information on the duration of lactation is quite contradictory, from 5-6 months to 12 months or even longer. Until the end of September (close of the hunting season and specimen collection), calves were found feeding exclusively on milk. In the stomach of animals of the next color stage, i.e., gray belugas, milk was also found together with food gathered independently, but the age of these animals could not be established. It would probably be more correct to assume that the period of lactation extends for six months (Kleinenberg et al., 1964). At the same time, the teeth of young belugas emerge roughly at 10 months of age and hence it is possible that milk suckling of the calves ceases in this period. The female usually delivers a single calf although, as among other cetaceans, twins are sometimes encountered. One female contained three embryos (two males and one female). It has been suggested that female belugas are capable of mating at a short interval (two to four weeks) after parturition. This suggests the possibility of annual births but investiga- tions carried out on various beluga populations have shown that some percentage of mature females do remain barren every season. The ratio between barren and gestating females in a herd varies considerably in different years as also in different regions of habitation. It has not been possible to establish the pattern of barrenness and hence the reproduc- tion cycle of this animal can only be surmised. In some cases, it appears 781 probable that а female may give birth for five to six or more consecutive years, after which she rests for a year (or more). In other cases, such rest periods might set in after two or three years (Tomilin, 1962; Kleinen- berg et al., 1964). It is also possible that the reproduction cycle varies for females of different age groups. Therefore, the tempo of population replenishment has yet to be established and recommendations for the rational commercial exploitation of the beluga are difficult to make. In the period of mating it was sometimes observed that two or more males chased a single female while groups of 10-20 adult males, isolated from mixed groups, were encountered concurrently. This suggests the possibility of polygamy among belugas (Sergeant, 1962а). Growth and development. Measurements of beluga embryos from Greenland waters showed that in the period of embryonal growth, devel- opment is relatively uniform (Table 53). Since the period of births is very prolonged, embryos of different size may be encountered at any time of the year (Fig. 335). Thus in the northern seas of the Soviet Union, the length of measured embryos varied from 7 to 168 cm, the embryos investigated in July being larger than those found in August and September (Kleinenberg and Yablokov, 1960). The maximum dimensions of embryos differ in different regions of the species’ range. In the Kara Sea, the largest embryo was 150 cm long, in Hudson Bay 151 cm, Barents Sea 170 cm, Sea of Okhotsk 175 cm, waters of western Greenland 183 cm, and in the Gulf of St. Lawrence 213 cm. Newborn calves are often smaller than the larger embryos. Thus the length of the smallest calves (from among those measured) was (in cm): in western Greenland 116; Sea of Okhotsk 130; Kara Sea 140; Bar- ents Sea 147; Gulf of St. Lawrence 147; and Hudson Bay 153. The average body length of a newborn calf, covering all the regions, is assumed to be 150 cm (Kleinenberg et al., 1964). The growth rate of belugas is the highest in their first year. The average monthly increment has been put at 30 cm; in the lactation period Table 53. Body length of beluga embryos, cm (Nielson and Dagerbol, 1930; after Tomilin, 1957) Month Number of © Body length Average specimens November 36 32-78 60.2 December 40 46-104 87.3 January 11 88-112 96.3 February 19 122-142 129.9 March 19 113-183 142.2 April 6 142-158 149.1 782 Fig. 335. Embryo of beluga (figure Бу М.М. Kondakov). the increment should be more than after changing over to independent feeding. By the end of the first year the animal reaches a length of almost 300 cm. The dimensions of belugas at the age of up to one year from the Sea of Okhotsk varied from 140 to 280 cm, the peak of the growth graph falling in the 180-200 cm interval. Further growth of belugas is identified from the color changes of the 581 animals in three stages. Animals of the first color group, called “gray” animals, have a bluish-gray body and are immature. The next color group, the “blue” group, covers animals transiting from gray to white. Mature as well as immature animals of both sexes are encountered in this group. Finally, the “white” group comprises adult mature animals of various sizes with a pure white body color. The age of transition from one color group to another has not been established. It has been assumed that the average annual increment in the second and third years is 75 cm and in the fourth year about 45 cm among males and about 30 cm among females (Sergeant, 1959a). The males soon outgrow females and hence the average size of adult males is more than that of females (Table 54). Age determinations of females based on dentine layers in the teeth showed that the immature group comprised animals of three to four years (body length 202-317 cm) but females aged two to five years could be. placed in the group of potentially mature animals. The minimum age of mature females is three years but the maximum number of northern bel- ugas mature at age four to five years or more (body length 315 - 435 cm). Among mature belugas, the growth rate decreases sharply. In the first year after sexual maturity, the female adds an average of 20 cm, in the second year 5-7 cm, and in the third year 2-3 cm. In the fourth to the fifth years of maturity, growth ceases and the animals attain physical maturity. By then, they measure 355 -360 cm in body length and fall in the age group seven to nine years. The body length of females in the Far East in the first year of maturity rose by 28 cm, in the second and third years by 7-9 cm, and in the fourth and fifth years by 2-4 cm. Their 582 783 Table 54. Body dimensions of belugas of different color groups, cm (Kleinenberg ef al., 1964) Region Color Males Females ge) ee ИЕ ВНИИ. ИЕ ВЕЧЕ Number of Body Average Number of Body Average specimens length specimens length Gulf of St. Lawrence Gray 8 226-263 247 13 203-282, 239 Вше 12 282-345 311 18 282-343 318 White 59 329-447 406 38 310-409 372 Kara Sea Gray 1) 239-340 276 15 224-341 272 Вше 20 300-393 336 17 274-365 322 White 35 343-472 400 26 326-390. 353 Sea of Okhotsk Gray 34 195-310 252 28 170-357 286 Blue 173 187-479 351 98 251-422 18531 White 136 359-500 447 53 360-418 388 physical maturity set in in the fifth to the sixth year after sexual maturity. The average body length of physically mature females in the Far East was 390 cm (Kleinenberg её al., 1964). Male belugas from Hudson Bay attain sexual maturity at six to nine years (12-18 dentine layers) at a body length of 275 -320 cm and females at six years (12 layers) at a body length of 275 cm. Belugas on the eve of the “white” stage have 13 to 18 dentine layers. Up to 35 layers are seen in very old animals. The maximum number of layers counted is 50 (Sergeant, 1959a). Longevity of the belugas has not been established. Enemies, diseases, parasites, mortality, and competitors. The killer whale and the white polar bear may be regarded as enemies of the beluga albeit they inflict little damage on the population. Apart from diseases caused by parasites, no other has been recorded. Ectoparasites are not known. Thirteen species of helminths are known among belugas: trematodes three, cestodes one, nematodes eight, and acanthocephalans one. None of the three species of trematodes has been reported in any other ani- mal. The trematode Odneriella seymouri Price was found in the Bering, Okhotsk, White, and Barents seas. Leucasiella arctica Delamure and Kleinenberg, parasitizing the rectum, has been detected in the Barents Sea. Leucassiella mironovi Krotov and Delamure was found in the small intestine of belugas from Aniv Bay and La Perouse Strait. The cestode Diphyllobothrium lanceolatum Krabbe, reported before for three or four species of pinnipeds and possibly common porpoises, is evidently a facultative parasite of the beluga. Four of the eight known species of nematodes have been exclusively reported in the beluga. Anisakis (Anisakis) kukenthalu Cobb, parasitizing 784 the stomach, was found in the Arctic Ocean (Spitsbergen), the Barents, Kara, North, and Okhotsk seas, and in Aniva Bay (Sakhalin). Sztenu- rus arctomarinus Delamure and Kleinenberg was detected in the lungs of belugas from the northern Arctic Ocean, and White and Barents seas. Otophocaenurus oserskoi Skrjabin, localizing in the auditory organs, was found among belugas from the waters of Sakhalin, northern Arc- tic Ocean, and White and Barents seas. Stenurus pallasii van Beneden, parasitizing the auditory organs, bronchi, and circulatory system, was detected in the Arctic. Anisakis (Anisakis) simplex Rudolphi, very widely distributed among marine mammals, was found in the gullet, stomach, and intestine of belugas and also in ten other species of toothed whales, two species of baleen whales, and Steller’s sea lion. It is known from various places in the North Sea, eastern Kamchatka, Japan, and New Zealand. Terranova decipiens Krabbe, parasitizing the stomach and intes- tine, has been detected in the beluga only once (Kara Sea), whereas it occurs in 12 species of pinnipeds, 2 species of toothed whales, and 2 species of baleen whales in the North Sea, the Atlantic and Pacific oceans, and in Antarctica. The nematode Stenurus minor Kuhn, found among belugas in the northern Arctic Ocean, North Atlantic, North, Black, and Azov seas, and along the Asian coast of the Pacific Ocean, parasitizes the auditory organs, bronchi, heart, and blood vessels of only common porpoises in addition to the beluga. Finally, Crassicauda gili- akiana Skrjabin and Andreeva parasitizes the same organs of the beluga and common porpoise. It has been detected in the Amur estuary and in the waters of Kuril Islands. The lone species of acanthocephalan parasitizing the intestine of beluga, Corynosoma strumosum Rudolphi, is known in 11 other species and subspecies of pinnipeds, common porpoises, the sea otter, and also land carnivores and ichthyophagous birds. Among marine mammals, this helminth has been found along the Atlantic coast of Europe, in the Baltic Sea, Lake Ladoga, along the Canadian coasts, in almost all the northern waters of the USSR, around the Kuril Islands, in the waters of Sakhalin, along the coasts of California, and in the Caspian Sea (Delamure, 1955; Tomilin, 1962). Population dynamics. Changes of beluga populations relative to nat- ural factors are not known. The comparatively low scale of hunting per- haps does not influence the numbers of this species. Field characteristics. The pure white color of adult animals distin- guishes this species from all other marine mammals. Further, the beluga has no dorsal fin. The corpus adiposum [melon] is prominent, reaching almost the end of the broad snout; a “beak” is absent. The teeth are conical and develop in both jaws; canines are absent. In all the color 583 785 groups, coloration is monochromatic. Belugas usually stay together in groups which gather into herds of various strengths. In the period of en masse arrival of fish which serve as their food, belugas form herds of thousands of animals. This species is essentially a coastal form but is also encountered amidst ice at times of high density. (V.A.) Economic Importance Although beluga hunting has been carried out for several centuries in our country, it has never been extensive. In the 1920s to the 1930s, beluga hunting in the seas of the Soviet north and Far East expanded but was not intensive. In the years of maximum catch (1931-1935), the number caught in the Soviet Union did not excede 5,000. The maximum catch occurred in the Far East up to 1940 but thereafter dropped to a few hundreds per year. After 1950, hunting intensified somewhat in the White, Barents, and Kara seas, but nonetheless the annual catch has never exceeded a couple thousand animals. Outside the limits of our waters beluga hunting is also not significant. Formerly, the most productive area was the Gulf of St. Lawrence where some 2,000 belugas were caught annually, dropping to a few hundreds in 1932 - 1938; hunting ceased altogether in 1940. In Hudson Bay, from 1948-1960 more than 1,000 belugas were caught only in three years, the catch varying from 600 to 900 in the other years. In the waters of Greenland and Spitsbergen, the annual catch used to vary widely, from a few tens to several hundreds. In the rest of the Canadian Arctic regions, the catch was invariably low. Statistics of beluga hunting are far from perfect and the figures cited should be regarded as highly tentative. Nevertheless, the catch of beluga throughout its range has never been much and hence the products of beluga hunting have never been of commercial or economic importance. The techniques of catching the beluga are diverse and specific to each region. The oldest equipment for catching the beluga in the Euro- pean north is the White Sea sweep net whose length depends on the capability of the cooperative (number of operators and karbases* avail- able) and may go up to almost 2,500 m./In principle, this is a dynamic method of catching but is extremely primitive in practice. A composite net 2,400 m long is cast in the open sea, not farther than 7 to 8 km from the coast; the operation involves 24 boats and 170 people. The depth of casting should not exceed the wall height of the sweep net. Two sec- tions each of the net set up on two boats are stitched together before setting out to catch and these boats enter the sea in pairs. Such pairs are * Large boats with high sides used in marine and river transport—Translator. 584 786 deployed at some distance from each other in a semicircle open on the side of the anticipated approach of the herd of belugas. As the animals enter this semicircle, each pair of boats begins to diverge in the opposite direction, casting the sweep net into the water and joining it with the successive pairs to ultimately form a circle with the sweep net. This circle is gradually closed, its diameter reduced, and the boats enter the circle of the sweep net. The belugas are killed from the boats by shooting or battering with ice chisels. The killed animals are towed to the coast and dressed. The production cost of hunting by this method is extremely high (Klumov, 1936b, 1939). Another, much older piece of equipment used in catching is the Nor- wegian stationary net. Its wall height varies from 8 to 14 m and length 450 to 1,200 m. The net is set up only at places of regular arrivals of belugas at an angle of 30 to 35° to the coastline. A brigade of catch- ers consists of 8 to 12 persons with some boats. The men in the boats approach the beluga herds from the sea side and attempt to chase the animals into the net inlet by raising a racket, after which the inlet is sealed by a net. The belugas trapped within the net are battered with ice chisels and dragged to the coast (Heptner, 1930). The fabric mesh or so-called “polovinka” [meaning half] is similarly passé. Its length is 25-50 m, height 8-12 m, and mesh size 50-60 cm. These nets are set up on anchors perpendicular to the coast in a check- ered pattern at a distance of not less than 15 m from each other. The lower seine rope should reach the bottom. A team of five persons can operate 30 to 40 such “рооушкКаз”. The animal passes through the mesh but its caudal fin becomes entangled and the net twists around it. With no possibility of floating, the animal succumbs to asphyxiation. Best results are obtained on dark autumn nights when the belugas cannot see the net (Klumov, 1939). These traps have been modernized in recent years, with plant fiber replaced by capron [nylon] or flexible galvanized steel cables, which are less visible in the water. Hunting is done in the open sea from a ship (in the region of Novaya Zemlya and in Yenisey Strait), which chases a herd of belugas into the net. Such innovations have significantly increased the catch in such traps (Butorin, 1957). The development of beluga hunting in the Far East took a different course. The prototype of the beluga net here was an ordinary casting net used for catching salmon. Special coastal nets of this type, up to 1,000 m or more in length and made of thick strong material, were used for catching belugas. The ends of the net, laid on a kungas,* were affixed to the coast. As a herd approached, the boat with the kungas in tow * Open fishing boat with a carrying capacity of 3-5 tons—General Editor. 787 surrounded it and drew the second end of the пе! to the coast. The net was then hauled in manually or with a winch to the coast and the captured belugas gradually removed, killed, and dressed (Fig. 336). Later, this coastal net was elongated to 2,500 m and was gathered on two kungases standing together with two boats on anchors more than 1 km from the coast. As a beluga approached, the kungases were towed simultaneously in the opposite direction which hastened trapping of the animal and enlarged the area of catch. Both ends of the net were drawn to the coast and its further hauling in and removal of the animals done as in the coastal method of catching. In the course of further modernization (I.K. Nepomnyashchii), a deep net was designed with a length of about 1,000 m and a wall height of 18 m. Using two boats and one kungas, a herd of belugas was surrounded comparatively far from the coast, the net with the caught belugas was towed to the coast, and the trapped animals were recovered from the large net by the conventional method. This did not make for venturing beyond 10 km from the coast, towing took several hours, and if the weather turned foul, there was danger of losing not only the catch, but the net itself. A new net 1,600 m in length with a wall height of 18 m was subsequently designed. It was cast from two motor boats without a kungas. On sighting a herd of belugas, these boats diverged in the Fig. 336. Caught belugas laid out on the coast. Sakhalin, 1938 (photograph by V. A. Arsen’ev). 585 788 opposite direction and surrounded the herd with the net. After this, much of the net was gathered in so that a small circular section remained under the water. The boats, traveling in the opposite direction, gradually tautened and transformed this circular section into a line. The belugas caught in the net died of asphyxiation, were dragged to the approaching kungas, and hauled to the coast (Fig. 337). The team of catchers using these nets consisted of 25 to 30 persons (Arsen’ev, 1940). At places with a high variation in sea level during low and high tides, the belugas trapped in the net were left until low tide set in and then dragged to the coast for killing and dressing. The coastal people sometimes shoot belugas with rifles but many of the killed animals are lost. Sometimes the beluga is first harpooned manually and then shot. This method was in vogue for sometime in the Gulf of St. Lawrance and other regions of the Canadian Arctic. At some places the belugas are chased into narrow straits and the exit sealed off by a net. In most cases, these primitive methods of hunting result in a low catch. In dressing the killed belugas, the subcutaneous fat layer together with the skin is removed first. The fat layer is later separated from the skin and oil obtained from the blubber. The skin, now free of fat, is salted and after a few days, the thick layer of epidermis recovered. The skin is again salted and, after sometime, packed in barrels for dispatch Fig. 337. Towing the trapped belugas. Sakhalin, 1938 (photograph by V.A. Arsen’ev). 586 587 586 789 to leather manufacturers. The remaining carcass is processed into edible parts and those to be used in fertilizer meal. Where there are no process- ing facilities, the carcass is used as feed for fur-bearing animals. The oil melted from the blubber is used for commercial purposes. The oil from the head (corpus adiposum [melon]) and from the hollow of the lower jaw is collected separately. This oil possesses the property of remaining fluid at low temperatures and is used a special lubricant. The skin of the beluga can be used in making shoe soles and other footwear articles as well as halters and reins. Further, the intestine of the beluga can be satis- factorily used in the sausage industry; 30 m long, the intestine lengthens three times over after removal of the muscular layer. Blood accounts for roughly 5% of the overall weight of the animals. The valuable product, black albumin, can be produced from the blood (Dorofeev and Arsen’ev, 1936; Druker and Gakichko, 1936). The average weight of males and females of different color groups of belugas are given in Table 55. Table 56 gives the weight of the body parts of some animals. The ratio of the weight of different body parts of belugas without classification into age (color) groups is shown in Table 57 (those sections of the carcass used commercially were weighed). The beluga reserves in all the regions of its extensive range are not as large as often assumed and hence their catch should be based on a knowledge of the biology and population of these animals. Beluga hunt- ing has been restricted in recent years but could be enlarged without affecting these reserves. This could be done first in the Far East seas where not more than a thousand belugas have been caught annually since the 1940s. In the northern seas the increase should be more cau- tiously approached since hunting here is already vigorous although the possibilities of commercial exploitation of the animal population have not been fully realized. It should not be forgotten, however, that in Hud- son Bay, for example, some restrictions have been imposed on hunting © since even the comparatively low catch over several years has reduced Table 55. Average weight of whole animals from Sakhalin Bay, kg (Arsen’ev, 1935b) Sex Color group White Blue Gray Sucklings Total No. Av.wt. No. Av.wt. №. Ау. м. №. Av.wt. №. Av. wt. Males 120 — 880.1 3811 489.4: 21 310.4 2 150.0 181 726.2 Females 81 6566 31 506.1 Zi 28184 4 224 1432 53726 ? 7 — 3 — 63 == 2 — 15 — Total DOSE SOO een. 494.35 111 266.5 8 122.9, 399. 2577.4 586 587 790 Table 56. Weight of body parts of belugas from Sakhalin Bay, kg (Druker and Gakichko, 1936) Body part Color group of belugas White Blue Gray 1 2 3 1 2 a ani Skin: Epidermis (‘“‘jacket’’) 34.5 aan 28.2 26.5 24.6 14.2 Dermis 18.5 57.2 27.3 13.9 11.8 9.1 Fat of: Trunk 240.8 151.5 294.3 146.6 114.5 66.6 Неаа 4.1 TES) 10.9 4.1 2.8 1.4 Jaw 0.8 0.5 0.3 0.6 0.6 0.2 Flesh 242.2 184.4 217.4 208.6 155.7 58.6 Bones 68.2 ТР 86.0 65.3 37.3 25.3 Flippers 8.1 9.1 9:7 9.1 52 32 Caudal flukes 8.9 9.0 12.5 9.5 5.2 3.5 Viscera: Liver, heart, lungs, 723 49.0 65.6 lel 35.5 15S) kidneys, and stomach Intestine 22.6 16.0 18.1 19.0 11.5 5.5 Table 57. Weight of carcass sections of belugas from Sakhalin Bay, kg (Dorofeev and Arsen’ev, 1936) Number Av. м. Wt. of skin Wt. of carcass Wt. of skin Wt. of skin of of one with fat and without viscera = and epidermis without belugas animal epidermis without fat epidermis Ау. wt. Yof Av.wt. %of Av.wt. %of Av.wt. Hof body body body body wt. wt. wt. wt. 66 509.4 262.5 51.5 246.9 48.5 52.3 10.3 22.5 8.6 the beluga reserves. Investigations should continue for a more precise determination of numbers in each population. (V.A.) Genus of Narwhals (Narwhals or Unicorns) Genus Monodon Linnaeus, 1758 1758. Monodon. Linnaeus. Syst. Nat., ed. X, I, p. 75. Monodon monoceros Linnaeus, 1758. 1772. Ceratodon. Brunnich. Zoologiae fundamenta, р. 48. Substituted for Monodon Linnaeus, 1758. 1804. Narvalus. Lacépéde. Hist. Nat. Cétacées, p. 37. Narwalus vulgaris Lacépéde = Monodon monoceros Linnaeus, 1758. (V.H.) 588 791 Dimensions are the largest in the family, the body length reaching 610 cm. Compared to the beluga, the neck is poorly developed. The head is obtuse and spherical. The mouth section is small. The body of young animals is monochromatic, dark (bluish-gray, slaty, or slaty-blue) but in adults light-colored with numerous dark spots of irregular shape not exceeding 5 cm in diameter on the head, dorsum, and flanks. The spots on the upper parts of the head, neck, and caudal stem sometimes merge to form an overall dark background. The skull structure in general is the same as in the genus Delphi- napterus but asymmetry is more sharply manifest. Teeth are lacking in the lower jaw. The upper jaw has only one tooth in each half (in females, they usually do not emerge). The left tooth in males is transformed into a spiraled screwlike tusk reaching 3 m in length. The right tooth, how- ever, is more often concealed within the gum. Sometimes two tusks grow in males. From time to time, tusks are seen among females too. Verte- brae 50 to 55; cervical vertebrae not fused. Phalangeal formula: I, _,, II; _ 9, Ш4-в, [V>-4, and V,_-,. Compared to the beluga, the sternum of narwhals is short and broad. Narwhals seek food in the pelagic part of the sea. They feed mainly on cephalopods (teuthophagous) but also consume fish, possibly crus- taceans as well. The periods of mating and parturition are protracted. Narwhals are distributed in the waters of the Arctic and North Atlantic. There is no special hunting for narwhals. The genus comprises a single species: the narwhal or unicorn, М. monoceros Linnaeus, 1758. (V.S.) NARWHAL OR UNICORN Monodon monoceros Linnaeus, 1758 1758. Monodon monoceros. Linnaeus. Syst. Nat. ed. X, I, p. 75. Northern seas of Europe and America. 1781. Monodon narhval. Borowski. Gemeinn. Naturgesch. d. Thierreichs, 2 Did: 1804. Narwalus vulgaris. Lacépéde. Hist. Nat. Cétacées, р. 37, 142, pl. 4. Substituted for Monodon monoceros Linnaeus, 1758. 1811. Ceratodon monodon. Pallas. Zoogr. rosso-asiatica, I, p. 295. Seas of the Russian arctic (“In mare arcticum Ruthenum imperium alluente”). (У. H.) Diagnosis Only species of the genus. 589 792, Description Males can attain a body length of 610 cm and females 420 cm. A rounded small head is set off from the trunk by a weakly dis- cernible neck (Fig. 338). The spermaceti case slightly overhanging the upper jaw gives the head its roundness. The upper lip projects slightly forward above the fleshy lower lip. The slightly convex dorsum carries a low fold of skin up to 5 cm thick and 75 cm long. The asymmetry of the skull (Fig. 339) is manifest in the more intense development of maxillae and premaxillae on the left side while the cra- nium is better developed on the right side. The importance of the tusk (tooth) to the narwhal is not clearly understood. It has been suggested (Thompson, 1939) that the spiraled tusk serves to stabilize the animal body against rotation along the axis during circular movements of the caudal flukes during swimming. It is perhaps more correct to regard the tusk as a secondary sex characteristic of males (Chapskii, 1941; Sleptsov, 1955) serving as a defense organ. It is possible that the animal uses this tusk to pierce holes in the ice through which not only he himself, but also other members of the herd, i.e., females and young animals without tusks, can breathe (Tomilin, 1957). Cervical vertebrae 7, thoracic 11-12, lumber 6-10, and caudal 26-27. пи The main measurements of the skull (Tomilin, 1957), average of one to four animals are (in cm): condylobasal length 61; zygomatic width 39; length of rostrum 28; width of rostrum at base 21; length of lower jaw 50; and mandibular symphysis 6. (V.S.) Geographic Distribution Basin of the northern Arctic Ocean. This species lives mainly in the arctic zone, less often seen close to the coasts.!? Geographic Range in the USSR Narwhals have been observed in the Barents Sea in Novaya Zemlya at 76° N lat. (Russkaya Gavan’), in Franz Josef Land archipelago, in the midpart of the Kara Sea, around Dickson Island, and in the Chukchi Sea. In 1938, during the drifting of ice breakers “Sedov,” “Sadko,” and “Malygin,” narwhals were sighted no less than 15 times in the region 81° 21’-82° 15’ М lat. and 136° 15’ - 138° 15’ Е long. They were sighted 19 Distribution is equally known from the records of animal remains as from field observations. 793 ут А РРР РРР ЕЕ = РР РИ, 22:57: 589 from the floating arctic stations in June, July, and August of 1950 north- west of Wrangel Island, in 1955 north of De Long Islands, and in 1956 between Severnaya Zemlya and Franz Josef Land. Remains of narwhals have been found in the estuaries of rivers Olenek, Anabar, Khatanga, and Lena, and also on Lyakhovsk Islands, Cape Serdtse-Kamen’, and in the Chukchi Sea. In the summer months narwhals are most common in the waters from 80° to 85° N lat. 589 590 794 Ie = 7 Fig. 340. Skull of a narwhal with two tusks (figure by М.М. Kondakov). Geographic Range outside the USSR American Arctic, north of Labrador and Alaska and waters of Green- land (Fig. 341); known in the Canadian archipelago in Hudson Bay, Davis Strait, and in Baffin Bay right up to Smith Strait, in the waters-of Spitsbergen, later in the waters of Cape Barrow and in the Beaufort Sea close to the Colville estuary. Beached narwhals or their remains have been found at some points along the coasts of Greenland, in the Beau- fort Sea, near Cape Gal’skett, and in Kiwalik Bay in the Chukchi Sea at 60° N lat. and 162° E long. (Tomilin, 1957, 1962; Uspenskii, 1958; Hall and Kelson, 1959; Gest, Buckley, and Manville, 1960). (V.A.) Geographic Variation Not established. Biology Population. One of the comparatively rare forms of cetaceans, this species does not form large congregations. It is more abundant in the waters of Greenland than in other parts of the range. 795 (лэмэзту “WA ) 504220и0щ иороиой! Чечилеи dy} JO a8ueI $10945 ‘1$ "ВЫ 065 59 — 796 Food. {Information on this aspect is very scant. Cephalopods evidently serve as the main sustenance, followed by fish to a lesser extent. Among cephalopods, the squids Gonatus fabricii and possibly Rossia, Bathipoly- pus, Ommatostrephes, and some others have been detected. Fish food includes cod, flounder, halibut, skate, salmon, and herring (Smirnov, 1935; Tomilin, 1957). These animals feed predominantly in the pelagic region. The oral cavity is well adapted to feeding on cephalopods. Most of the above fish represent deep-dwelling forms. Behavior. There are quite a number of references in the literature to narwhals being encountered in large herds numbering thousands of animals. In our times, however, such large herds have not been reported. Narwhals are most often encountered in small groups or singly; rela- tively large collections are very rare. Herds consisting of a relatively large number of animals are probably of mixed composition, with adult males, females, and young animals. Small herds, in some cases, consist exclu- sively of females with calves. Some twenty instances are known when even a relatively small herd consisted exclusively of adult males. The narwhal usually remains far from coasts and approaches only sometimes in search of food. The animal can remain submerged under water for quite a long time, after which it surfaces eight or nine times successively at intervals of roughly 3 sec. During surfacing, the animal makes a loud expiration, flexes its body arcuately, but does not usually display the tusk above the water. Only when a group of narwhals dives in a small air hole, do their tusks project above the water (Gorbunov, 1940). In July, 1955, in the western part of Franz Josef Land archipelago (Cambridge Bay), a group of narwhals, a hundred strong, was sighted in a state of intense excitation. Sometimes the animals flexed arcuately, leaped out of the water like dolphins or, while diving under the ice, cast their head down such that the caudal flukes rose high above the water. While emerging from under the ice they often lifted the entire anterior half of the body with the tusk above the water surface. Meanwhile, at some distance, some ten narwhals remained immobile on the surface. The diving narwhals were probably feeding. At night, the animals were quiet and stationary with tusks visible above the sea surface. Some of the tusks crossed like swords (Sushkina, 1956). Once a narwhal was seen following a ship. Instances are known of groups of narwhals abandoning the ice and remaining a long time in small air holes, thereby falling easy prey to hunters. Mixed herds of narwhals and belugas have also been recorded. Narwhals produce various sounds, such as loud gurgles, resonant groans, and howling hoots (Tomilin, 1962). 592 797 Seasonal migrations and transgressions. Observations оп these aspects are very scant. In the waters of the Soviet Union, the zone of polar ice edges probably serves as the southern boundary of distribution in winter. In summer the animals ascend northward and are usually encountered beyond the 80th parallel. In Baffin Bay they arrive with the ice in Smith Strait in winter but again return south in the autumn, reaching Disko Strait by December (70° N lat.). Only some stray animals penetrate south of Sukkertoppen region (about 65° N lat.) while the main herd remains in the north. Narwhals are common from Angmagsalik village to Scoresby Sound (65 to 70° М lat.) on the east coast of Greenland in summer months (May to August) but the animals may ascend northward up to perhaps 80° N lat. or beyond. Some distant migrations of narwhals into the south are known. Their remains have been found in Mezensk Bay of the White Sea, along the Murman coast close to the Pechora estuary, at Jan Mayen, Iceland, Faroe Islands, and along the coasts of Norway, Scotland, and England. One instance of the death of a female narwhal was recorded in the Elba estuary in Holland. In the Pacific Ocean, transgressions are known in Bering Strait and toward St. Lawrence Island. One dead narwhal was found in the region of Port Moller in the central Alaskan peninsula. In the last century, one narwhal was found on Bering Island (Commander Islands) (N. Smirnov, 1935; Tomilin, 1957; Gest, Buckley, and Manville, 1960). Reproduction. Mating and parturition may occur at almost any time of the year (N. Smirnov, 1935). The birth of a narwhal calf was witnessed from the floating station SP-5 in the waters between Franz Josef Land and Severnaya Zemlya on July 30, 1956. It was 1.5 to 2 m long (Uspenskii, 1958). Very small as well as almost mature embryos were found at the same time (Fig. 342). The duration of gestation is not known. Usually one calf is born and twins are rare. The body length of a newborn calf hovers around 150 cm. Suckling calves are bluish-gray or slate-colored. As the ica й мой Ds i A z В Е. . be VELHO. pete pep Zz LG ip CEG ETE Fig. 342. Newborn narwhal (figure by N.N. Kondakov). 592 798 © animal grows, the ventral part of the adult gradually lightens (white or yellowish) while the flanks and dorsum darken with innumerable random dark brown spots spots of irregular shape. On the dorsum, especially the upper part of the neck and head, the spots are darker and denser than on the flanks. The periods of color variation as also the degree of their consistency with the changing age of the animal have not been established (Tomilin, 1957). Enemies, diseases, mortality, parasites, and competitors. One of the possible enemies of the narwhal is the white polar bear; the young are threatened by Greenland sharks. Only bone diseases are known in the narwhal. Two species of lice have been found in the skin folds of the nar- whal: Cyamus monodontis and C. nodosus. Among endoparasites, three species of nematodes are known. Anisakis (Anisakis) simplex Rudolphi, parasitizing the gullet, stomach, and intestine, has also been found in ten. species of toothed whales, two species of baleen whales, and in Steller’s sea lion. It has been detected in the North Sea and in the North and South Pacific Ocean. Terranova (Terranova) decipiens Krabbe, common in pinnipeds, seems doubtful in the narwhal. It definitely occurs in two species of baleen whales, in toothed whales, and in common porpoises. Torynurus alatus Leuckart is known only in the narwhal from the coast of Greenland. It has been detected in the skull cavity, eustachian tube, venous vessels, and lungs (Delamure, 1955). Field characteristics. Spotted coloration of the dorsum and body flanks is a characteristic feature of the species. Teeth one pair but usually do not emerge among females; one tooth among males turns into a long (up to 3 m) spiraled tusk projecting forward. A dorsal fin is absent. This species inhabits predominantly the Arctic Zone and can often be seen in air holes among highly compact ice. (V.A.) Economic Importance Only the local Greenland people regularly catch some narwhals and uti- lize their products to meet personal needs. Elsewhere in the range these cetaceans are caught merely incidentally in small numbers. Hence this species has almost no economic importance. The skin of the narwhal is quite tough and can be used for the same purposes as the skin of the beluga. The carcass of the animal is useful for preparing the same products as those from other marine mammals. The large tusk of the male can evidently be carved or sold as a sou- venir because of its rarity. Narwhal tusks were highly prized at one time (Vinogradov, 1949; Tomilin, 1957). (V.A.) 593 799 SUPERFAMILY OF SPERM WHALES Superfamily PHYSETEROIDEA Gill, 1872 Family of Sperm Whales Family PHYSETERIDAE Gray, 1821 Dimensions range from medium to large. The body form is diverse: the anterior half of the trunk of sperm whales is relatively short but very thick; in dwarf sperm whales, it is more elongated while the thickness is maximum between the flippers and the dorsal fin. The head may be very large, up to one-third of the body length, and obtusely flattened or slightly rounded anteriorly, or markedly smaller and more proportionate and rounded. The dorsal fin is low and humplike or quite high and falcate. The rostrum of the. skull is broadened, flattened, and: has a large depression on the dorsal surface. The lower part of the spermaceti sac lies inside this depression; among males, this sac projects forward in relation to the skull bones. The occipital bone forms a high crest. The lachrymal is fused with the zygomatic. The petrosal is enlarged toward the skull. The narrow lower jaw does not extend beyond the anterior section of the head: the symphysis of both its halves usually forms not less than one-third of its length. The teeth, located in deep alveoli, are preserved in most animals only in the lower jaw, being either rudimentary or altogether absent in the upper jaw. Keratinized projections on the palate replace the teeth functionally. Sperm whales feed mainly on cephalopods of different sizes, includ- ing very large ones (teuthophagous). These whales are distributed throughout the World’s Ocean. The northern limit of the range extends into the Pacific Ocean from Maina- Pylgo village and Cape Navarin to the Pribilof Islands and in the Atlantic, from Davis Strait to the region west of Jan Mayen and Spitsbergen and eastward up to Kanin Nos Peninsula. The southern boundary of distri- bution reaches the edges of drifting ice in the Antarctic. Sperm whales are common and abundant while dwarf sperm whales are few and very rarely encountered. Diaphorocetus Ameghino, found in the Lower Miocene, already had a skull with features characteristic of Physeteridae Linnaeus, though it was not as large as the modern forms, and had teeth in the upper jaw. Reduction of upper teeth is noticed among representatives of sperm whales from the Middle Miocene (for example, in Aulophyseter Kellog). The family is usually (Simpson, 1945) divided into three subfamilies, of which one is extinct (Hoplocetinae Cabrera, six genera) and two 594 800 contemporary: Physeterinae Flower (sperm whales proper) and Kogiinae Gill (dwarf sperm whales). The total number of genera is 19. Physeterinae includes ten extinct genera and one contemporary; Kogiinae comprises one extinct and one contemporary genus. Thus the contemporary gen- era of the family are: sperm whales Physeter Linnaeus and dwarf sperm whales Kogia Gray. In the context of systematics, the contemporary fam- ily of sperm whales is not a complex group and its division into genera raises no doubts. However, subspecies have not been well studied. Only one subfamily of sperm whales proper, physeterinae, with one genus of true sperm whales, Physeter Linnaeus, has been established for certain in the fauna of the USSR. The presence of the subfamily of dwarf sperm whales, Kogiinae, with the single genus Kogia Gray, has not been established but is entirely possible. Only sperm whales are of practical importance. They are hunted in large numbers, especially in our Pacific Ocean waters. (V.S.) Subfamily of Sperm Whales Subfamily Physeterinae Flower, 1864 Genus of Sperm Whales Genus Physeter Linnaeus, 1758 1758. Physeter. Linnaeus. Syst. Nat. ed. X, I, p. 76. Physeter catodon Lin- naeus. ‚1761. Catodon. Linnaeus. Fauna suecica, р. 18. Catodon macrocephalus Linnaeus = Physeter catodon Linnaeus. (V.H.) Large whales with sharply manifest sexual dimorphism: males up to 18-20 m long and females up to 11-13 m. The large head constitutes one-third of the total length of the body. The dorsal fin is weakly developed, low, and usually rounded in the form of a prominence. A few very low tubercles occur posterior to it. The broad flippers have a rounded outer margin. The caudal flukes are broad. The blowhole (nostril) is present in the anterior section of the head to the left. The length of the rostrum constitutes more than one-half the condy- lobasal length of the skull. Its dorsal surface has a deep depression. The skull bones are sharply-asymmetric. The right nasal bone fuses with the right premaxilla. The pterygoids are adjacent for a significant distance. Vertebrae usually 50. Teeth 257° 0°). The narrow scapula has a large coracoid process. Ribs up to 11 pairs, of which only 3 are joined to the sternum. The body color is dark and the abdomen only slightly lighter than the upper part. 595 801 The main food object is cephalopods while fishes represent ап addi- tional item. The ability to dive deep and remain long under water is a characteristic feature of sperm whales. These are polygamous animals. Various researchers have put the duration of gestation at 10 to 16.4 months. The periods of mating and parturition are protracted, Females apparently bear offspring once in two years. These whales are distributed predominantly in the warm and temper- ate waters of the Pacific, Indian, and Atlantic oceans (for more details, see “Description” of the species). Females, compared to males, are more thermophilic. Significant migrations are a characteristic feature. | In USSR waters, they are encountered in the Barents Sea and in the seas of the Pacific Ocean. Fossil remains have been found in the Upper Miocene and Pleis- tocene of North America and in the Lower and Middle Miocene of Europe. These whales are of great economic importance. A large number is caught every year. The genus comprises a single species: the sperm whale, Physeter catodon Linnaeus, 1758. (V.S.) SPERM WHALE Physeter catodon Linnaeus, 1758 1758. Physeter catodon. Linnaeus. Syst. Nat., ed. X, I, p. 76. Orkney Islands. 1758. Physeter macrocephalus. Linnaeus. Syst. Nat., ed. X, I, p. 76. “Euro- pean Ocean”. 1818. Physeterus sulcatus. Lacépéde. Mem. Mus. Hist., Nat., Paris, 4, р. 474. Japan. 1822. Physeter australasianus. Desmoulins. Dict. class. Hist. Nat., 2, p. 618. Moluccas and New Zealand. 1851. Catodon australis. Wall. Mem. Australian Mus., I, р. 1. Australia.! (V.H.) Diagnosis Only species of the genus. 1 After Hershkovitz (1966); А.С. Tomilin (1957) attributes this name to MacLeay, 1851. 802 Description The dimensions of this species are the largest of any toothed whale.” The body is teardrop-shaped and considerably thickset in the anterior half (Fig. 343). The anterior part of the huge head terminates bluntly in a “forehead”. The dimensions of the head are mainly determined by the massive spermaceti sac situated above the skull in the depressions of the maxillae and projecting beyond the anterior tip of the jaw. The зрегтасей sac consists of lower [melon or “Junk” and upper [spermatic organ or “сазе”| sacs made up of connective tissue with a high content of an only liquid, i.e., spermaceti (Fig. 344). The sacs are surrounded by connective tissue and muscles. Liquid wax (cetin) is the main constituent of spermaceti (96.99%). Adult sperm whales carry 1.0 to 5.7 tons of spermaceti. The head of males is relatively larger (up to one-third of body length) than in females (slightly more than one-fourth body length). With age, the head section of the sperm whale is relatively enlarged while the caudal end is correspondingly shortened (Ivanova, 1955). The relative increase in head size commences even in the fetal growth period (Ivanova, 1959). The cross section of the head appears rounded from above and keel- shaped from below (anterior to the jaw). The narrow lower jaw with the mouth closed is held to the head by the connective tissue edges [lips] of the upper jaw. The unpaired blowhole is S-shaped and located, unlike 596 in Other cetaceans, in the anterior part of the head left of midline and is 595 Fig. 343. Sperm whale, Physeter catodon, male and female (figure by N.N. Kondakov). Data on the weight of the animals and of the various parts of their body are given later under “Economic Importance” and Table 62 (р. 838). 595 596 803 Fig. 344. Section of the head of the sperm whale, Physeter catodon (figure by N.N. Kondakov): 1—blowhole; 2—left nostril; 3—right nostril; 4—upper sper- тасей sac [spermaceti organ or “сазе”]; 5—lower зрегтасей sac [“melon” or “Junk”; 6—sinew-tendon layer. 20 to 60 cm long. The eyes are roughly at the center of the head at the level of the posterior corners of the mouth. The section of the orbital aperture is 6-12 cm long and the diameter of the orb 15-17 cm. Small (up to 1 cm long) crescent-shaped ear openings are located posterior to the eyes and slightly below them. The head is set off from the trunk by a faintly perceptible neck. The trunk attains maximum thickness in the zone of the flippers but thins out gradually thereafter. The flippers are short and broad, with rounded outer edges. The large lobes of the caudal flukes are separated by a deep notch. Two to six very low tubercles are located posterior to the humplike dorsal fin. A rather low leathery keel runs along the lower side of the caudal stem. The integument of the sperm whale is quite thick (Table 58). In some sperm whales the thickness of the abdominal skin [epidermis plus dermis] may go up to 50 cm. The thickest epidermis and the densest 7 14227 O/ VLE 2 OM as 4 7. РА й Fig. 345. Sperm whale, Physeter catodon (ventral view) (figure by М.М. Kondakov). 596 597 804 Fig. 346. Head of a male sperm whale, Physeter catodon (figure by М.М. Kondakov). Table 58. Skin [epidermis plus dermis] thickness in different sections of the trunk of the sperm whale (in cm) (V.E. Sokolov) Sex Date of length, Abdomen Dorsum birth m Body Lateral ee ee ee ee АЗ 08 рате Under Around Before Above Anterior around of head flippers navel vulva _ flippers to flippers dorsal fin Female Aug. 7 10.2 15 10 = 21 — 6.5 6 Female Aug. 7 9.8 18 14.3 22 20 25 95 8.5 Female Aug. 7 9.3 16 11.5 15.5 12 16 10 7.5 Мае Sept.13 148 31 30 = 38 33 22 8 collagen fiber fascicles in the dermal layer occur on the anterior surface of the head. This is explained not only by the fact that this part of the head faces maximum resistance against water while swimming, but also by the fact that males in the period of mating butt each other with their “foreheads” just as rutting rams do (Sleptsov, 1955). The epidermis of the sperm whale is characterized by prominent projections of epidermal barriers extending into the dermal papillae. These projections, in many cases, have thickened distal and thin proximal sections. On the dorsum and flanks the skin surface is corrugated. In the depressions between the 597 598 805 tubercles, the epidermis forms projections into the dermal layer. On the throat 10 to 40 longitudinal furrows occur, up to 1 cm in depth and 5 to 50 cm in length (Tomilin, 1957). It is probably because of these that the throat can expand slightly while swallowing a large quarry (Beddard, 1900). A typical leathery callosity on the dorsal fin is regarded as a sec- ondary sex-related feature of females (Kasuya and Ohsumi, 1966). The body color is monochromatic dark (black, blackish-brown, blackish-cinnamon) with a much lighter-colored abdomen bearing an irregular white patch. The outline of this patch varies sharply in different animals. The sperm whales encountered in the waters of Japan can be classified into four color groups (Omura, 1950b): (1) with continuous dark gray color all over the body; (2) with a slightly lighter coloration on the lower side of the head and the lower jaw; [(3) with a light coloration on the lower side of the head and the lower jaw—this is an addition to Omura’s classification]; (3) with a light coloration all over the head; and (4) with a very light coloration all over the trunk. Animals of the third group are the rarest. Sperm whales caught off the Kuril Islands are mainly of two colors, dark and light, and form no less than eight variations of the white pattern (Ivanova, 1959*). With age, the color of the animals turns somewhat lighter. Many white and gray patches, bands, scratches, etc., formed as a result of skin infections caused by ectoparasites, lampreys, and suckers and hooks [jaws resembling the parrot’s beak] of squids are seen On the skin surface. Such patches are more among females than males (Omura, 1950b). In the sharply asymmetric skull the upper lateral sections of the maxillae form crests laterally surrounding the base of the spermaceti sac. The horizontally disposed palatines are covered from the rear by the pterygoids. The halves of the lower jaw form a symphysis constituting up to 50% (or slightly more) of the entire jaw length (Fig. 349). The relative length of the rostrum and mandibular symphysis increases with age (Tomilin, 1957). Cervical vertebrae 7, thoracic 11, lumbar 8, and caudal 24. Infor- mation on the fusion of the cervical vertebrae is contradictory. Some scientists assume that the cervical vertebrae of the sperm whale fuse into two independent groups of 1 to 3 and 4 to 7 vertebrae (Sleptsov, 1955). According to other authors, the sperm whale represents a unique case among cetaceans with its atlas remaining free while all the other vertebrae are fused (Tomilin, 1957). The level of the transverse processes of the lumbar vertebrae is much lower than that of the thoracic. In the five-digit limb (Fig. 348) the humerus is sometimes fused with the ulna and the radius. Phalangeal formula: I,, I, HI;, 1V4, and V3 (Beddard, 1900). &06 597 Fig. 347. Head of a female sperm whale, Kuril Island, 1962 (photograph by A.V. Yablokov). Ge S : 598 Fig. 348. Flipper of the sperm whale, Physeter catodon (figure by N.N. Kondakov). The peg-shaped homodont teeth of the sperm whale, often slightly curved midlength, can reach large proportions, up to 27 cm in length (Yablokov, 1958a), with the largest occurring in the middle of the jaw. The teeth are larger among males than females. Large teeth are encoun- tered only in the lower jaw. The teeth on the upper jaw are small, not strong, or may be altogether absent. The number of upper teeth among | 599 807 Fig. 349. Skull of the sperm whale, Physeter catodon (figure by М.М. Kondakov). males varies from nil to 19 and among females from nil to 10 (Sleptsov, 1955). The tooth crowns in embryos may have three cusps (Nishiwaki, Hibiya, and Ohsumi, 1958). The teeth of calves usually emerge by the end of the lactation period. With advancing age, the size of the teeth increases considerably.* Mandibular teeth begin to wear out a few years after birth (the anterior and middle ones in the first instance), some- times right up to the base, although the upper and lower teeth may not come into contact with each other. The gaps between the teeth vary from 3.5 to 15 cm. The teeth are firmly set in the alveoli and rise above the gum level only to one-sixth to one-fifth of their length; intense growth of cement is a characteristic feature. The volume of the pulp cavity can measure up to one-half that of the tooth. The teeth of adult animals have no enamel layer (Yablokov, 1958a). The total length of the largest male caught was 20.7 m, of the female 15.8 m. The average length of 2,510 males caught in the Northern hemi- sphere was 14.6 m and of 15,526 males caught in the Southern hemi- sphere 15.9 т; the average length of 1,190 females (waters of Japan) was 10.64 m (Tomilin, 1957). ЗА method was suggested for determining.:the age of sperm whales by counting the number of layers in a longitudinal slice of tooth (Nishiwaki, Hibiya, and Ohsumi, 1958). ‘599 808 The average main body measurements as percentage of body length among males and females caught off the Kuril Islands (Tomilin, 1957) are: distance from tip of snout to blowhole in males 2.30 (two mea- surements) and females 1.50 (2); distance from anal opening to notch between caudal flukes 30.30 (94) and 32.20 (46); height of dorsal fin 2.40 (147) and 2.60 (58); length of dorsal fin 8.00 (145) and 8.10 (62); length of flippers 9.6 (147) and 9.7 (74); maximum width of flippers 4.90 (26) and 4.90 (20); anterior-to-posterior width of caudal flukes 9.40 (192) and 9.70 (88). The main skull measurements (see Fig. 349) of three male sperm whales (Tomilin, 1957; Omura et al., 1962) are (in cm): condylobasal length 510, 450 and 360; interorbital width 220, 170; length of rostrum 370, 320, and 250; width of rostrum at base 150, 170, and 120; and length of lower jaw 450, 300 [The authors gave the skull measurements in decimeters instead of centimeters. I have added a zero to them to convert them to centimeters—Ed.] (V.S.) Geographic Distribution Sperm whales inhabit the warm and temperate zones of the World Ocean and are encountered almost everywhere. The summer ranges of males and females are diverse: the former are encountered at very high lati- tudes, reaching the arctic zone in the Southern hemisphere; the latter do not emerge beyond the temperate waters throughout the year. Geographic Range in the USSR Constitutes a small section of the total range of the species. These ani- mals are encountered in the basin of the Atlantic Ocean in the southern part of the Barents Sea: along the Murman coast, possibly up to Kanin Peninsula in the east (Fig. 350). In the Pacific Ocean they are distributed in the Sea of Japan where they are sometimes sighted in Peter the Great Gulf and even transgress into Zolotoi Rog Bay. They are encountered in the waters of the Kuril Islands, predominantly from the Pacific Ocean side. Large males inhabit the Sea of Okhotsk, mainly its northeastern section. Farther northward, these animals are known in Kronitskiy and Kamchatka gulfs off the southeastern coast of Kamchatka and around the Commander Islands. They have been sighted off Cape Navarin in the southern part of the Gulf of Anadyr. Geographic Range outside the USSR Distributed almost everywhere in the North Atlantic Ocean (Fig. 351). Throughout the year, sperm whales are sighted off the Cape Verde, 809 ‘(aa,uasIV “W'A) USSN ЭЦ: Ш! ‘Uopowws sajaskyg ‘этеца шло@з ay] Jo э8иеы “OSE ‘9 009 “(лэ.иэзлу У А) иоро1р> dajaskyg ‘aye шлэ4$ ay) Jo э8иел $910э4$ ‘16$ “314 109 SSS SSS ШУ 000$ 000% 0006 0002 000: 0 0001 602 811 Canary, Azores, and Madeira islands and along the coasts of Spain and Portugal, off the Antilles and Bermuda islands, and along the coasts of Florida, i.e., up to 520- 15° N lat. They are quite common in the Mediter- ranean Sea. They are not infrequent in the waters of Great Britain, off the Orkney and Shetland islands, widely distributed along the coasts of Norway, in the waters of Iceland, and off the Faroe Islands. Data on their distribution along the American mainland between Florida and Newfoundland are altogether lacking although some random observa- tions suggest that these animals are encountered here. In the North Atlantic, sperm whales reach Newfoundland, Labrador, Davis Strait, and the coasts of southern Greenland. They inhabit the waters of Jan Mayen (70° N lat.) and even Spitsbergen (around 80° N lat.). Information on the distribution of sperm whales in the equatorial section of the Pacific Ocean is highly approximate. They have been assumed to be quite common close to Kalimantan, Celebes, Mariana, Marshall, and Bonin islands, and are possibly encountered in the Sulu and Banda seas, i.e., not only reach the equator, but sometimes even the equatorial waters of the Southern hemisphere. The contemporary regions of wintering of northern sperm whales are possibly bounded by the line: Bonin Islands - Hawaii- California coast (Berzin and Rovnin, 1966), 1.е., within 15 to 20° М lat. Sperm whales are common along the southern and eastern coasts of Japan but rare in the East China Sea and in the southern part of the Sea of Japan. In the northeastern half of the Pacific Ocean, sperm whales are distributed from California to the Gulf of Alaska and the Alaskan Peninsula and along the islands of the Aleutian range. In the Bering Sea, they are common in Bristol Bay and in the waters of the Pribil of Islands, occupy the eastern part of the sea bound by the shallow-water line running from Bristol Bay to Cape Navarin on the Asian coast and probably up to St. Lawrence Island. In the Southern hemisphere, sperm whales occupy the entire zone of tropical and temperate waters. In the Atlantic Ocean, they are known from the coasts of Brazil to the Falkland Islands and Tierra del Fuego in the south, and sighted in the waters around Tristan da Cunha, Ascension, and St. Helena islands. In the eastern part of the Atlantic, they are com- mon from the equatorial waters of the Gulf of Guinea to the southern tip of Africa. In the Indian Ocean, the largest number probably inhab- its the waters of southeastern Africa and Madagascar; they are known off the Mascarene, Seychelles, Cocos, and Christmas islands, and along the coasts of Australia. Penetrating northward, the distribution crosses the equatorial zone and this animal is known in the Arabian Sea and transgresses into the Red Sea. In the southern half of the Pacific Ocean, 603 812 sperm whales are abundant along the coasts of Peru and common in the waters of Chile. They are encountered close to numerous islands, including Fiji, Samoa, etc., and are common in the waters of Australia and New Zealand. In the summer months male sperm whales are widely distributed in the Antarctic waters and reach the southern Arctic zone (Kirpichnikov, 1950b; Tomilin, 1957; Berzin and Rovnin, 1966; Berzin, 1971). In spite of the fact that the two ranges of the species—northern and southern—are distinct, there is no real demarcation in the distribution of sperm whales. They are encountered in the equatorial zone of the Northern and Southern hemispheres throughout the year and hence it is very difficult to draw the boundary of distribution in either hemisphere. While a demarcation is possible in the Atlantic Ocean (though highly approximate), it is difficult to do so in the Pacific Ocean. Apparently the equatorial zone serves as the area of contact between the northern and southern populations of the sperm whale but information is not sufficient to resolve this problem. (V.A.) Geographic Variation A comparative morphological study of sperm whales from different parts of the World Ocean revealed no significant differences in the body proportions (Ivanova, 1955, 1961). However, some difference in the overall dimensions does exist between the northern and southern sperm whale populations (Nishiwaki, 1955). Based on this and the time differential in the main biological rhythms of the whales of the North- ern and Southern hemispheres, two subspecies of sperm whales are recognized (Tomilin, 1957). Only one of these subspecies is found in the USSR. Northern sperm whale, P. c. catodon Linnaeus, 1758 (syn. macro- cephalus). Animals of relatively smaller dimensions, with the males measuring an overall length of 14.6 m (North Pacific Ocean). They are encountered in the Barents, Japan, Okhotsk, and Bering seas, and the Pacific Ocean. Outside the USSR, they are found in the North Pacific and Atlantic oceans. Systematic relationships between the sperm whales of the North Pacific and Atlantic oceans have not been studied. Evidently these pop- ulations are highly isolated (amphiboreal distribution) and possibly may not be identical. The name sulcatus Lacépéde could be applied to the Pacific population. 813 Outside the USSR, some scientists recognize the form Р. с. aus- tralasianus Demoulins, 1827,* characterized by very large dimensions (average length of males 15.9 m). These animals are encountered in the warm and temperate waters of the Atlantic and Pacific oceans in the Southern hemisphere, the Indian Ocean, and the Antarctic. The interrelations between the two subspecies of sperm whales in the equatorial zone, especially in the Pacific Ocean, have not been studied. It is possible that many local herds of sperm whales exist in different parts of the species range; sometimes such herds can be localized in a relatively small water body (Klumov, 1955) (see “Seasonal Migrations and Transgressions”). (V.A. and V.H.) Biology Population. The sperm whale can be regarded as one of the more abun- dant species of large whales. The population is perhaps most abundant in the Pacific Ocean. The probable population of the sperm whale in the North Pacific Ocean has been put at roughly 150,000 (Nishiwaki, 1966). Up to 4,000 - 5,000 sperm whales were caught in some years off the coasts of Chile and Peru, indicating a large population of these whales in the South Pacific Ocean; they are also abundant in the other waters of the Southern hemisphere. Thus, in some years in the Antarctic alone, up to 6,000-7,000 male sperm whales were caught. Insofar as the North Atlantic is concerned, even approximate population data are not avail- able. It is probable that the population here is relatively small, which is Supported by the comparatively low level of hunting. The maximum number of sperm whales is caught in the waters of the Azores where the total catch sometimes does not exceed 1,000 per year. Food. Two groups of animals, cephalopods and fish, constitute the main food of sperm whales in both the Northern and Southern hemi- spheres, with cephalopods predominating. The food of sperm whales has been studied in great detail in the North Pacific Ocean. Data for the other regions of the World Ocean are fragmentary and incomplete (Table 59). Three species of squids predominate in the food of the sperm whale from the Bering Sea: Gonatopsis borealis, Gonatus magister, and Moroteuthis robusta; Meleagroteuthis separata is of lesser importance (Table 59). Among the fishes, redfish occupies first position of importance as a food item. It is followed by the smooth lumpsucker 1 А.С. Tomilin (1957) designated this form as P. с. australis MacLeay, 1851, which is hardly correct (see synonyms and note on p. 801). 604 814 Bering Sea (Berzin, 1959) Cephalopods Gonotopsis borealis Gonatus magister Gonatus fabricti Moroteuthis robusta Meleagroteuthis separata Galiteuthis armata Onychoteuthis banksti Chiroteuthis veranyi Fish Smooth lumpsucker, Aptocyclus ventricosus Perches, Sebastodes (few species) Rat-tails, Coryphaenoides sp. Lancetfish, Alepisaurus aesculapius Skates, Raja (two sp.) Pacific lamprey, Entosphenus tridentatus Sculpins, Myoxocephalus sp. Table 59. British Columbia (Pike, 1950b) Kuril Islands (Betesheva and Cephalopods Gonatus fabricii Moroteuthis robusta Fish Sea perch: Sebastodes ruberrinus Sebastodes sp. Ragfish, [costeus aenigmaticus Skate, Raja rbina Arctic lamprey, Lampetra Japonica Salmon, Salmonidae Akimushkin, 1955; Akimushkin, 1957; Betesheva, 1960, 1961; Tarasevich, 1963) Cephalopods Gonatus magister Gonatus fabricii Taonius pavo Moroteuthis robusta Onychoteuthis banksii Architeuthis japonica Meleagroteuthis separata Chrioteuthis veranyi Galliteuthis armata Stigmototeuthis dofleini Octopodoteuthis longiptera Crystoloteuthis berhingiana Paroctopus conispadiceus Japatella heathi Grimpoteuthis albatrossi Alloposus mollis Octopus gilbertia Cirroteuthis sp. Amphitretus sp. Octopus sp. Octopodidae sp. Fish Black rat-tail: Coryphaenoides acrolepis Albatross rat-tail, C. pectoralis Piked dogfish, Squalus acanthias Lancetfish, Alepisaurus aesculapius Pacific saury, Cololabis saira Pacific cod, Gadus morhua macrocephalus Alaska pollock, Theragra chalcogramma Smooth lumpsucker, Aptocyclus ventricosus Humpback salmon, Oncorhynchus gorbuscha Navaga, Eleginus navaga gracilis Podonema, Podonema longipes Sharks, Somniosus sp. Anglerfish, Oneirodes sp. Sculpins, Cottidae gen. sp. Salmon, Oncorhynchus sp. Greenlings, Pleurogrammus sp. Skates, Raja sp. 815 Food objects of sperm whales Japan (Mizue, 1951b) Azores Islands (Clarke, 1956) Antarctic (Korabel’nikov, 1959) Cephalopods Squids Fish Pacific cod, Gadus macrocephalus Alaska pollock, Theragra chalcogramma Scorpionfishes: Sebastodes flammeus Sebastodes iracudus Pacific saury, Cololabis saira Pacific sardine, Sardinella melanosticta Round herring, Etrumeus micropus Japanese anchovy, Engraulis Japonica Pacific mackerel, Pneumotophorus japonicus Spotted mackerel, Scomber tapeinocephalus Sharks, Selachii Cephalopods Histioteuthis bonnelliana Cucioteuthis unguiculatus Tetronychoteuthis dussumierii Lepidoteuthis physeteris Ancistrocheirus lesneuri Loligo forbesi Architeuthis sp. Fish Long-finned tuna (albacore), Thunnus alalunga Anglerfish, Ceretias holbolli Atlantic footballfish, Himantolophus greenlandicus Barracuda, Sphyraena sp. Cephalopods Onychoteuthis banksii Architeuthis sp. Squid (genus not identified) Fish Anglerfish, Ceratias holbolli Patagonian toothfish, Dissostichus eleginoides Southern putassu, Micromesistius australis Skate, Raja griseocaudata 604 816 which, however, is encountered in the stomach of sperm whales more often than any other fish. The ratio of the various groups of animals in. the food of the sperm whale is as follows: of the 107 stomachs with food, 69 (64.4%) contained only squids, 28 (26.2%) squids and fish, and 10 (9.4%) only fish. The importance of cephalopods and fish in the food of sperm whales undergoes variation relative to the region of habitation. In the waters of the Commander Islands, 74.4% of the stomachs anal- ysed contained squids exclusively, 23.9% squids together with fish, and only 1.4% fish exclusively; for the region of the western islands of the Aleutian range, these values were respectively: 54.0, 26.9, and 19.1%, and in Olyu- torskiy Gulf and adjoining waters 35.7, 35.7, and 28.6% (Berzin, 1959). The stomach of sperm whales from the waters of the Kuril Islands (360 stomachs) contained 28 varieties of cephalopods (21 squids and 7 octopuses). Cephalopods constitute about 95% (by weight) of the food intake of the sperm whale and fish less than 5%, although the latter were _ detected in almost one-third of the sperm whales investigated. Squids are 605 606 of basic importance among cephalopods; octopuses constitute no more than 4% of the intake. But only seven species of squids are of prac- tical importance as food: Gonatus magister, G. fabricii, G. simile, Tao- nius pavo, Galliteuthis armata, Chiroteuthis уетапу, and Maleagroteuthis separata constitute up to 80% of the cephalopods consumed, with 60% accounted for by the three species of the Gonatidae family alone. Cuttle- fish, abundant in the Far Eastern seas, have not been found in the stom- ach of the sperm whale (Betesheva and Akimushkin, 1955; Akimushkin, 1957; Betesheva, 1960, 1961). In the waters of Japan an analysis of 1,627 stomachs showed that squids represent the main food here too, although octopuses were found quite often but in small quantities. All the other food objects were comparatively rare. The male sperm whale feeds on a more diverse diet than the female (Mizue, 1951). The species of cephalopods were not indicated in this report. In the waters of British Columbia, where only 50 stomachs were investigated, squids were similarly found to be of primary importance. Frequency of Encounter (Number of Stomachs) of Various Objects in the Food of Sperm Whales from the Waters of British Columbia (Pike, 1950b) Squids 35 Ragfishes (Icosteidae) 16 Sea perch 16 Skates Э Salmon (?) 3 Lamprey 1 606 817 Squids serve as the main food of Antarctic sperm whales too: fish was found in only 6 (5.2%) of the 129 stomachs investigated (Kora- bel’nikov, 1959). According to the latest data, the large Antarctic tooth- fish is often found in the stomach of sperm whales from various regions of the Antarctic (V.L. Yukhov). Cephalopods have been recorded as the main food of sperm whales from many other regions of the World Ocean: the North Atlantic, the Mediterranean Sea, tropical zone, and along the coasts of Australia (Tomilin, 1957) (see Table 59). Sperm whales consume mainly comparatively small squids, 30-40 to 100 cm and less often 150 cm long, but sometimes the stomachs have contained giant animals. Thus a sperm whale 14.3 m long killed in the Azores in 1955, contained a mollusk (with tentacles) 1,049 cm long, weighing 184 kg (Clarke, 1956) (Table 60). In the Azores the main food of males as well as females consists of comparatively small squids of nearly identical size even though the male is considerably larger than the female. Sponges, crabs, crustaceans, and even bits of the skin of pinnipeds, sometimes found in the stomach of sperm whales, are merely items swallowed incidentally. Stomachs with a large volume of food are relatively rarely seen. Of the 264 stomachs (Kuril Islands) analyzed, 13 (4.9%) were well filled, 28 (10.6%) were moderately full, 125 (47.4%) contained little food, and 98 (37.1%) were empty (Tarasevich, 1963). The weight of the stomach contents of 120 sperm whales showed 33% with 4 to 10 kg of food, in some cases going up to 200 [sic] kg (Betesheva and Akimushkin, 1955). Most often, the stomachs revealed only the “beaks” of squids, sometimes Table 60. Sizes of squids consumed by sperm whales (Clarke, 1956) Squids found Mantle length Male sperm whales Female sperm whales of squids, cm wl es el УРНЫ Number % Number % 60-90 41 59 28 67 90-150 26 37 13 31 150-180 2 3 = = 180-240 1 1 1 2 Total 70 100 42 100 Average length 95 92 of squids, cm 607 818 more than 10,000. The maximum number of beaks found was 28,000, corresponding to the consumption of 14,000 squids. This, however, does. not represent a one-time intake since beaks are digested very slowly (or not at all) and probably accumulate over a rather long period. It is interesting that the squid Ommatostrephes sloanei-pacificus, inhabiting the upper horizons of the sea and most abundant in the Far Eastern waters, is of almost no importance in the food of the sperm whale and that the abundant cuttlefish, living close to the surface, is not touched at all. The main food consists of bathypelagic species of cephalopods and fish, most of which do not rise to a depth of less than 500 m and some of which live at depths of 1,000 m or less (Akimushkin, 1954b). Thus, sperm whales catch their quarry at depths of not less than 300 to 400 m, where these whales have almost no food competitors with the possible exception of beaked whales (Ziphiidae). In search of food, sperm whales can descand right to the ocean floor since their stomachs sometimes contained bottom-dwelling deepwater animals. In the 1960s, with the increasing geographic spread of sperm whale hunting, more information has become available about the food of these whales in regions earlier not studied. In the open waters of the northeast- ern part of the Pacific Ocean, the predominant contents of the stomach of sperm whales were the remains of squids, mainly Taonius pavo, Chi- roteuthis veranyi, Meleagroteuthis separata, Galiteuthis armata, and Gona- tus magister. Among fish, the following were encountered: Alepisaurus aeusculapius, Pseudopentaceres richardsoni, Sebastodes alutus, and Aptocy- clus ventricosus. The stomach of sperm whales from the waters of central California revealed the squids Moroteuthis robusta and Gonatus borealis, and among fish, the sharks Apristurus brunneus and Squatina californica, and sablefish, greenling, and lanternfish (Tarasevich, 1968; Berzin, 1971). In the waters of New Zealand and the nearby islands, squids were found in 84% of the stomachs studied. Remains of scorpionfish, skates, sharks, some invertebrates (fire salpians, shrimps) and even brown algae were detected. In the southern part of the Indian Ocean the stomach of sperm whales predominantly contained squids, including large ones (up to 9 m [sic] in length). Barracuda and sometimes porcupinefish were among the fishes frequently encountered in the stomach contents. On the whole, the main food of the sperm whale every where is cephalopods (squids up to 80% of intake and octopuses), including roughly 40 species and constituting not less than 95% of the total weight of the food intake. The stomach of sperm whales also contained the rem- nants of over 50 species of fish but, in spite of this diversity, fish account for no more than 5% of the total intake. Most of the animals serving as food objects of the sperm whale are deepwater species (Berzin, 1971). 608 608 819 Relatively often, the stomach of sperm whales revealed altogether extraneous matter (Fig. 352): rubber boots, wire reels, glass fishing buoys, rubber gloves, plastic toys (motor cars, pistols, dolls, pails, etc.), jugs, plastic bags, coconuts, vinyl chloride bags, fishing tackle with hooks, empty bottles, apples [sic], and many others (Berzin, 1971). Daily activity and behavior. Sperm whales are polygamous animals with sexual dimorphism distinctly manifest in their dimensions. In warm waters they form groups consisting usually of 10 to 15 females and a large male. But groups of hundreds of animals are not uncommon. Adult males that are not members of these groups (harems) remain aloof, often indi- vidually; they gather into relatively large groups only sometimes at food sources. The latest observations have shown that the males included in harems are not the largest and oldest but much younger males of 13 to 14 m length with high sexual prowess. They are the leaders of harems and drive away the old males, who subsequently remain aloof. All of this calls for a detailed review of the contemporary concepts of harems, their composition, behavior, interrelationships between individual age and sex groups in small and large collections, and other aspects of fam- ily and herd behavior of the sperm whale (Berzin, 1971; V.A. Zemskii, D.D. Tormosov, Yu.A. Mikhalev). Fig. 352. Foreign objects recovered from the stomach of sperm whales (photo- graph by A.A. Berzin). 609 820 In the regions of summer habitation, depending on age and size of animals, male sperm whales generally form groups of definite composi- tion. The groups most often consist of comparatively same-sized animals rather than animals of different sizes. Of the 23 groups analyzed in the Gulf of Alaska, 18 (78.3%) consisted of whales of nearly similar size and only five (21.7%) of animals of different sizes. It has been suggested that the reason for animals of the same size grouping together lies in their identical abilities for getting at food available at different depths. Large whales can dive deeper and feed at places where much smaller animals cannot. The same-sized groups of very small whales usually con- sist of nearly same-aged animals (with a difference of one or two years). In groups consisting of much larger whales, the age differences of the members are greater (sometimes 10 years). This factor probably deter- mines the high individual variations in the dimensions of adult whales whose growth has ceased (Tarasevich, 1967a). The nature of diving of the sperm whale is typical they can remain submerged under water for up to an hour and sometimes even longer (two hours is considered maximum). Before prolonged submergence, the sperm whale dives sharply, its body is steeply curved or humped and the bent caudal stem is visible on the sea surface. The animal dives almost vertically and, in most cases, the caudal flukes are displayed above the water. The appearance of this “butterfly form” above the sea surface serves as a positive indication that the whale has submerged for a long time and may surface very far from the site of submergence. - The maximum depth of submergence of the sperm whale has not been established. Nineteen instances are known of damage to underwater cables caused by sperm whales entangled in them, exclusively in tropical and temperate waters (between 46° S lat. and 46° N lat.). The depth at which dead sperm whales were detected ranged from 118 to 1,116 m. Having become entangled in the cable, the sperm whale grips it with the teeth, tears the insulation, and thereby damages it. In 1951, a cable connecting Lisbon and Malaga revealed this type of damage at a depth of 2,200 m, so far the maximum known depth. In most cases, the depth of cable damage exceeded 500 m, sometimes 1,000 m. Most often, the lower jaw and caudal fin of the sperm whales were entangled in the cable. It has been assumed that the whale held the cable in its mouth while chasing for food and then began to whip it when the slackened cable wound around it. It has been suggested that sperm whales become entangled at low depths and then gradually slip down along the slope of the sea bed. However, the nature of the entanglements revealed that they occurred at the places where the animals were actually found (Khizen, 609 821 1957; Yablokov, 1962). Sperm whales perhaps are actually capable of descending to a depth of 1,000 m or more. After prolonged residence under water, the sperm whale rests a long time on the surface (Fig. 353). At this time, it remains almost stationary, only slightly moving forward; in a horizontal position, it rhythmically submerges and blows every 10 to 15 sec. The number of blows varies and probably depends largely on the time spent under water. Usually there are 20 to 40 blows but up to 75 in some cases. Compared to the older animals, the young ones produce fewer blows. The former evidently dive deeper and for a longer duration. During this characteristic rest period, the sperm whale remains very quiet and unconcerned, lying on the surface for 10 min or longer; in most cases, a whaling boat can approach such a whale within shooting range. During hunting around Antarctica there have been instances of a whaling boat literally approaching a sperm whale so close as to hit the animal with its stempost (V.A. Arsen’ev). Sometimes a sperm whale rising from the deep emerges almost ver- tically from the water, thrusting about half of its trunk clear. In other cases, the animal leaps high at an angle to the sea surface and falls on the water with a loud splash. The speed of the sperm whale is comparatively slow. The so-called moving or migrating whales (traveling from one region to another) usually travel straight at 5 to 7 miles/hr. They swim close to the water surface, often make shallow dives, and produce many blows. Feeding Fig. 353. Sperm whales at sea (figure by NN. Kondakov). 610 612 822 animals move more slowly, probably at not more than 3 miles/hr. When searching for food, they remain submerged a long time, changing course under water regularly and very sharply. A frightened or wounded sperm whale moves at maximum speed, quite often along a straight course, and produces blows frequently. The maximum speed of a large sperm whale hardly exceeds 10 miles/hr. In any case, ships covering 12-12.5 miles/hr invariably overtake an escaping sperm whale after 40-50 min of pursuit. A sperm whale wounded by a harpoon sharply dives deep but soon reemerges on the surface, rising almost vertically, and displaying its huge obtuse head. If the movement of the whale boat is not arrested in time, the surfacing whale can appear underneath it and damage the bottom severely by colliding against it with its head. Instances are known of such situations wherein the whale damaged the whaling boat by distorting the propeller blades or even the crest of the shaft. These cases formed the basis for the widespread belief that a wounded sperm whale often actively attacks a ship. Such attacks are no doubt unintentional and inflicted at random by the surfacing sperm whales (V.A. Arsen’ev). Instances of sperm whales being beached, either singly or in groups, are not infrequent along the coasts of Denmark, France, Florida, Cal- ifornia, Tasmania, and New Zealand. Eight cases have been registered of groups of beached whales ranging from 16 to 36 animals. Sometimes Fig. 354. Typical blows of the sperm whale. Pacific Ocean (photograph by M.M. Sleptsov). 613 823 such groups consisted exclusively of females but often of lone males. Twenty-two males cast up on January 16, 1954 on the coast of California were roughly identical in size (Gilmore, 1957; Tomilin, 1962) (Fig. 355). Seasonal migrations and transgressions. The nature and courses of migration of sperm whales have not been thoroughly studied. The scheme of migrations is generally as follows: in the winter months, most of the sperm whales gather in warm and tropical waters and move in sum- mer into temperate waters, although some males even depart into the cold waters of the Northern and Southern hemispheres. The main fac- tor determining the courses of migrations and the distance of travel of sperm whales in cold waters is the effect of such currents as the Kuroshio in the Pacific Ocean and the Gulf Stream in the Atlantic. The zone of influence of these currents provides favorable conditions for the survival of the main food objects of the sperm whale, 1.е., cephalopods, whose entry into the north in summer months is determined by the intensity and direction of the currents. Sperm whales move into relatively high latitudes in pursuit of these cephalopods. Early in spring, in the North Pacific Ocean, sperm whales begin to migrate from the warm waters northward. Around the Bonin Islands the first females are sighted February end and by March their catch is maximum. Most of the whales migrate along the eastern coast of Japan. Some whales migrate northwest and are caught in small numbers on the western coast of Kyushu Island. They are sometimes encountered in the East China Sea. Sperm whales are not caught off the Korean coasts but Fig. 355. A group of beached sperm whales. California (from R.M. Gilmore, 1956). 611 824 some numbers enter the Sea of Japan from the south (Omura, 1950). Some lone sperm whales, quite rare in fact, are noticed in Peter the Great Gulf and sometimes transgress even into the Bay of Zoloti Rog (Vladivostok). In the waters of southern Japan, sperm whales are seen early in spring, up to April; the earliest to arrive here are young, already mature males. In June-July, most of the sperm whale herds*concentrate in the waters of Japan; at this time, males, females, and calves live here. In July, they move northward but the groups break up in the waters north of 45° М lat. Males separate from the general groups and go northward while females with calves inhabit the much warmer waters. The hunting of sperm whales in the waters of Japan is most fruitful from June through November (Nishiwaki, 1966). In the central part of the Kuril range, the first sperm whales, excep- tionally large single males, are sighted early in April and groups begin arriving by May. By mid-May, herds of sperm whales do not actually transgress north of the 48th parallel. In the northern part of the Kuril range, lone male sperm whales are seen at April end to early May and by May end the so-called harem whales begin to arrive. By mid-June, stable collections, called herds, of sperm whales of different sexes and ages are formed here. Large males by this time move farther northward in large numbers (Tarasevich, 1965). In the first half of June, mixed groups of sperm whales are fairly uniformly distributed from the Pacific Ocean side of the middle and northern sec- tions of the Kuril range and their herds depart farther from the coasts. Adult lone males by this time are seen in the Sea of Okhotsk, which is not at all visited by females or young animals. In July, the number of sperm whales around the Kuril Islands is far smaller and only minimal groups are seen throughout the region. It can be assumed that herds of sperm whales often travel at this time toward southeastern Kamchatka and are partly scattered over this immense water body. At the end of July in the central part of the Kuril range (Bussol Strait-Simushir Island), new herds of sperm whales of medium size, con- sisting mainly of young males, are seen approaching from the south. At the very end of July, new herds are again seen in the region of Urup—Simushir. These herds contain many females and some large males. These are clearly the returning harems. During the first 20 days of August sperm whales remain in this region, even slightly extending southward (up to Friza Strait), but do not advance northward at all. At this time herds consist of very small animals but large whales begin to be sighted again by the end of August. They probably now begin to move out of the Sea of Okhotsk southward. 612 825 This pattern of arrivals confirms the assumption that during summer two narrowly localized sperm whale herds, i.e., southern Kuril and north- ern Kuril populations, inhabit the Kuril Islands. Whales of the southern Kuril herd inhabit the northern parts of the Japanese Islands and the southern parts of the Kuril range, reaching its middle in the north. The northern Kuril herd inhabits the waters of the northern Kuril Islands and is confined to the coasts of southeastern Kamchatka but may reach the waters of the Commander Islands (S.K. Klumov). In September, the maximum number of sperm whales is seen in the Kuril waters since the whales returning from the northern waters now begin to arrive. October is the month of en masse departure of sperm whales southward and their herds around the Kuril Islands gradually thin out. Individual animais or even small groups may, in favorable years, remain for wintering close to the Kuril range (Sleptsov, 1955). The northern boundary of distribution of adult females and young sperm whales of both sexes is regarded as 51°N lat. (Omura, 1950) although they are encountered even farther north, up to 52-54° N lat. in Kronotskly and Kamchatka straits and around the Commander Islands (Kirpichnikov, 1950b; Sleptsov, 1950; Tomilin, 1957). Hence, in favorable years, females can reach even up to 53-54° N lat. The usual boundary of distribution of adult males is 61 -62° М lat. (Cape Navarin) where whales from both the western and eastern coasts of the Pacific Ocean live. It is more probable that the region of Olyutorskiy Gulf (60° N lat.) serves as the northern boundary of distribution for males of the Asian herd although some stray whales have reached 65°30’ М lat. under favorable conditions (Omura, 1950). Migrations of the sperm whales of the American herd are as fol- lows. From the wintering regions, disposed along a tentative line from the Hawaiian Islands to the coast of California, sperm whales begin to migrate northward at March end to early April. The period of migra- tions is considerably prloonged and hence they are sighted even in May in the southern parts of the range. The large extent of the wintering range along the longitude makes for several courses of migrations. One group moves along the coasts of the American mainland, another moves within 145 - 150° W long., and yet another between 162 and 167° W long. In April or early May, all these groups reach roughly 50° N lat. where the comparatively linear course takes a complex route, At these latitudes the males from the mixed groups finally separate out, with some ani- mals moving into the middle and eastern sections of the Gulf of Alaska and the Pacific waters of the Aleutian range, and others occupying the northernmost part of the Gulf of Alaska, moving far westward right up to the Commander Islands, entering-the Aleutian Straits into the Bering 613 826 Sea (Bristol Bay and Pribilof Islands) and, spreading along the east- ern Shallow-water section of the sea, reach Cape Navarin and the Gulf of Anadyr. These are exclusively males of various dimensions and ages which gradually isolate from the mixed herds and gather in the regions of food concentration, at times forming significant herds. It has been assumed that the much younger, although sexually mature males are the first to separate from the common.herds, followed by the larger older animals. This is possibly associated with the concluding phase of repro- duction. The northern boundary of distribution of groups of females and young animals in all probability is 50-51° М lat. They are confined predominantly to the Gulf of Alaska but are not infrequent in these same latitudes around the Aleutian Islands, sometimes reach the west- ern section of the range and, more rarely, even the Commander Islands. Under favorable conditions, in some years small groups penetrate even farther north and are seen in the Bering Sea (Berzin and Rovnin, 1966; Tarasevich, 1967a). i The return of groups of sperm whales to the wintering regions com- mences in September. The sequence and periods of migrations of indi- vidual age and sex groups have not been unraveled. Migrations proba- bly occur by the same routes as when moving northward. In November, the sperm whales practically abandon the northern regions of the range although some stray animals or small groups may overwinter there. Such instances are known even in the waters of the Commander Islands. Sperm whales reside almost everywhere around the central and southern Kuril Islands, and sometimes comparatively large herds are seen here in winter. Wintering sperm whales have been recorded even in the Gulf of Alaska (Zenkovich, 1936b, 1952; Sleptsov, 1955; Tomilin, 1957). The periods of migrations of sperm whales in the North Atlantic are roughly the same as in the Pacific Ocean. After overwintering in the southern regions of the range, the whales begin to move north along the western and eastern coasts of the ocean in early spring. Mixed groups of sperm whales probably do not proceed north of 50°N lat. but mature males of different ages reach Iceland, Greenland, Davis Strait, and Jan Mayen and Spitsbergen islands. This is somewhat more northward than in the Pacific Ocean due to the influence of powerful Gulf Stream cur- rents. Some males, depending on hydrological and trophic conditions, may sometimes overwinter in the north but most of the whales return to the southern waters in autumn. Regular seasonal migrations are also known among populations of the Southern hemisphere but their extent differs for mixed groups and adult males. Mixed herds of sperm whales apparently do not travel beyond the south 40° latitudes since only one case has been recorded of a 614 827 female sperm whale being caught off South Georgia Island (54° 30’ $ lat.). In the summer months, however, males are widely distributed in the waters of Antarctica and reach the ice there—which is not the case in the Northern hemisphere. In autumn males abandon the Antarctic Zone and return to their points of wintering. Sperm whales have not been observed at any time of the year in the Gulf of Guinea (Kirpichnikov, 1949b, 1950b). Observations over several years in the various zones of the South- ern hemisphere revealed that groups of small sperm whales, consisting of immature males and females as well as adult females, remained mainly in tropical waters. Large males were quite rare among them. The approxi- mate body length of the small sperm whales was 7 to 10 m. In the subtropical zone small groups of sperm whales, consisting of males that had attained sexual maturity but were still not participating in reproduction (approximate body length 12.0-12.5 m), were found. Groups of males about to attain maturity (body length 11.0-11.5 m) were encountered separately. The large males here were larger than in the tropical region but nevertheless constituted an insignificant minority. In the zone of temperate waters (40° latitudes) there were no imma- ture males or small whales measuring 7-8 m in body length. Mature males 12-13 m long predominated while large males constituted up to 20% of the total strength. In the Antarctic waters (south of the 50th parallel), only large males 14-15 m long or more were encountered. Females and young sperm whales of either sex were totally absent (Tormosov, 1970). Reproduction. Female sperm whales usually do not emerge beyond the warm and temperate waters and hence the seasons of mating and parturition among them are not as sharply manifest as among those whales whose females perform regular migrations into the cold waters of both hemispheres. Births occur among sperm whales throughout the year but the maximum number occurs in a relatively brief period. For the Northern hemisphere, this evidently is the early autumn months. In the North Atlantic, more births have been recorded from May through November with the maximum number taking place from July through September (Clarke, 1956). On the eve of parturition, females concentrate in quiet zones where conditions are more favorable for the newborn. Such zones in the Pacific Ocean are the waters of the Marshall and Bonin islands, eastern coast of Japan, and to a lesser extent the waters of the southern Kuril Islands and the Galapagos Islands; in the Atlantic Ocean these are the waters of the Azores and Bermuda islands and the coasts of the African provinces of Natal and Madagascar. Sperm whales gather in regions with clean deep 615 828 water оп the leeward side of islands or reefs (Sleptsov, 1955; Tomilin, 1957). In the Southern hemisphere, births occur from December through April with a maximum in February. They take place in regions of quiet and relatively warm waters, where sharks, killer whales, and other ene- mies of the newborn sperm whales are few. The optimum water surface temperature would seem to be 15-17°C. For the Atlantic and Indian oceans, such waters are found between 35 and 39° S lat. and it is here that the majority of female sperm whales undergo parturition. In April, 1962, close to Tristan da Cunha Islands, the process of par- turition was observed from a helicopter (F. Khomchik). Among several groups of sperm whales, numbering 25-30 animals each, one group of six was obviously isolated. These whales dived continuously together in a row, churning the water, which soon became blood-stained and a new- born whale was shortly sighted on the water surface. The newborn imme- diately began to swim alongside its mother. The two were accompanied by four other sperm whales, probably also females. Observers have noted that during birth the female assumes a vertical posture with almost one-quarter of the body length projecting above the water. The caudal flukes in the newborn remain coiled for sometime in the form of a small tube. Other adult females dive under the newborn calf and help it to remain afloat on the sea surface until the caudal flukes unfold (Tormosov, 1970). Gestation in the sperm whale is far more prolonged than in most of the other species of cetaceans. It has been assumed to extend for 16-18 months (Matthews, 1938b; Nishiwaki, Hibiya, and Ohsumi, 1958; Ohsumi, 1965). The ratio of male to female embryos is close to 1:1. Of the 1,068 embryos studied in the northern part of the Pacific Ocean, males accounted for 48.1%. In the waters of Chile and Peru, of 1,118 embryos, males were 56.2% and in the waters of Africa (491 embryos) 45.6%. Of the total of 2,677 embryos studied, males accounted for 51.0% (Ohsumi, 1965). Females usually deliver a single calf; twins are very rare (0.66%, Clarke, 1956). The periodicity of reproduction has been put at three (Clarke, 1956) or even three to four years (Ohsumi, 1965). Growth and development. Embryonic growth of the sperm whale con- tinues for up to 16 (probably even 18) months. Thus the growth of embryos proceeds more slowly than among other species of whales, which are characterized by a very short period of embryonic growth (Fig. 356). The growth pattern of sperm whale embryos is depicted in Table 61. The length of the newborn calves varies from 350 to 500 cm (Tomilin, 1957). The average size of the newborn from the North Atlantic is 392 cm (Clarke, 1956) and from the Southern hemisphere 415 cm (Lous, 829 ИРИ, ( 2 РСС р 94472 р 2222 2222222 777 = 7 Е a Се 77 4 Fe POO PA GLE tt AZZ eg Paar 7 614 Fig. 356. Embryonic growth of the sperm whale, Physeter catodon (figure by N.N. Kondakov). 615 Table 61. Growth of sperm whale embryos (in cm) (Ohsumi, 1965; Berzin, 1971) Month of growth Average length, according to Ohsumi Berzin 151 6 10 2nd 24 20 3rd 52 30 4th 76 50 Sth 103 70 6th 127 90 7th 155 5 8th 183 140 9th 210 170 10th 234 215 11th 262 260 12th 289 290 13th 314 320 14th 341 350 1518 369 380 16th 393 400 1959). The average size of calves, based on data for both hemispheres, is 405 cm (Ohsumi, 1965). The largest embryo from the North Pacific Ocean was 460 cm long (Berzin, 1964a). A newborn calf can weigh up to 1,000 kg. The duration of milk suckling, on analogy with the other species of large whales, was initially considered to be 6-7 months (Vinogradov, 616 830 1949; Sleptsov, 1955), later 13 months (Clarke, 1956), but according to the latest data, 24-25 months (Ohsumi, 1965). The age of the sperm whale has been determined by the number of layers in the dentine formed as a result of seasonal metabolic variations associated with changing conditions of habitation. Right up to the fill- ing up of the pulp cavity, two layers of dentine are laid every year: one very broad, light-colored, and intensely calcined, the other very narrow, dark-colored, and less calcined (Fig. 357). These two layers form a sin- gle annual ring whose number reflects the age of the whale (Nishiwaki, Hibiya, and Ohsumi, 1958; Berzin, 1961). The question of the number of layers laid in the course of a single year has not yet produced a com- monly accepted opinion. It is possible that the annual rings may be two, or a single annual ring may consist of different numbers of intermediate rings. This aspect continues to be studied. By the end of the first year, sperm whales of the North Pacific Ocean reach a length of 6 m or slightly more, 1.е., the increment in body length at the end of the first year is almost 2 m (all figures are average values). The growth tempo subsequently slows down somewhat and by the third year the animal reaches a length of 8 m, adding another 2 m in these two years. By age 3.5 -4 years, the female sperm whales have attained sexual maturity and their growth tempo subsequently is markedly slower than that of the males, with the annual increment not exceeding 50 cm. The onset of physical maturity and growth cessation among females occurs at the age of 15 years at a body length of 11 m. Females 11.7 m long are over 30 years of age and have already lost their middle pairs of Fig. 357. Tooth of the sperm whale, Physeter catodon. A—external view; B—longitudinal section (figure by N.N. Kondakov). 616 831 teeth. It may tentatively be assumed that at maximum age females have attained a length of 12 m or slightly more. Males attain sexual maturity at 5 years of age and a body length of 9.5 m but their growth tempo subsequent to reaching sexual maturity does not decrease initially. For the next few years the annual increment averages 80 cm, then gradually decreases thereafter. Growth cessation is noticed at the age of 23-25 years at a body length of about 16 m (Berzin, 1961, 1964a). The body length of male antarctic sperm whales at the time of attaining sexual maturity is 11.5-12.5 m because of the generally larger dimensions of whales of the Southern hemisphere (Matthews, 19386; Nishiwaki, 1955; see under “Geographic Variation’’). Under conditions of intense contemporary hunting practices, the maximum length of male sperm whales very rarely reaches 18 m and whales larger than this size are not found at all. Age determination revealed the oldest females to be 30 years (body length 10.7 m), over 30 years (body length from 10.9 to 11.7 m), and over 35 years (11.1 and 12.4 m); males were 27 years (15.6 and 14.8 m), 29 years (16.2 m), and over 30 years (14.8 m) or even 32 years. It has been assumed that age determination of animals older than 35 - 40 years is not possible since the pulp cavity is totally sealed. Be that as it may, this age is not a maximum for the sperm whale (Berzin, 1971). Enemies, diseases, parasites, mortality, and competitors. There is only one known case of a piece of the rostrum of a swordfish being found in the trunk of a beached sperm whale (Tomilin, 1957). Diseases have not been studied. Only dental caries, distortion of the lower jaw, and some other diseases are known. . A significant film of diatomaceous algae with Cockoneis ceticola predominating, has sometimes been observed on the body of the Antarctic sperm whales. In tropical waters minute ulcers have been detected on the skin of sperm whales, which form scars during the residence of the animals in the cold seas: the nature and origin of these ulcers are not understood. Crustaceans of Penella sp., common in rorqual, infect individual sperm whales occasionally. Barnacles are also rare and found in small numbers. They comprise three species of the genus Conchoderma (С. auritum, С. virgatum, and С. cuvieri). Coronula sp. has been encountered as a rare exception. The whale lice Cyamus physeteris and C. catodontis are common parasites of most sperm whales. They infest more often the urogenital and anal openings, less so the wrinkled flanks, and very rarely the dorsum (Clarke, 1956; Tomilin, 1957). Thirty-one species of helminths are known in sperm whales. These include trematodes 1, cestodes 12, nematodes 10, and acanthocephalans 8 species. The single species of trematode, found in the Kuril waters, 832 Zalophotrema curilensis Gubanov, parasitizes the liver of only the sperm whale. Of the 12 species of cestodes, 5 are found exclusively in sperm whales. Tetrabothrium curilensis Gubanov and Hexagonoporus physeteris Gubanov parasitize the small intestine and Tetragonoporus calyptocephalus A. Skr- jabin localizes in the bile ducts of the liver; all three have been found in Kuril waters. Multithuctus physeteris Clarke parasitizes the bile ducts of the liver and Polygonoporus giganticus A. Skrjabin the intestine in the Antarctic. The cestode Tetrabothrius affinis Lonnberg, found in the waters of Norway, Sourh Africa, New Zealand, and Antarctica, para- sitizes the intestine of the sperm whale and three species of baleen whales. Priapocephalus grandis Nybelin was found in the intestine of the sperm whale and four species of baleen whales in the Pacific Ocean. Phyl- lobothrium delphini Bosk, parasitizing the skin and found at many places in the Atlantic Ocean, the Mediterranean Sea, waters of the Commander Islands, Australia, and Antarctica, has been detected in the sperm whale, six Other species of toothed whales, Weddell seal, and the Arctic whale. Further, some cestodes not identified to species level, have been detected in the sperm whale. These are: species of Tetraphyllidae (larvae) in the subcutaneous adipose tissue, Diplogonoporus sp. in the bile ducts of the liver, Trigonocotyle sp. in the small intestine of the sperm whale and Baird’s beaked whale, and Trypanorhyncha sp. (larvae) in the stomach of the sperm whale, sei whale, Minke whale, and Steller’s sea lion. Of the 10 species of nematodes, 5 are found exclusively in the sperm whale. The stomach parasite, Anisakis (Anisakis) catodontis Baylis, has been detected in the waters of South Africa; Anisakis (Skrjabinisakis) physeteris Baylis in the North Pacific Ocean, waters of South Africa, and Antarctica; and Anisakis (Anisakis) ivanizkii Mosgovoy around the Com- mander Islands. Placentonema gigatissima Gubanov was found in the pla- centa (Kuril Islands). Anisakis (Skrjabinisakis) skrjabinit Mosgovoy раг- asitizes the stomach and small intestine of the sperm whale and spotted dolphins (Commander Islands, Sea of Okhotsk, and Antarctica). Anisakis (Anisakis) dussumierit Beneden, parasite of the stomach and large intes- tine of the sperm whale and spotted dolphins, was detected in the waters of Japan and the Commander Islands. Anisakis (Anisakis) simplex Rudol- phi, found in the sperm whale and widely distributed in marine mam- mals (North Sea, eastern Kamchatka, Japan, and New Zealand) para- sitizes the gullet, stomach, and intestine. In the northwestern part of the Pacific Ocean, Anisakis (Anisakis) pacificus A. Skrjabin was detected in the stomach of the sperm whale, killer whale, and fin whale. More- over, Anisakidae g. sp. and Tetrarhynchidae m. sp. (not identified to the species level) also parasitize the sperm whale. 618 833 Two species of acanthocephalans parasitize the intestine only of the sperm whale: Corynosoma curilensis Gubanov was found off the Kuril Islands and Corynosoma mirabilia A. Skrjabin in Antarctic waters. Corynosoma strumosum Rudolphi, parasite of the intestine, is known only in belugas, common porpoises, 11 species of pinnipeds, and land mammals and birds. Bolbosoma physeteris Gubanov was detected in the intestine of the sperm whale and the killer whale in the waters of the Kuril Islands. Bolbosoma capitatum Linstow parasitizes the intestine of the sperm whale and two more species of toothed whales from the Atlantic Ocean and the Mediterranean Sea. Bolbosoma brevicolle Malm, detected in the large intestine and rectum of the sperm whale, is also known in five species of baleen whales; it has been found in the North Atlantic, in the waters of South Africa, and near South Georgia Island. Bolbosoma turbinella Diesing parasitizes the intestine of the sperm whale, beaked whales, and five species of baleen whales from the Atlantic and Pacific oceans of the Northern and Southern hemispheres. Finally, Bolbo- soma tuberculata A. Skrjabin, detected in the South Atlantic and Indian oceans, parasitizes the intestine of the sperm whale, sei whales, and Bryde whales (Margolis, 1954; Delamure, 1955; Margolis and Pika, 1955; A. Skrjabin, 1958, 1959, 1960, 1961, 1970; Berzin, 1971). Natural mortality of the sperm whale has not been studied nor have the reasons for the death of besched whales been ascertained. The sperm whale has almost no trophic competitors with the possible exception of members of the family of beaked whales. Population dynamics. Population variations in sperm whales due to natural factors are not known but due to hunting are significant. Hunting of sperm whales using manual harpoons and sailboats commenced in the first half of the eighteenth century and continued over the next one hun- dred years on an extensive scale. By the mid-nineteenth century, sperm whale reserves had shrunk markedly in all the seas; hunting became unprofitable and almost ceased. This promoted a significant restoration of the reserves of the sperm whale. The second stage in sperm whale hunting commenced with mod- ern techniques using whaling fleets and reached a high magnitude. As a result, by the 1960s there was a sharp reduction in the population of sperm whales in some regions, accompanied by a reduction in the aver- age size of the animals caught, increase in the relative number of much younger animals in the catch, and some other features of diminishing reserves. This situation was mainly noticed in the waters of Japan and the Kuril Islands and to a lesser extent in the other regions, including the Antarctic where only adult male sperm whales that have not participated in reproduction are known to gather. 834 Field characteristics. The huge angular head occupies over one-third of the body length. The blowhole in the sperm whale is at the end of the snout on the left side and not on the upper part of the head, as in all other whales; hence the blow of this animal is not vertical but set forward to the left at an angle of roughly 45° to horizontal. The blow is broad, “bushy,” with a height of not more than 3-4 m. The sperm whale can be recognized with certainty from a distance by the nature of the blow. The body color is monochromatic, usually dark. While diving to a depth it often exhibits the caudal flukes. After prolonged submergence, it lies on the water surface almost without movement and allows the very close approach of ships. (V.A.) Economic Importance The development of sperm whale hunting intensified mideighteenth cen- tury following the rapid decline of the catch of right whales whose’ herds had been destroyed. Hundreds of sailboats, working mainly in the tropical waters of all the three oceans, 1.е., predominantly in the reproduction sites of sperm whales, were engaged in hunting them. This hunting proved highly profitable since the demand for spermaceti was inexhaustible and its price very high. Spermaceti was used mainly in the candle-making industry: candles made from it burn with a bright flame and without soot. By the mid-nineteenth century, kerosens was being used for illumination and the demand for spermaceti fell sharply. By this time a perceptible reduction in number of sperm whales had also made hunting unprofitable and it almost ceased circa 1860. The let up in hunting of the sperm whale continued for about half a century. Early in the twentieth century, the hunting of sperm whales (mech- anized by now) was resumed. It was particularly well developed in the North Pacific Ocean, along the coasts of South America, and later even in the Antarctic. Following the reduction in the population of the main commercial species of baleen whales, the importance of the sperm whale rose gradually in all the regions of the World Ocean although not to the same extent everywhere. In the North Atlantic Ocean the sperm whale is of utmost commer- cial importance in the southern part of the range. Thus in the Azores it represents a unique commercial species of whale. From the end of the 1930s to the end of the 1950s, 500 to 700 sperm whales were caught annu- ally. From the beginning of the 1960s, the volume of hunting decreased somewhat—from 400 initially to 300 whales later to 145 in 1969. Around Madeira Island, too, the sperm whale was caught exclusively but the mag- nitude of hunting was relatively small. Commencing from the early 1940s, 619 835 the catch did not exceed 200 a year and after 1967 did not reach even 100. This species is of some commercial importance in the whaling industry of Spain, Portugal, and the northern part of western Africa where sperm whales have accounted in some years for 30 to 80-90% of the 200 to 300 whales caught annually. In the northern waters these whales are of some importance in Norway and Iceland although the number caught is small. On the Norwegiian coasts no more than a hundred sperm whales are caught per year, or 15 to 40-50% of the total catch; they numbered 120-170 animals per year or 25 -45% of the total catch in the waters of Iceland from the end of the 1950s to the mid-1960s. At the end of the 1960s, the annual catch was 100 sperm whales. The more abundant sperm whale population of the North Pacific Ocean ensures a larger catch. The catch is particularly high in the coastal stations of Japan and from large floating whaling stations. Up to 1950, less than 1,000 sperm whales were caught off the coasts of Japan, which represented about hair the total catch of whales of all species. From 1950, hunting activity rose considerably and the number of sperm whales caught ranged between 1,200 and 2,600 animals per year. In 1968, the sperm whales caught here numbered 3,747 and in 1969—3,668. In some years sperm whales accounted for 60-70% or even more of the total catch. Until the mid-1950s, when the “pelagic” fleet was not large, the catch of sperm whales usually did not exceed 500-700 animals per annum. But from the mid-1950s, with the introduction of new Soviet and Japanese whaling fleets, it began to rise rapidly: initially up to 2,000 and then 4,000, reaching almost 8,000 sperm whales a year by the early 1960s. Between 1965 and 1969, the number of sperm whales caught totaled 10,500 to 12,500 per year or 50-70% of the total catch of the fleet. Sperm whales caught from the coastal bases on the Kuril Islands between 1950 and 1962 totaled 1,400 to 2,000 (70-80% of the total catch). The whaling industry of the Kuril Islands came to a halt in 1964. Large-scale hunting of sperm whales in the northeastern part of the Pacific Ocean also ceased. From the coastal bases of British Columbia, from the end or the 1940s to the early 1960s, 120 to 320 sperm whales were caught per year but whale hunting ceased in 1967. Off the coasts of California, the catch never exceeded a few tens (sometimes up to a hundred) of sperm whales per year. The sperm whale catch in the Southern hemisphere has been large with several areas participating in the hunt: waters of Chile, Peru, South Africa, pelagic hunting in the Antarctic, and, in recent years, in the more northern waters. In the African waters (Natal and Cape Province) whale hunting was resumed in 1948 and 400 to 1,000 sperm whales of a total of 1,200 to 2,700 whales were caught every year until 1956. From 1957, 620 836 the catch of sperm whales increased considerably and 1,200, 2,500, and even 3,600 sperm whales were caught every year (50-70% of the total catch). Off the coasts of Chile and Peru, sperm whales represented the main species of whales hunted and accounted for 70.9 and even 100% of the catch. The maximum number of these whales caught from the mid- 1950s to the mid-1960s ranged from 2,500 to 5,000 animals per year. At the end of the 1960s, hunting fell slightly to less than 2,000 animals. After the Second World War, the “pelagic” whaling fleet of the Antarctic developed rapidly and caught a large number of sperm whales (excep- tionally large males). But, of the 2,000-4,000 to 6,000-7,000 animals caught, sperm whales in a season constituted only 10-15% and later 20-22% of the total catch of the whaling fleet. From 1967 to 1969, the catch of sperm whales in the Antarctic fell to 2,500 to 2,600 animals per year but the Antarctic whaling fleet began catching these whales north of 40° S lat. along the course into Antarctica and back. This catch accounted for 2,000 or more sperm whales in a season. In Australia hunting of the sperm whale commenced only in the mid-1950s, following a reduction in the population and subsequent total ban of hunting of humpback whales (these were the lone target of hunting in these waters). From the mid- 1960s, the number of sperm whales caught in Australian waters totaled 500 to 600 per year. Modern hunting of sperm whales, like that of other large whales, is carried out in special steel ships equipped with harpoon guns. The whale is killed with a harpoon with a grenade screwed to its tip and primed with gun powder. The grenade fires 4 sec after shooting and by this time the harpoon has penetrated the whale body (Fig. 358). The firing can be fatal or otherwise depending on the resistance of that part of the trunk struck by the harpoon. The wounded whale is dragged toward the boat by a rope tied to the harpoon, using a special winch, and the animal finally killed by shooting a second (or third) time. Instances of a whale requiring up to six shots are known. The behavioral aspects of the sperm whale make for some typi- cal features in hunting them. After prolonged submergence, the sperm whale usually remains a long time on the sea surface. This facilitates the approach of the ship within shooting range while the animal’s attempt to escapeton a straight course enables the chasing whale boat to catch up with it quickly. Thus most of the sperm whales sighted from the ship can be killed. But if the marked sperm whale succeeds in diving before the ship can approach, it is futile to wait for it to surface again. Air is pumped into the body cavity of the killed sperm whale so that the carcass floats and can be towed to the floating base or the coastal station for dressing. First the blubber is removed in layers and then the 620 621 837 Fig. 358. Harpooned sperm whale (photograph by V.A. Arsen’ev). head is severed from the trunk. Using a steam or electric hoist, the large stock of fat in the spermaceti sac is recovered, sliced into large lumps, then charged into a boiler for melting the spermaceti. Then the viscera are removed from the carcass and the flesh from the bones. The bones of the skeleton and skull are cut by steam-powered saws into bits that can be fed into fat-melting boilers to ensure more complete conversion into oil. The main product obtained from the sperm whale (Table 62) is the oil from the blubber, bones, and viscera. The physical and -chemi- cal properties of the oil of the sperm whale differ from those of baleen whales (low specific gravity, low saponification number, low iodine num- ber, etc.). The oil is used for commerical purposes by various industries. A large male (length 14-16 m), on average, yields about 10 tons of fat. Spermaceti has a characteristic chemical composition and is collected and processed separately from the blubber. Spermaceti is used by per- fumeries in preparing cosmetic media (creams and pomades) and to a lesser extent in other industries as well. The liver of the sperm whale contains the maximum amount of vitamin A compared to the liver of all other species of whales: about 6,000 units, on average, versus 1,500 units in a female fin whale (Mrochkov, 1953). The mechanical method of pressing the subcutaneous fat layer yields more oil than digesting in a boiler (Dormenko, 1952; Zaikin, 1953). In the former method the upper 838 layers of the integument can be used in making leather goods. For this purpose two layers of 5-6 mm thickness each are taken from the carcass of the whale and a third additional layer up to 8 mm thick from its head. Much deeper layers can also be used in the leather industry but only after special compaction using special fillers. Table 62. Weight of the body parts of the sperm whale, kg (Zenkovich, 1937b; M.V. Ivashin) Body part Far East (Kronotsk Bay) Antarctica Male 13.5 m Male 18.0 m Male 16.1 m Subcutaneous fat 4,955 12,663 11,450 Spermaceti 620 5,757 4,400 Spermaceti sac 3,981 10,343 8,665 Flesh 5,640 7,697 7,850 Skull 1,800 6,087 4,100 Lower jaw 320 880 1,050! Vertebral column 1,986 3,387 3,4552 Caudal flukes 314 DS Ribs 965 2,147 3,2903 Flippers 485 978 Pelvic bones 108 200 Sternum 209 329 Viscera 1,282 2,199 3,340 Total 22,665 53,365 47,600 Liver — — 620 Liver and kidney 331 368 == Heart 113 211 — Lungs 161 317 — Lungs, heart, neck, and gullet — — 1,300 Stomach and intestine — — 700 Intestine 254 465 — Testes 10.5 16.5 — Penis 88 106 Remaining parts 323 716 720 1Lower jaw together with tongue. Vertebral column together with caudal flukes. 3Ribs together with flippers. Length of whale, m Area of raw skin, т? 8.9 19.2 9.5 25.5 10.5 28.4 11.0 33.0 14.5 55.5 15.3 71.0 622 839 Soles made from the skin of the sperm whale are not inferior (some- times even superior) in wearing properties to those made from cattle hide. However, the skin of the sperm whale always suffers from defects, such as flaws of 5-10 mm in diameter and scars and scabs sometimes up to 50 cm? or more in area. As these damages are randomly scattered all over the skin, cutting is difficult and much wastage results (Bulgakov et al. 1954). Hence the use of sperm whale skin in making leather goods has not developed. The lower layers of the skin, after mechanical degreasing, can be used in making gelatin and glue (Kolchev, 1954). The teeth of the sperm whale are excellent material for carved articles (chess pieces, handles, etc.) and almost indistinguishable from those made of ivory. Sperm whales carry ambergris in the form of a hard waxlike substance present in large lumps of indeterminate form. Crude ambergris varies from gray to black and smells like wet soil. It has a granular and layered structure. Its specific gravity varies from 0.730 to 0.780; it softens from the warmth of the hand, melts at 60°C without bubbling, and volatilizes at 100°C. It dissolves well in hot alcohol and burns with a light blue flame and slight resinous smell. Although ambergris is known from ancient times, its origin has yet to be ascertained. It is sometimes found floating on the water or cast on seashores. In the remote past it was regarded either as refuse of birds or as an exudation of plants and roots of trees. Later, the remains of chitioous “beaks” of cephalopods consumed by the sperm whale were detected in the ambergris, which suggested its origin from the intestinal tract of the whale. Its formation is thought to be a reaction to infection by parasites inhabiting the intestine or a product of the normal secre- tion of the rectal glands. The third point of view would appear to be more correct, according to which ambergris is formed as a result of the retention of undigested remnants of “beaks” of cephalopods, which are enveloped in special secretions in the digestive tract of the sperm whale. Depending on its quality, ambergris costs 150-200 rubles per kg. The production of synthetic ambergris has somewhat affected the cost of the natural product but the quality of the synthetic is inferior to the natural. In olden times ambergris was used as an antispasmodic and in Chinese medicine as a stimulant or antiseptic. The chief value of ambergris lies in its exclusive property of absorbing odor and exceptional retention of it. Therefore, natural ambergris is used in producing perfumes of superior quality. Ambergris is found in the intestinal tract of sperm whales in lumps weighing several hundreds of grams to tens of kilograms. Remarkably large lumps of ambergris have also been found: one lump weighing 392 kg was recovered in the Azores and another weighing 420 kg off the coast of 623 840 Australia. In our country a lump of ambergris weighing 351 kg was recov- ered in 1958. In subsequent years, ambergris production has exceeded 3 tons (Ivashin, 1966). Ambergris as a raw material requires sometime for maturation and hence all varieties of ambergris do not command the same price and some are even totally unsuitable for use in perfumeries. Following the conclusion of the International Whaling Convention, 1946, some restrictions were imposed on the hunting of sperm whales. According to the international hunting rules, killing of sperm whales less than 10.7 m long was prohibited from the coastal whaling stations and less than 11.6 m long from the floating fleets, even though many sperm whales of these sizes are mature animals. In spite of these restrictions, highly intense sperm whale hunting continued and enormous catches led to a steady depletion in numbers, especially in the better exploited populations. Now a restriction has been imposed on killing sperm whales by prescribing annual quotas in all parts of the World Ocean, taking into consideration the strength of individual populations. In most cases, it would be extremely useful to restrict the killing exclusively to male sperm whales which, given the prevailing polygamous mode of life of these whales, could not adversely affect the normal replenishment of the various populations. (V.A.) Subfamily of Dwarf Sperm Whales Subfamily KOGIINAE Gill, 1871 Genus of Dwarf Sperm Whales Genus Kogia Gray, 1846 1846. Kogia. Gray. Zoology of the Voyage of H.M.S. Erebus and Terror, I, p. 22. Physeter breviceps Blainville. : 1876. Cogia. Wallace. Geogr. Distr. Anim., 2, p. 208. Correction of Kogia Gray. Nom ргаеосс. (V.H.) Dimensions the smallest in the family, with a total length up to 3.4 m. The body form is dolphin-shaped. The head is small, proportional, constitutes 1/8th to 1/6th of the body length, and is rounded anteriorly. The spermaceti зас” is well developed on the head but much smaller than in the sperm whale. The blowhole is situated on the parietal section of the head, almost at its center. The blowhole opening is crescent-shaped. The end of the lower jaw falls far short of the anterior end of the head. The fairly high dorsal fin lies roughly midbody. The flippers are relatively 5 See description under sperm whale, р. 799. 625 841 narrow and short. The body is dark-colored dorsally and light-colored ventrally. The rostrum is broad at the base, constricted sharply in the anterior part, and is shorter than the cranium. The depression on its dorsal sur- face is small and has a longitudinal crest formed by the premaxillae and maxillae. The nasal bones are not fused with the premaxillae. The max- illae have very large preorbital processes. The zygomatic bone does not articulate with the squamosal. The symphysis of the lower jaw is shorter than half its length. Teeth: 8-16 pairs on the lower jaw and 1-3 pairs in the anterior section of the upper jaw (but sometimes totally absent in the latter). They are thin, curved, and the sharp tip set posteriorly. Enamel is absent. Vertebrae 52-57: cervical 7, thoracic 12-14, lumbar 9-12, and cau- dal 21-27. The cervical vertebrae are fused. The flattened scapula has a small acromion and large coracoid processes. Ribs 13-14 pairs. The sternum is short. The phalangeal formula in the forelimbs is subject to individual variation: I,, Пб-з, II,-3, [У4 -з, and V>-7. The population of the dwarf sperm whale is small. Biology has not been well studied. It lives singly or in small groups and apparently feeds mainly on cephalopods. The periods of mating and parturition are pro- tracted. Gestation continues for about nine months. Females evidently reproduce annually. They live in the warm waters of the Indian, Atlantic, and Pacific oceans (Fig. 359). There is no special hunting of dwarf sperm whales in view of their rarity. The genus comprises two species (Handley, 1966): 1) dwarf sperm whale, Kogia breviceps Blainville, 1838; and 2) Owen’s dwarf sperm whale, Kogia simus Owen, 1866. Dwarf sperm whales are not observed in USSR waters but summer transgressions are possible (especially of Owen’s dwarf sperm whale) into the Sea of Japan and toward the southern part of the Kuril Islands. (V.S.) DWARF SPERM WHALE Kogia breviceps Blainville, 1838 1838. Physeter breviceps. Blainville. Ann. Franc. étrag. d’Anatomie Physiol., 2, p. 337, pl. 10. Cape of Good Hope. (V.H.) Diagnosis Dimensions largest in the genus, with the overall body length varying from 2.7 to 3.4 m. The relatively low dorsal fin lies slightly posterior to “(лэчэзту УЛ) 21801 ‘зэеЧм шлэ4$ улемр Jo зпиэ8 ay) Jo э8иез ‘66$ “314 051 06 тот ost Е , з A! ae = ON = see = = `. а т >... O} Е BST. в == хх = г. z ee, => 3 7 у = 79 625 843 midbody. Teeth are absent in the upper jaw. Mandibular teeth 12-16 pairs (less frequently 10-11 pairs). (V.S.) Description The body dorsally, including the caudal flukes and flippers is dark, almost black. The flanks are light gray and the abdomen light-colored. Poste- rior to the eyes and anterior to the flippers, one or two light-colored projections run bottom upward into the dark-colored region (Fig. 360). Light-colored spots are visible on the ventral surface of the caudal flukes. The body dimensions of adult male and female dwarf sperm whales from New Jersey and Texas states (USA) (Tomilin, 1957) are respectively (in cm): body length 299 and 274; distance from tip of snout to center of blowhole 32 and 35, up to anterior margin of dorsal fin 154 and 150; distance from anal opening to notch between caudal flukes 97; length of flippers 41 and 43; height of dorsal fin 9 and 13; length of dorsal fin 14 and 41; and width of caudal flukes (between apices) 50. The animals range in weight from 318 to 408 kg. The condylobasal length of the skull (Fig. 361) is 391-469 mm. The mandibular symphysis is long (8.6- 12 cm), with a keel on the ven- tral surface. The alae of the pterygoids and basioccipitals are elongated. The foramen magnum is situated roughly midpoint of the skull height. (V.S.) Geographic Distribution and Biology Inhabits predominantly the warm belt of the Pacific Atlantic, and Indian oceans. Distribution has been studied mainly from the finds of animals cast on coasts or shoals. More than 15 finds of beached dwarf sperm whales are known on the coasts of South Africa, Australia, and New Zealand, and over 50 on the Atlantic coast of the USA. Finds in the Pacific Ocean are not rare near Japan, the East China and South China seas, Gulf of Siam, along the coast of North America (Washington and Fig. 360. Dwarf sperm whale, Kogia breviceps (figure by N.N. Kondakov). 625 844 Fig. 361. Skull of the dwarf sperm whale, Кода breviceps (figure by М.М. Kondakov). 626 California states), and along the coasts of Mexico, Peru, Chile, Fiji Islands, Australia, Tasmania, and New Zealand. In the Indian Ocean, it is distributed from southern Australia to South Africa. In the Atlantic Ocean, beached animals are known on the coasts of North America from Nova Scotia to Florida and Texas, and on the coasts of South Africa, France, and Holland. In the Atlantic Ocean, these whales probably move into the open sea even in latitudes more northern than Holland (Dell, 1960; Tomilin, 1962; Gaskin, 1966; Hershkovitz, 1966; Handley, 1966). The residence of these animals in the waters of the Soviet Union has not been established. Transgressions are possible in the summer months into the Sea of Japan and the southern part of the Kuril range. Geographic variation of the species has not been studied. Information on the biology of dwarf sperm whales is very fragmentary. Remains of cephalopods (Sepia officinalis and others) have been detected in their stomach. Cephalopods apparently serve as their main food. Bones and otoliths of fish (Тисйо4оп sp. and others), 845 crabs (Carcinides maenas), shrimps (Paciphola pacifica, Pandalus sp., Pandalopsis sp., Penaeus sp., Hymenodora sp.) have been found less often. The period of mating is greatly prolonged and newborns have been encountered even in November and February; however, the majority of these whales probably mate in summer. Gestation is thought to extend for 9-10 months. The length of a newborn calf is around 110 cm; whales 153-171 cm long were still suckling and a female with a body length of 188 cm was immature. Females attain sexual maturity at a body length of 218-219 cm. A female about 3 m long, beached on a Dutch coast in December, 1925, contained an embryo 20 cm long. Females give birth to a single calf. An instance is known of a female, accompanied by a small calf, who was simultaneously pregnant and lactating. Dwarf sperm whales live singly, in pairs, or in small groups of up to five animals. Their movements are slow. Penela sp. was found among the ectoparasites. Larvae of the cestodes Monorygma grimaldii Monier and Phyllobothrium delphini Bosk and the nematodes Crassicauda magna Jonston and Mawson and Pseudoterra- nova mogiae Jonston and Mawson, have been detected in the abdominal cavity and subcutaneous tissue. A female 277 cm long weighed 369 kg while large males НЫ up to 500 kg. There is no special hunting for these whales. The Japanese industry while catching various species of dolphins occasionally nets dwarf sperm whales also. Although their flesh is edible, these whales are not of economic importance (Delamure, 1955; Tomilin, 1957, 1962; Nishiwaki, 1965; Handley, 1966). (V.A.) OWEN’S DWARF SPERM WHALE Kogia simus Owen, 1866 1866. Physeter (Euphyseter) simus. Owen. Trans. Zool. Soc. London, 6 (Г), р. 30, pls. 10-14. (V.A.) Diagnosis Dimensions the smallest in the genus, with overall length ranging from 2.1 to 2.7 m. The relatively high dorsal fin is roughly midbody. The upper jaw usually has 1-3 pairs of teeth and the lower 8-11 pairs (less often, 13 pairs). (У.5.) Description The body is dorsally dark gray and ventrally white. 627 846 The body measurements of a male and two females from Japan (Yamada, 1954) are: overall length 2.22 m; distance from tip of snout to blowhole 16.3, 20, and 20 cm, from tip of snout to apex of flippers 78.5, 87, and 86 cm; length of base of dorsal fin 34, 33, and 42 cm; height of dorsal fin 13, 11, and 17.5 cm; and distance from caudal notch to anal opening 68.5, 82, and 82 cm. The weight of the animals varies from 136 to 272 kg. The condylobasal length of the skull is 262-302 mm. The mandibular symphysis is short, 37-46 mm, and smooth on the ventral surface. The alae of the pterygoids and basioccipitals are short. The foramen magnum lies considerably below midskull height. The skull measurements of a male and two females from Japan (Yamada, 1954) are (in mm): condylobasal length 271, 284, and 297; length of rostrum 140, 165, and 160; width of rostrum at base 128, 136, and 140; interorbital width 218, 240, and 259; and length of lower jaw 224, 253, and 255. (V.S.) This species is encountered in the coastal waters of South Africa, southern Australia, India, Sri Lanka, Hawaiian Islands, Japan, and off the east coast of the USA. It is known predominantly from beached animals. The maximum number of beached animals has been found on the coasts of Japan and on the eastern coasts of the USA (Handley, 1966). Geographic variation has not been established. Biology has not been studied. Differences between the dwarf sperm whale and Owen’s dwarf sperm whale are not widely recognized and hence some data on the biology ascribed to Kogia breviceps may actually pertain to Kogia simus. (V.A.) Family of Beaked Whales Family ZIPHIIDAE Gray, 1865° Body dimensions medium and large. Males are larger than females or vice versa. The body is more or less spindle-shaped. Anteriorly, the snout forms a narrow “beak” sharply demarcated from the frontal adipose body among bottlenose and beaked whales or gradually rising above the head in the rest of the members of the family. The blowhole is crescent-shaped and its bulge may face the head or the tail. The small dorsal fin is situated at the level of the anal opening or slightly anterior to it. There is no notch 6 Hyperoodontidae, according to some authors. 629 847 between the caudal flukes ог it is very small. The flippers are low. The body is usually monochromatic brown or gray but somewhat lighter on the ventral side. The rostrum is long and narrow, longer than the cranium, with an open mesorostral groove (closed in old animals). The maxillae, premax- illae, and frontal bones bear crests. The palatine bones form part of the anterior wall of the nasal passage. The petrosal bones are fused with the skull. The lachrymal bones are large and not fused with the zygo- matic bone. The pterygoids are large and separated by some distance. The lower jaw is wider than the upper and projects forward slightly. Numerous teeth are seen in the embryos, which reduce in number subsequently. The adults of most of these species have just a few teeth only in the lower jaw. The number of functional teeth varies from г in Tasmacetus to § in Berardius and $ in the rest of the genera. The number of vertebrae does not exceed 50 and from 2-7 cervical vertebrae are fused. Ribs do not exceed 10 pairs and their sternal sections are not ossified. The system of air sacs is simple, without preorbital and postorbital lobes. In adult animals, rudiments of the olfactory nerves are preserved. The stomach comprises 4-14 chambers. The members of this family feed mainly on cephalopods (teuthophagous) but also consume fish. They are capable of diving deep and remaining submerged for a long time. Some perform annual migrations. They live singly or in herds, usually comprising 10-15 animals. These animals are distributed in the warm, temperate, and cold waters of the World Ocean (Fig. 362). The oldest of the Ziphiidae is evidently Notocetus Moreno from the Lower Miocene of Patagonia, resembling in many features the Miocene Squalodon Grateloup (family Squalodontidae Brandt) and having, in par- ticular, numerous teeth. Reduction of teeth among the primitive Ziphi- idae occurred in the Miocene and typical members of contemporary beaked whales appeared at the end of this period. The family consists of 17 genera, of which 5 are extant: Tas- macetus Oliver (Tasmanian beaked whales), Berardius Duvernoy (Pacific beaked whales), Mesoplodon Gervais (“sword-tooth dolphins’), Ziphius G. Cuvier (Cuvier’s beaked whales), and Hyperoodon Lacépéde (bottlenose whales). The generic division of the family has given rise to no doubts among researchers. However, the systematics of the species of beaked whales requires further development (especially of genus Mesoplodon). About 15 species are usually recognized in the family. Beaked whales are of little economic importance. | ues ЩИ ИЗ ‘НИЦ АИ | I x | ИН и at pa ты © "| is Е | MTA A ий |} | | | [ИН Ш | in) AAT | UPL La MUAH) ПИ Mil iil! 9. «ар. 1 | НИНА О ИИ (4 yi Ш ЩИ ИИЦ 1000 0 1000 2000 3000 4000 5000 Кт и Н | Fig. 362. Range of the family of beaked whales, Ziphiidae (V.A. Arsen’ev). 628 849 All of the genera, except Tasmacetus, are encountered or could be encountered in USSR waters. Our hunting of these animals is in small numbers. (V.S.) Genus of Pacific Beaked Whales Genus Berardius Duvernoy, 1851 1851. Berardius. Duvernoy. Ann. Sc. Nat., Zoologie, 15, p. 52. Berardius атоихи Duvernoy. Dimensions large, the largest in the family. The frontal projection is quite high. The “beak” is highly elongated and somewhat flattened. The low dorsal fin is situated above the anal opening and is not flexed along the posterior margin [falcate]. The flippers are relatively short. The lower jaw is slightly longer than the upper one. The color is dark brown, the under varying from grey to whitish. Asymmetry of the skull bones is less manifest than among the other members of the family Ziphiidae. The rostrum is narrow, its length more than double that of the cranium, and it is slightly flattened dorsoventrally. The maxillary crests are low. The premaxillae are symmetrical and their width almost equal throughout their length. The suture between the _ ргетахШае and maxillae does not extend beyond 20% of its length on the dorsolateral surface of the rostrum. The frontal bones have broad orbital processes. The large, massive, rounded, and almost identically sized nasal | bones project markedly into the frontals and occupy the highest position on the skull. The mesethmoid bone is only partly ossified. The anterior section of the lower jaw has two pairs of teeth, of which the anterior larger ones are located at the tip of the lower jaw and remain exposed even when the mouth is closed. The posterior pair of teeth emerge 10-20 cm behind the anterior ones. A few more teeth may remain concealed in the gums. Cervical vertebrae 7, thoracic 10-11, lumbar 12, and caudal 17-19; total 46 - 49. The three anterior cervical vertebrae are fused. The sternum consists of five sections. Biology has not been well studied. These animals remain confined in groups of roughly 20. They feed mainly on cephalopods. Mating and parturition are protracted. Gestation continues for 10 months. These animals are encountered in the Pacific and Indian oceans and in the South Atlantic. The genus consists of two species: Arnoux’s beaked whale (southern beaked whale), Berardius arnouxi Duvernoy, 1851, and Baird’s beaked whale (northern beaked whale), Berardius bairdi Stejneger, 1883. 630 850 Only Baird’s beaked whale is encountered in USSR waters.’ There is no special hunting for Baird’s beaked whale; it is caught only incidentally. (У.5.) BAIRD’S BEAKED WHALE® Berardius bairdii Stejneger, 1883 1883. Berardius bairdi. Stejneger. Proc. Ц. 5. Nat. Mus., 6, р. 77. Staraya Gavan’, east coast of Bering Island. 1883. Berardius vegae. Malm. Bihang Svenska Vet. Acad. Handl. 8 (4), p. 109. Commander Islands. 1947. Rostrifer nestorésmirnovi. Zenkovitsh (Zenkovich). Zh. “Rybnoe Khozyaistvo,” No. 10, p. 15. Nomen nudum. 1955. Berardius vegana. Bourdelle and Grassé. Traité de Zool., 17, р. 429. Erroneously for vegae Malm. (V.H.) Diagnosis Only species of the genus found in waters of the USSR. Description | The head is relatively small (Fig. 363). The “beak” is flattened dorsoven- trally. The “forehead” of males rises steeply upward and is set off from the “beak” by a distinct transverse groove (Fig. 364). In females the fore- head rises more gently and the transverse groove is absent. The blowhole is crescent-shaped and its bulge faces forward. Two (less often, three or four) deep longitudinal grooves occur on the throat (distinctly visible even in embryos; Fig. 364). The shape of the dorsal fin is variable—from low humplike to relatively high and well-proportioned. It is two or three times longer than its height. The flippers are placed close to the ventral surface of the body. They are quite short and broad (length only 2.5 -3.2 times their width). The caudal flukes have a small notch. The body color turns dark with age. In young animals the upper portion is gray with a brown tinge and the abdomen and flanks somewhat 7 Вана’; beaked whale, caught by our whalers in the Far East, was long erroneously classified as a bottlenose whale (Hyperoodon). Only after 1950 (Tomilin, 1952, 1957) was it demonstrated that these animals were actually Baird’s beaked whales. Thus all the infor- mation about bottlenose whales in the Pacific Ocean reported until 1956 pertains to this species. 8 Sometimes, especially in the earlier literature, and also in foreign publications (Her- shkowitz, 1966), also referred to as the “Pacific beaked whale”. (V.H.) 630 631 631 851 777.7 ae 72 LZ РРР 2227 Е 3 22.25... ИИ АР LLY SEE 7777 —_ Я LP = LE LZ РРР ЕЕ 2 И: РРР = и Be 5 CALLE iy ico и р Fig. 363. Baird’s beaked whale, Berardius Бата (figure by М.М. Kondakov). Fig. 364. Head of Baird’s beaked whale, Berardius bairdi, front view. Pacific Ocean (photograph by M.M. Sleptsov). lighter. In adult animals the dorsum and flanks are dark brown and the abdomen slightly liger in color (Sometimes may be white). White spots are seen on the skin surface of adult whales in the navel region between the flippers and under the throat. It is possible that these spots reflect an age-related variation of skin pigmentation (Tomilin, 1957). White scars (probably caused by bites of males) are more numerous on the dorsum than on the abdomen. Both the pairs of teeth occur in the region of the mandibular symphysis. The anterior teeth are set on the very tip of the jaw and may rise to a height of 79-89 mm and the posterior ones to 53-67 mm (Omura et al., 1955; Tomilin, 1957). Rear teeth are usually absent in females. In the newborn and suckling calves the teeth are not erupted. In young animals the anterior teeth are conical and may be longer than in adults. With advancing age, the teeth wear down somewhat and become more massive and flattened laterally, especially that part in the alveolus. Barnacles of the genus Conchoderma sometimes colonize on the anterior teeth. 852 631 Fig. 365. Head of Baird’s beaked whale, Berardius Бат, ventral view. Pacific Ocean (photograph by M.M. Sleptsov). 632 Fig. 366. Baird’s beaked whale on a deck. Pacific Ocean (photograph by М.М. Sleptsov). The main measurements of four adult females (Tomilin, 1957) are (in cm): body length 1,225, 1,080, 1,095, and 1,090; distance from tip of upper jaw to blowhole 132, 120, —, and —; distance from anal opening 853 to posterior edge of caudal flukes 355, —, 323, and 324; length of flippers 152, 108, 118, and 125; height of dorsal fin 30, 30, 31, and 33; and width of caudal flukes (between apices of lobes) 310, —, 270, and 268. The dimensions of males are less than those of females. The maximum known length of a female is 12.2 m, of a male 11.9 m. The measurements of the skull (Fig. 367) of two adult females and a young male Baird’s beaked whale (Omura et al., 1955 and Tomilin, 1957) are respectively (in cm): condylobasal length 152, 142, and 106 (tip of rostrum broken in the male); zygomatic width 75 (measurement not taken in the second female and male); length of rostrum 96, 92, and 52; width of rostrum at base 47, 43, and 31; length of lower jaw 133, 130, and 90; and length of mandibular symphysis 29, 27, and 17. (V.S.) AS ey ed SSS == SSS => SSSSSS SSSR SSeS SSS ~ 7 ИИА "И 1 AAW NY NYS BAW LY SRS ГИЯ! Ss ЧИ! 0; 632 Fig. 367. Skull of Вана’; beaked whale, Berardius Бата! (figure by М.М. Kondakov). 632 633 854 Geographic Distribution? These animals are distributed in the North Pacific Ocean. Geographic Range in the USSR (Fig. 368) In the Sea of Japan, it is distributed in Peter the Great Gulf and along the southern coasts of Sakhalin; in the Sea of Okhotsk, along the islands of the Kuril range and the coasts of western Kamchatka, close to the eastern coasts of Sakhalin, in the Gulf of Sakhalin in waters of the Shantarsk Islands and [ona Island, in the central part of the Sea of Okhotsk; in the Pacific Ocean, in waters of the Kuril Islands, along the southeastern coast of Kamchatka, in Avachinsk Bay and Kronotskiy Gulf, and around the Commander Islands; and in the Bering Sea, around Karagin Island and in Olyutorskiy Gulf. It apparently does not penetrate farther north than Cape Navarin (about 62°N lat.). Geographic Range outside the USSR (Fig. 369) The southern boundary of distribution probably traverses at the latitude of southern Japan. It is known in the Sea of Japan as well as the Pacific Ocean side of the islands of Japan but is more common and abundant in the Pacific Ocean than in the Sea of Japan. On the eastern side of the North Pacific Ocean, it lives from the coasts of California along the entire North American continent to the coasts of the Alaskan peninsula and Aleutian Islands. In the Bering Sea, it is distributed from Bristol Bay and the Pribilof Islands along the boundary of the eastern Bering Sea shallow-water zone to northwest of Cape Navarin on the Asian coast and the southern part of the Gulf of Anadyr (Zenkovich, 1939; Sleptsov, 1955; Tomilin, 1957; Chapskii, 1963). (V.A.) Geographic Variation Geographic variation has not been established. It is possible that Baird’s and Arnoux’s beaked whales (bairdi and arnouxi) represent only subspe- cific forms of the same species (Hershkowitz, 1966). Further, there is justification to assume that the species forms only some local herds or populations and not subspecies. For example, the Pacific Ocean coasts of Honshu and the Sea of Okhotsk’s coast of Hokkaido are evidently inhab- ited by various populations of Pacific beaked whales (Omura, Fujino, and Kimura, 1955; see below). (V.A.) 9 [See footnotes on page 854][sic; pp. 843 -844—Gen. Ed.] 635 855 Biology Population. Only a few hundred beaked whales are caught every year in the North Pacific Ocean. It may be concluded from this that the population of these whales is comparatively small. However, in most parts of the range the beaked whale is caught only incidentally while hunting for other whales. This no doubt has a bearing on their overall catch. Food. Data on the food of beaked whales are very meagre. Evidently cephalopods and to a lesser extent fish represent the main food. Food Objects of Baird’s Beaked Whale (Betesheva, 1960, 1961) Cephalopods Fish Gonatus magister Alaska pollock, Theragra Gonatus fabricii chalcogramma Gonatus borealis Podonema, Podonema longipes Ommatostrephes sloani- Navaga, Eleginus navaga pacifius gracilis Albatross rat-tail, Coryphaenoides pectoralis Black rat-tail, Hemimacrurus acrolepis = [Coryphaenoides acrolepis| There are references to finds of redfish (ocean perch), skates and their eggs in the stomach of beaked whales (Zenkovich, 1939; Tomilin, 1957). Daily activity and behavior. Pacific beaked whales are usually confined to groups of 5-7 each or sometimes up to 20 animals (Zenkovich, 1939). They dive for 8-20 min (less frequently up to one hour) after which they “rest” on the sea surface for 3-4 min producing 10-15 blows in this period. The blow of the beaked whale is low, 1.0-1.5 m in height (Tomilin, 1957). On surfacing after prolonged submergence, the forehead of the whale is seen first on the surface followed immediately by a small blow. The anterior part of the dorsum and the dorsal fin are seen next. During intermediate submergences, only the upper end of the dorsal fin is exposed since the whale floats right on the water surface almost without flexing the body. Before undertaking a fresh prolonged submergence, at the moment of appearance of the dorsal fin, the body flexes and the head is concealed under water. Next the anterior part of the trunk is gradually flexed more sharply and assumes an almost vertical posture. The dorsal fin and the caudal stem adjoining it then gradually rise above the water and, when they are highest above the surface, the entire body 856 25 140 150 те J | затык у 2 160 170 633 Fig. 368. Range of Вана’; beaked whale, Berardius bairdi, in the USSR (V.A. Arsen’ev). begins to descend slowly and that part of the trunk visible above the water gradually submerges into the sea, the last to disappear being the caudal flukes. However, the caudal flukes, unlike in sperm whales, do not rise out of the water. On surfacing, the beaked whales form a typical chain, sometimes in twos or even threes, one behind the other (Fig. 370). The animals leap from the water at an angle to the sea surface roughly equal to one- third the length of the trunk, then fall on the water with a loud thud (Tomilin, 1957). Sometimes they completely breach the water and dive with the head down like a dolphin without landing on the flanks or on the abdomen, as generally happens in the case of large whales. The herds usually differ in composition. Adult males and females with calves remain singly, far away from the coasts, forming independent herds, while young animals gather in separate groups close to the coasts and transgress even into bays and gulfs (Zenkovich, 1939). Mixed herds are also noticed, with males, females, and young animals living together 857 180 | | | ! в | Е: И, т к : Е ИНН И ТО Wace Kiss eit HL! Hi, ! : AAS @ a \ j “eit Ри! | |] м i р сх Ц ИЯ. Fig. 369. Species range of Ваша’ beaked whale, Berardius bairdi (V.A. Arsen’ev). 634 858 2 Ри... -/ р PN ane a И LZ 4 ZI AE BE eee = 22 AZ А, att. Le, ee ie SOO DES LA GI Е LZ РРР = Digs ЕЕ Ct a ee eZ oe ene za LEAF AEG gee ae’. РОЗИ РР р р АС 635 Fig. 370. Ваша’; beaked whale, Berardius bairdi, at sea (figure by М.М. Kondakov). (Tomilin, 1962). Many cases of beached Baird’s beaked whales have been recorded: near Centerville and Santa Cruz on the Californian coast, in Tokyo Bay in Japan, and on the Pribilof and Commander Islands. Seasonal migrations and transgressions. Beaked whales perform reg- ular seasonal migrations but their periods and courses have not been established. It is known that they appear in spring in the northern part of the range, in the Sea of Okhotsk and the Bering Sea, and leave for the south in autumn for the waters of California and southern Japan 636 where they winter. Since more beaked whales are caught along the Pacific Ocean coast of Honshu than in the Sea of Japan, it has been assumed that most of the herd wintering in southern Japan migrates north along the eastern side of the islands of Japan. Here they are seen in early May with the maximum number sighted in the waters of Japan in July and August, fewer in September, and hunting ceases altogether in November. Apparently a part of this herd (population) spends the summer months in the waters of Japan and another part farther north since, in May, June, and July, they are noticed along the eastern coast of Kamchatka and in August near the Commander Islands, Karagin Island, and in Olyutorskiy Gulf (Zenkovich, 1939). In the Sea of Japan, beaked whales are seen earlier than along the coasts of the more southward Honshu Island. In March they are sighted in Peter the Great Gulf and in April, along the northeastern coasts of Hokkaido in the Sea of Okhotsk. In May the hunting of beaked whales in Hokkaido increases considerably, lessens in June, and ceases almost alto- gether by July and August. In September and October the second arrival of beaked whales occurs off the Hokkaido coasts and-their catch in these months increases again. It is possible that an independent population of 859 beaked whales lives here, migrating in summer into the Sea of Okhotsk and again appearing in autumn during autumn migrations to the winter- ing sites. A preponderance of males has been recorded in the annual catch of beaked whales from the coastal stations in the waters of Japan. In five years of hunting, they totaled 68.3% of the total catch. It would seem that whales of different sex (and probably of age) groups migrate at different periods by different routes (Omura, Fujino, and Kimura, 1955). Reproduction. Mating and parturition occur in winter months in the wintering areas, but have not been established very accurately. The majority mate in February with most births recorded in December. Both these periods are perhaps protracted by a few months. On August 5, 1957, a female 1,076 cm long held an embryo 12 cm long. Apparently this female had mated in summer. On the other hand, on August 21, 1927, a female caught in the same region was found with an embryo 242 cm long. This female had perhaps mated in winter. Gestation extends for about 10 months (Omura, Fujino, and Kimura, 1955; Omura, 1958). Growth and development. Adults and embryos differ in body propor- tions (Omura, Fujino, and Kimura, 1955). Among embryos: (1) the head is relatively large; (2) flippers are shifted closer to the tail; (3) flippers are relatively large; (4) navel caudally situated although position of the anal opening is the same as in adults; (5) dorsal fin is relatively very high although its relative length along the base is the same as in adults; and (6) caudal flukes at the point of their articulation with the caudal stem are relatively broader. The average size of newborn calves is 4.6 m. Calves 580 cm long were still suckling. The animals attain sexual maturity at three years of age or slightly later at an average body length of 10.1- 10.4 m (females) and 9.7- 10.1 m (males) (Table 63). The smallest pregnant female was 9.8 m long and the largest imma- ture female 10.4 m (Tomilin, 1962). Enemies, diseases, mortality, parasites, and competitors. It would appear that Pacific beaked whales have no enemies. Their diseases have not been studied. From among the ectoparasites, whale lice of the genus Platicyamus have been found. Some instances of barnacles (Conchoderma) adhering to the teeth have also been recorded. Overgrowth of diatomaceous algae Cockoneis sp. and Navicula sp. and adhesions caused by suction of the Pacific lamprey (Entosphenus tridentatus) have also been observed. Nine species of endoparasites are known: trematodes one, cestodes two, nematodes four, and acanthocephalans two. The trematode Oschmarinella sobolevi Skrjabin has been detected in the bile ducts of the 860 637 Table 63. Dimensions of Pacific beaked whales caught in the waters of Japan (Omura, 637 Fujino, and Kimura, 1955) Length of whale, m Caught Males Females Total 5.5 — 1 1 5.8 — 1 Л 6.1 — 1 1 7.0 7 3 5 73 1 4 5 7.6 1 1 2 7.9 1 — 1 8.2 6 7 8 8.5 4 4 8 8.8 8 6 14 9.1 25 17 42 9.4 19 11 30 9.7 38 17 55 10.1 76 18 94 10.4 149 38 187 10.7 158 50 208 11.0 91 61 152 11.3 43 33 76 11.6 9 20 29 11.9 —- 4 12:2 — it 1 liver in beaked whales only from Kuril waters. Neither of the cestodes Trigonocotyle sp. and Tetrarhynchidae gen. sp. has been identified at the species level. Of the four species of nematodes found in beaked whales, Anisakis (Anisakis) simplex Rudolphi, parasitizing the gullet, stomach, and intestine, is very widely distributed among marine mammals. It is known among many species of toothed and baleen whales and Steller’s sea lion among Pinnipedia (detected in the North Sea, off the coasts of Kamchatka, Japan, and New Zealand). The nematode Anisakis skrjabini Mosgovoy, found in beaked whales and sperm whales in the waters of the Commander Islands, the Sea of Okhotsk, and in the Antarctica, localizes in the stomach and small intestine. Crassicauda giliakiana Skrjabin and Andreeva has been found in the kidneys and ureter of beaked whales and belugas of the Sea of Okhotsk and Delamurella hyperoodoni Gubanov in the lungs (tracheae) of only the beaked whale of the Sea of Okhotsk. The acanthocephalan Bolbosoma nipponicum Yamaguti detected in the intestine of beaked whales from Kuril waters is also known among three species of baleen whales and two species of pinnipeds of the Kuril Islands and the seas of Japan and Okhotsk. Echinorhynchus gadi, detected in the 638 861 stomach of beaked whales, is а known parasite of fish (Delamure, 1955; A. Skrjabin, 1959, 1960). The extent of mortality among beaked whales is not known. These whales face some competition for food from sperm whales. Field characteristics. The body length is around 10 m (for adults) and the color monochromatic—dark brown on top and lighter below. The “beak” is flattened. The anterior mandibular section has two pairs of teeth. The much larger first pair projects outward with the mouth closed. The dorsal fin is high and well-proportioned and situated above above the anal opening. These animals live in small groups, dive simultaneously, and never display the caudal flukes. The blow is small, at a height of 1.0-1.5 m, resembles a flash, and is bushy. (V.A.) Economic Importance Minimal but regular hunting of Baird’s beaked whale is carried out only in the waters of Japan. At other places this whale has no economic importance (Table 64). The Japanese and Soviet whaling fleets catch some ten beaked whales every year while hunting for large whales; the coastal stations of British Columbia and California on the North American coast also catch a few of these whales. As these whales are of small dimensions, the quantum of products obtained from them is also comparatively small (Table 65). The quantity of fat obtained from one average size whale is, on average, 2,700 kg (Khar’kov, 1940). Japanese whalers catch beaked whales using small boats equipped with small-bore harpoon guns in the same manner as in hunting large Table 64. Catch of beaked whales in the waters of Japan (international whaling statistics) Year Whales caught Year Whales caught 1948 73 1959 186 1949 92 1960 147 1950 186 1961 133 1951 252 1962 145 1952 321 1963 160 1953 262 1964 189 1954 — 1965 172 1955 258 1966 171 1956 297 1967 107 1957 186 1968 117 1958 229 638 862 whales using large whale boats. The animals caught are brought to the coastal stations. The fat of these whales is used only for commercial purposes. The fat from the jaws (from the mandibular hollow space) and around the jaws serves as raw material for producing a high-quality oil useful for lubricating fine mechanisms. The flesh is generally not used as food but, after cooking, can be used for feeding fur-bearing animals. In some cases, however, the flesh of these whales is processed for human consumption also. Based on their population, the hunting of beaked whales could be somewhat expanded. Rules for controlling hunting have not been formu- lated and, at the present rate of their exploitation, there is yet no need for such. (V.A.) Genus of Beaked Whales [Sword-tooth Dolphins] Genus Mesoplodon Gervais, 1850 1850. Mesoplodon. Gervais. Ann. Sc. Nat., Zoologie, 14, p. 16. Delphinus sowerbiensis Blainville = Physeter bidens Sowerby. (V.H.) Medium body dimensions, with a length up to 6.7 m. The dorsal fin is low and its apex flexed posteriorly. The flippers are pointed at the tips. The bulge of the crescent-shaped blowhole faces backward. The sloping forehead transits gradually into a fairly long beak. The body color is dark (from black to gray) and the abdomen light- colored. The long and thin rostrum is narrower on top than in other mem- bers of the family of beaked whales and exhibits mesorostral ossification, Table 65. Weight of body parts of Baird’s beaked whales (Zenkovich, 1937b; Tomilin, 1951; Sleptsov, 1961) Body part Body length and sex Female Female ? 10.0 m 10.8 т 11.1 м Subcutaneous fat 3,337 2,258 5,616 Flesh 2,400 2,838 1,9741 Skeleton 1,953 1,256 3,050! Viscera 958 1,148 399 Total 8,648 7,500 11,039 Note: The heart, lungs, and liver of another female, 11.1 m long (pregnant), weighed 54, 281, and 64.5 kg respectively (Khar’kov, 1940). ['These figures are possibly reversed.—Ed.] 863 formed through ossification and fusion with the surrounding bones of the vomer. The supraoccipital and interparietal bones form a high crest. The frontal bones have small orbital processes. The mandibular symphysis is considerable in size (one-fifth to one-third the jaw length). One pair of teeth, strongly compressed laterally, is located at the level of the anterior third of the lower jaw (at the end of the jaw only in М. muirus) and projects outside the mouth when it is closed, while cov- ering the upper jaw from the sides. The roots of the teeth run obliquely to the longitudinal axis of the jaw. Among some species of these beaked whales, the teeth are sharply enlarged; they are larger among males than females. Numerous minute teeth are concealed in the gums of the upper and lower jaws, particularly among young animals (they may be resorbed with advancing age). Cervical vertebrae 7, thoracic 9-11, lumbar 9-12, and caudal 18-21; total 46-48. The atlas and axis are fused; sometimes a third cervical vertebra is attached to them. The spinous vertebral processes are very large. The sternum has four (less often five) sections. Of the five digits on the forelimb, the second and third are the longest. Biology very poorly studied. These are pelagic whales, rarely encoun- tered along coasts. They remain singly, in small groups, or much larger herds. They apparently feed mainly on cephalopods. The periods of mat- ing and parturition are very prlonged. The distribution is extremely wide: in the eastern part of the Atlantic Ocean, from Norway and the British Isles to Madeira, the Mediterranean Sea, and coasts of South Africa; in the western Atlantic, from Newfound- land and Canada; and in the Caribbean Sea, from Trinidad and the coast of Argentina (42° S lat.) to the Falkland Islands. In the Pacific Ocean, in the east from the Bering Sea to La Jolla in California in the north and along the coasts of Chile in the south, and in the west from the Bering Sea to Japan and from Australia to New Zealand; and the Indian Ocean (Hershkowitz, 1966). Abundant everywhere (Fig. 371). Fossil remains have been found in the Upper Miocene of North America and in the Upper Miocene and Middle Pliocene of Europe. The genus comprises 10 (Nishiwaki and Kamiya, 1958; Hershkowitz, 1966) or 11 (Moore, 1968) species: M. bidens Sowerby, 1804—North Atlantic from Norway, the Baltic Sea, and Great Britain in the east to the Mediterranean Sea (inclusive) and in the west from Newfoundland to Massachusetts state (USA); M. europaeus Gervais, 1848 - 1852—North Atlantic (English Channel and from New York to Florida, Gulf of Mex- ico, and also the Caribbean Sea from Cuba to Trinidad); M. mirus True, 1913—North Atlantic (British Isles and coasts of France and from Cape Breton and Nova Scotia peninsula to Florida), South Atlantic (southern Ц И in ИНЫХ Uystglystetadalateletate | Е a $ 3 8 8 8 Fig. 371. Range of the genus of beaked whales [sword-tooth dolphins], Mesoplodon (V.A. Arsen’ev). 640 641 865 coasts of South Africa); М. grayi Haast, 1874—Indian Ocean (coasts of South Africa and Australia), South Pacific Ocean (from New Zealand to Chilean coasts), South Atlantic from the coasts of Argentina and the Falkland Islands to South Africa, North Atlantic (Netherlands coast); M. ginkgodens Nishiwaki and Kamiya, 1958—North Pacific Ocean from Japan in the west to California in the east and coasts of Sri Lanka in the Indian Ocean; М. layardi Gray, 1865—South Pacific Ocean and Indian Ocean from Australia and New Zealand to the South Atlantic between the Falkland Islands and South Africa; М. densirostris Blainville, 1817—eastern part of the Atlantic Ocean (coastal waters of Madeira and South Africa), western Atlantic (Nova Scotia peninsula, Canada, and ' Bahama Islands), western part of the Pacific Ocean (Lord Howe Island and Queensland in Australia), North Pacific Ocean (Midway Island), and Indian Ocean (from South Africa and the Seychelles to apparently west- ern Australia); М. stejnegeri True, 1885—Pacific Ocean from the Bering Sea to Oregon state (USA) in the east and Japan in the west; M. bow- doini Andrews, 1908—New Zealand coast; M. carlhubbsi Moore, 1946 [1963]—North Pacific Ocean (east coast of Japan and west coast of the USA between 32° and 47° N lat.); and M. hectori Gray, 1871—Southern hemisphere, temperate waters. The systematics of the genus of beaked whales has not been suffi- ciently developed. Many species have been described on the basis of just one or two specimens and the differences between some of them repre- sent only individual variations. Thus, it is possible that M. stejnegeri, M. bowdoini, M. ginkgodens, and M. carlhubbsi, inhabiting the Pacific Ocean, belong to the same species, as conceded by some scientists who regard the differences between these species as inconsequential. The main dif- ferences are: the lateral thickness of the tooth crown is greater than its front-to-back width—in M. stejnegeri by seven times, M. ginkgodens six times, and М. bowdoini three or four times; and the ргетахШагу foramena lie at the same level as the maxillary foramena or posterior to it in M. stejnegeri and M. bowdoini, but markedly more anteriorly in M. ginkgodens. There are other differences also. The collection of more factual data should clarify the systematics of the genus. On the other hand, the differences between some species (groups of species) are quite significant. Thus, M. layardi Gray is isolated in a special subgenus, Dolichodon Gray, M. densirostris Blainville in subgenus Dioplodon Gervais, and all the rest of the species placed in the subgenus Mesoplodon Gervais (Moore, 1968). Only one species, Stejneger’s beaked whale, M. stejnegeri, has been reported in USSR waters. Some other species may also be encoun- tered: in the southern part of the Baltic Sea—Sowerby’s beaked whale, 642 866 М. bidens; in the seas of the Far East—M. carlhubbsi and М. ginkgodens. The latter two species could, however, be regarded as belonging to the species M. stejnegeri and hence are not described here. There is no special hunting of these beaked whales. (V.S.) STEJNEGER’S BEAKED WHALE Mesoplodon (Mesoplodon) stejnegeri True, 1885 1885. Mesoplodon stejnegeri. True. Proc. U.S. Nat. Mus., 8, p. 585. Bering Island and Commander Islands. (V.H.) Diagnosis Total body length up to 6 m. Color black, more light-colored ventrally. The antorbital notches in the skull are developed relatively poorly. The foramena of pair V of the nerves in the maxilla lies anterior to that in the premaxilla or at the same level. The palatine bones are not adjacent. The ratio of length of tooth crown to its width is more than in the other species of these beaked whales (6.4:1 or even slightly more). (V.S.) Description The lower jaw is longer than the upper by 15 mm. The flippers are relatively small. The caudal flukes are lighter in color than the trunk and may be white on the underside. Quite a large number of small white patches (traces of scars or parasitic infections) are present on the body surface (Fig. 372). The teeth are strongly built in the males (Fig. 373). but barely emerge from the gums in the females. The following are the main body measurements of an adult male Stejneger’s beaked whale (Tomilin, 1957) (in cm): body length 509; max- imum girth of body 254; distance from tip of lower jaw to base of flip- pers 112; distance from tip of upper jaw to anterior margin of blowhole 59; distance from anal opening to notch between the caudal flukes 141; Fig. 372. Stejneger’s beaked whale, Mesoplodon stejnegeri (figure by N.N. Kondakov). 867 (9) Fig. 373. Head of Stejneger’s beaked whale, Mesoplodon stejnegeri, lateral view (figure by N.N. Kondakov). length of flippers along the anterior (lower) margin 55; maximum width of flippers 16; height of dorsal fin 21; length of dorsal fin along the base 34; and spread of caudal flukes 128. The main measurements of the skull (Fig. 374) of male Stejneger’s beaked whales caught off the coasts of Oregon and California (two speci- mens), and Japan (one specimen) (Tomilin, 1957; Nishiwaki and Kamiya, 1959) are respectively (in cm): condylobasal length 81, 82, and 80; zygo- matic width 39, 38, and 38; length of rostrum 49, 51, and 49; width of rostrum at base 23, 22, and 16; length of lower jaw 69, 71, and 68; and length of mandibular symphysis 18,—, and 18. (V.S.) Fig. 374. Skull of Stejneger’s beaked whale, Mesoplodon stejnegeri (figure by N.N. Kondakov). 643 868 Geographic Distribution and Biology Pacific Ocean from the Bering Sea to the Californian coast in the east — and coasts of Japan in the west (information on distribution is based exclusively on the sites of animals stranded on coasts; field observations are almost nil). Geographic Range in the USSR (Fig. 375) Known only from the waters of the Commander Islands where three instances of beached beaked whales have been recorded on the shores of the Bering Sea. Their presence is possible in the waters of Kamchatka and the Kuril Islands. Fig. 375. Range of Stejneger’s beaked whale, Mesoplodon stejneger1, in the USSR (V.A. Arsen’ev). 643 645 869 Geographic Range outside the USSR (Fig. 376) Aleutian Islands (one record), Alaskan peninsula (one record), Pacific Ocean coast of North America (five records off the coasts of Washington, Oregon, and California states), and Japanese coasts (two records in the region of Ayukawa) (Tomilin, 1962; Hershkowitz, 1966). Geographic variation has not been established. Biology virtually not studied since the species is very rare through- out its range. Cephalopods apparently serve as the main food although some individual whales have been sighted at the sites of congregations of salmon. The very small number of observations indicate that they live singly or in very small groups (up to three animals). In life style, they are probably similar, if not identical, to Sowerby’s beaked whale (M. bidens Sowerby, 1804). Adult animals are about 6 m long and black with a light-colored underside. Teeth one pair, strongly compressed laterally, project from the sides, and cover the upper jaw when the mouth is closed. These whales are of almost no economic importance. (V.A.) SOWERBY’S BEAKED WHALE Mesoplodon (Mesoplodon) bidens Sowerby, 1804 1804. Physeter bidens. Sowerby. Trans. Linn. Soc. London, 7, p. 310. Elginshire, Scotland. 1817. Delphinus sowerbensis. Blainville. Nouv. Dict. Hist. Nat. 9, p. 177. Substituted for Physeter bidens Sowerby. (V.H.) Diagnosis Maximum overall body length up to 5.5 m. Body dorsally black or bluish-black and usually grayish or whitish ventrally. The antorbital notches are well developed in the skull. The foramena of pair V of the nerves in the maxilla is located posterior to the foramena in the premaxilla. The palatine bones are adjacent. The rostrum is broad at the base. The teeth lie posterior to the mandibu- lar symphysis. The ratio of length of tooth crown to its width is 3.5:1. (V.S.) Description The head is slightly compressed laterally. The “beak” is slightly flattened dorsoventrally (Figs. 377 and 378). The lower jaw is only п’еу). ge of Stejneger’s beaked whale, Mesoplodon stejnegeri (V.A. Arse Fig. 376. Species ran 644 645 645 871 Fig. 377. Sowerby’s beaked whale, Mesoplodon bidens (figure by М.М. Kondakov). V2) Fig. 378. Head of Sowerby’s beaked whale, Mesoplodon bidens (figure by N.N. Kondakov). insignificantly longer than the upper (roughly by 13 mm). The body color is variable—from black to gray with abdomen light, sometimes white. The anterior margin of the caudal flukes and the upper and lower jaws may be light-colored. Numerous whitish spots and bands—remnants of scars—cover the body. The teeth of males are much larger than in females; in the latter, the teeth in most cases do not emerge from the gums. The basic body measurements of two female Sowerby’s beaked whales are as follows (Tomilin, 1957) (in cm): body length 345 and 431; distance from tip of snout (in second animal from tip of lower jaw) to flippers 91 and 109.8, up to blowhole 44 and —; length of flippers 30 and 39.2, their width 12 and 13; distance from tip of snout to dorsal fin 204 and 275.8; height of dorsal fin 27 and 20.9, its length 49 and 36.6; and spread of caudal flukes 68 and 99.4. The basic skull measurements (Fig. 379) of three males caught off the Shetland Islands, Norway, and Sweden (Tomilin, 1957) are respectively (in cm): condylobasal length 74, 73, and 74; zygomatic width 29, 30, and 27; length of rostrum —, 48, and 50; width of rostrum at base 18, 19, and 17; length of lower jaw 70, 64, and 64; and length of mandibular symphysis —, 21, and 22. (V.S.) 872 Geographic Distribution North Atlantic Ocean from the coasts of Norway, the British Isles, and the Baltic Sea up to the Mediterranean Sea in the east and from New- foundland to Massachusetts state along the American coast (the distri- bution of the species was established exclusively from information on beached animals). Geographic Range in the USSR Stray transgressions are possible into our waters in the Baltic Sea. Geographic Range outside the USSR (Fig. 380) Finds of beached animals are known in Newfoundland and Massachusetts state on the North American coast. In the eastern part of the North Atlantic, they have been registered on the coasts of Norway, Sweden, England, Poland, Holland, the FRG, the GDR, Belgium, and France. This whale is encountered in the Mediterranean Sea (Sergeant and Fisher, 1957). (V.A.) ; 645 Fig. 379. Skullof Sowerby’s beaked whale, Mesoplodon bidens (figure by N.N. Kondakov). 646 646 Fig. 380. Species range of Sowerby’s beaked whale, Mesoplodon bidens (V.A. Arsen’ev). Geographic Variation Not established. Biology Very poorly studied. Absolutely nothing is known about the population of Sowerby’s beaked whales. Information on their food is not available. Behavior. These whales are seen more Often singly or in pairs. In the shallow-water bays of Khertvik on Gassen Island (Norway), the “tech- nique” of two of these whales stranded on the coast was observed on April 18, 1957. Initially, they swam along the coast to a shallow point almost touching the bottom. For a quarter of an hour they attempted to return to the sea but a low tide occurred and the exit from the bay became very Shallow. Then, the larger of the two whales turned directly toward the coast and soon found itself on a sandy beach. It quivered its tail vigor- ously and within minutes a large pit had formed in the sand. Meanwhile the smaller whale swam along the coast but having covered a distance of only 20 m slammed against some rocks and remained motionless; it resisted only slightly when its tail was looped. From the first sighting of these animals along the coast to their death, no more than 30 min elapsed. One was an adult female 505 cm in body length and the other a large female calf 315 cm long (Jonsgard and Hoidal, 1957). Seasonal migrations and transgressions. In La Mancha Strait and on the coasts of Great Britain, beached Sowerby’s beaked whales have been found in March, April, September, and December; in the Baltic Sea, 647 874 such finds have been recorded in February, June, August, and September. These animals are believed to inhabit these waters throughout the year. Their migrations have not been traced. Reproduction, growth, and development. Mating and parturition occur at the end of winter and spring. The period of mating is quite prolonged. Gestation extends for about a year. The body length of a calf at birth varies from 152 to 213-244 ( 183) cm. In the period of lactation, the calf adds 90-120 cm to its length. Suckling ceases at a body length of about 300 cm (Jonsgard and Hoidal, 1957). Enemies, diseases, mortality, parasites, and competitors. Enemies of Sowerby’s beaked whale are not known. The diseases have not been studied. An endoparasite (Conchoderma sp.) was found on the teeth of Sowerby’s beaked whale. Six species of helminths are known. Cysts of the trematode Monostomum delphini Diesing were found in the blubber of a whale from Gavra region but the highly superficial description does not permit positive identification of the species affinity of this parasite. The cestode Strobilocephalus triangularis Diesing, detected in the intestine of Sowerby’s beaked whale, is also known in the bottlenose whale and two or three species of dolphins. It has been found off the coasts of Portugel and Brazil. The cestode Tetrabothrium forsteri Krefft, parasitizing the intestine of Sowerby’s beaked whale, was also detected in two species of dolphins from the Mediterranean Sea and New Zealand waters. The widely distributed cestode Phyllobothrium delphini Bosk, parasitizing the skin of Sowerby’s beaked whale, is also known in seven species of toothed whales, the Arctic whale, and Weddell’s seal. It has been detected near the Azores, in the Mediterranean Sea, off the Commander Islands, along the coasts of Australia, and in the Antarctic. The nematode Anisakis (Anisakis) simplex Rudolphi is very widely distributed and, in addition to Sowerby’s beaked whale, parasitizes the gullet, stomach, and intestine of ten other species of toothed whales, two species of baleen whales, and Steller’s sea lion. It has been detected in the North Sea and off the coasts of Kamchatka, Japan, and New Zealand. The only species of acanthocephalans, Bolbosoma vasculosum Rudolphi, was found in the intestine of Sowerby’s beaked whale and the common dolphin from the Atlantic Ocean and the Mediterranean Sea (Delamure, 1955). Field characteristics. The maximum body length is 5.5 m. The body is almost black dorsally and slightly lighter ventrally. The lower jaw is almost not longer than the upper. The teeth almost do not emerge among females. With the mouth closed, a pair of large flat teeth projects upward along the sides of the upper jaw in males. 875 Sowerby’s beaked whale has по economic importance and there is no hunting of this animal. (V.A.) Genus of Goose-beak [Cuvier’s Beaked] Whales Genus Ziphius G. Cuvier, 1823 1823. Ziphius. G. Cuvier. Rech. ossem. foss., 5, р. 350. Z. cavirosiris G. Cuvier. 1846. Xiphius. Agassiz. Nomencl. Zool., p. 389. Correction for Ziphius G. Cuvier. (V.H.) Medium size, with body length up to 8 m. The “beak” is short and transitions smoothly into the low frontal pro- jection of the corpus adiposum. The jaw line is short. The dorsal fin is low and highly variable in shape in different animals, from crescent-shaped to triangular. The flippers are relatively long and narrow. The lower jaw projects forward more than the upper. The jaw line extends posteriorly to less than half the distance between the tip of the snout and the eye. The color is usually dark and subject to considerable individual vari- ation. The dorsum may be lighter in color. The skull is asymmetric. Among the members of the family, the ros- trum of the skull is the widest and shortest (only slightly longer than the cranium) in this genus. The vomer forms a stable mesorostral ossifi- cation. Longitudinal maxillary crests are very poorly developed. A large depression, housing the spermaceti sac, occurs on the dorsal side of the skull between the outer edges of the maxillae. The mesethmoid bone is ossified. Posterior to the nostrils, the maxillary, premaxillary, and nasal bones form a large projection in the form of an arch, which overhangs the nostrils. The nasal bones attain maximum width anteriorly. Teeth one pair, located on the anterior tip of the lower jaw, and projecting anterior to the upper jaw when the mouth is closed. The sternum consists of five sections. Cervical vertebrae 7, thoracic 9-10, lumbar 10-11, and caudal 19-22; total 46-49. The anterior three _ or four cervical vertebrae are usually fused. Phalangeal formula: [,, Из, III;_¢, [У4, and У, ->. Ribs 9 pairs. Biology not well studied. These animals represent forms of the open sea and feed apparently on mollusks. The periods of mating and partu- rition are prolonged. These whales are distributed in the World Ocean but are few in number everywhere. The genus consists of a single species: Cuvier’s beaked whale, Ziphius cavirostris G. Cuvier, 1823. There is no special hunting of these whales. (У.5.) 648 876 CUVIER’S BEAKED WHALE Ziphius cavirostris G. Cuvier, 1823 1823. Ziphius cavirostris. G. Cuvier. Rech. Oss. Foss. ed. 2, 5 (1), p. 350, pl. 27. French coast in the department of Rhone delta (Bouches- du-Rhone), Mediterranean Sea. 1826. Delphinus desmaresti. Risso. Hist. Nat. Eur. Mérid., 3, p. 23. Mediterranean Sea. 1883. Ziphius grebnitzkii. Stejneger. Proc. Ц. $. Nat. Mus., 6, р. 77. Com- mander Islands. (V.H.) Diagnosis Only species of the genus. Description The frontal projection is much lower than in bottlenose whales and Pacific beaked whales (Figs. 381 and 382). The characteristic small oral cavity is smaller than in other species of the family of beaked whales. Two longitudinal grooves traverse the throat. The bulge of the crescent- shaped blowhole faces backward. 648 Fig. 381. Cuvier’s beaked whale, Ziphius cavirostris (figure by N.N. Kondakov). 648 649 877 (9) Fig. 382. Head of Cuvier’s beaked whale, Ziphius cavirostris (figure by М.М. Kondakov). The body color is subject to great individual variation. Monochro- matic black, gray, or bluish-gray animals are encountered. In others, the ventral side may be lighter in color than the dorsal but the opposite, albeit less often, has also been recorded. Other color variations are likewise possible. Many depigmented patches of various shapes and sizes are seen on the skin. The head and dorsum (up to the dorsal fin) are sometimes very light-colored in old animals. The rostrum of the sharply asymmetric skull is poorly set off from the cranium. Adult males are characterized by a mesorostral ossification that is more prominent than in females and the young. This ossification lies in a very deep longitudinal depression of the rostrum between the premaxillae. The teeth of males are larger and more massive than in females (tooth diameter in males 25-29 mm, in females 10-14 mm). Young animals have 28-30 vestigial teeth in the gums of the upper and lower jaws. These teeth reduce in number as the animal ages. Cuvier’s beaked whale has 7 cervical, 9 thoracic, 11 lumbar, and 20 caudal vertebrae. The proportions of these sections to the length of the entire vertebral column are respectively 3, 17, 36, and 44% (Slijper, 1936a). The stomach of Cuvier’s beaked whale has five sections; the first and last are large and the three intermediate ones small; all of them are smooth-walled (Kenyon, 1961). The main body measurements of three females and one male Cuvier’s beaked whale (Tomilin, 1957; Kenyon, 1961) are respectively (in cm): body length 579, 584, 658, and 543; distance from tip of snout to posterior edge of dorsal fin 389, 391, 425, and 350, to the axillae 115, 142, 132, and 131, to the blowhole 60, —, 66, —; length of flippers 46, —, 56, and 42; maximum width of flippers 16, 17, 17, and —; height of dorsal fin 30, 19, 25, and 21; and width of caudal flukes from tip to tip 162, 182, 175, and 133. The largest of the animals caught had a body length of 793 cm. 652 649 878 Females are larger than males. Of 51 males and 34 females caught in Japan, the average length of the males was 550 cm (maximum 671 cm), of the females 580 (701) cm (Table 66). The following are the basic skull measurements (Fig. 383) of two adult males and a female Cuvier’s beaked whale (Tomilin, 1957) (in cm): condylobasal length 85, 81, and 88; zygomatic width 54, 52, and 53; length of rostrum 48, 46, and 47; width of rostrum at base 33, 32, and 48; length of left half of lower jaw 75, 74, and —; and length of mandibular symphysis 18, 16, and 18. (V.S.) Geographic Distribution Warm, temperate, and partly temperate-cold waters of the World Ocean. Geographic Range in the USSR Constitutes a very small part of the entire range of the species. Cuvier’s beaked whale has not been sighted with certitude in our waters of the basin of the Atlantic Ocean but may be encountered on out coasts of the Baltic Sea since it is known from Kattegat and waters of southern Sweden (transgressions): in the Far East, waters of southeastern Kamchatka and the Commander Islands where beached animals are a regular annual fea- ture. According to less reliable information, the animal has been sighted near the Kuril Islands and off the southern coast of Sakhalin (Sleptsov, 1961). Table 66. Dimensions of Cuvier’s beaked whales caught in Japan (Omura, Fujino, and Kimura, 1955) Body length, cm Number of whales Males Females Total 305 1 — 1 335 a — = 366 — 2 2 396 3 2 5 427 2 1 3 457 4 — 4 488 2 4 6 518 4 7 6 549 9 5 14 579 8 6 14 610 7 5 12 640 7 5 12 671 4 1 5 — 1 879 y =. а / ON РК) А РК м yr 649 Fig. 383. Skull of Cuvier’s beaked whale, Ziphius cavirostris (figure by М.М. Kondakov). Geographic Range outside the USSR (Fig. 384) Encountered in the North Atlantic Ocean along the coasts of the British Isles and southern Sweden, close to the coasts of Spain, Italy, France, and Corsica (Ligurian Sea), northern Africa, in the Mediterranean Sea, and in the Atlantic waters of North America. It is known in the Pacific Ocean around the Hawaiian Islands, in the East China Sea, along the coasts of Japan, and the Pacific Ocean coast of North America from Cali- fornia to the Pribilof Islands in the north. In the Southern hemisphere, it has been observed in the waters of Brazil, Argentina, South Africa, close to Sri Lanka, Polynesian Islands, around Australia, Tasmania, and New Zealand. Beached animals have been recorded at the following points: on the British Isles and Shetland Islands, in Kattegat Strait and on the coasts of Boguslan (Sweden), in the Baltic Sea, Bay of Biscay, Mediter- ranean Sea (near Nice, in the Rhone delta, at Malaga near Valencia, on Corsica, on the coast of Italy at Liguria, Messina Bay, and on the coast of Algeria), on the coast of North America in the states of New Jersey, Massachusetts, South Carolina, Rhode Island, California, and Alaska, and in British Columbia (Fig. 385). A large number of beached animals is known even in the Southern hemisphere (Tomilin, 1957; Clarke, 1958; Chapskii, 1963). Comparatively uncommon throughout its range. (V.A.) *(лэ.цэзгу “W'A) 5мколар smiydiz ‘этеца рэхеэа з.лэтлпо до э8ие1 $э1024$ ‘pee “314 059 i "7 651 р. 385. Above: Cuvier’s beaked whale near Cape Chernyi, Mednyi Island, July, 1962. Below: Cuvier’s beaked whale stranded on the shore of Bering Island, : May, 1960 (photographs by S.V. Marakov). 653 882 Geographic Variation Not studied. Biology Population. Comparatively common in our waters along the eastern coast of Kamchatka, especially around the Commander Islands; very rare or sighted occasionally elsewhere. Food. Data on this aspect are negligible. It can only be assumed that cephalopods serve as the main food of Cuvier’s beaked whale. Of the three stomachs dissected on the Commander Islands, one was empty and two contained the beaks of squids and crystalline lenses of cephalopods (Tomilin, 1957). The stomach of a female found on Amchitka Island (Aleutians) held the remains of 1,300 cephalopod mollusks—squids Gonatus sp.—in the form of beaks, lenses, etc. (Kenyon, 1961). Behavior. These whales live in small groups, remain on the sea sur- face for about 10 min, dive together, and spend 10 to 30 min or more under water. Seasonal migrations and transgressions. Along the Commander Islands (in the northern region of distribution), these animals are sighted in March and continue to be seen until autumn. Along the coasts of Japan, these whales are caught year-round although the best hunting season is from May through October, peaking in August. Hunting is carried out from coastal stations located exclusively on the Pacific Ocean coast of Honshu and on the northeastern coast of Hokkaido. In the Sea of Okhotsk and along the coast of the Sea of Japan, where Baird’s beaked whales are hunted, Cuvier’s beaked whales are not caught (Omura, Fujino, and Kimura, 1955). Apparently Cuvier’s beaked whales are also encountered along the coast of California almost year-round, as supported by the following records of whales stranded at roughly 41° N lat.: a large female on February 17, 1963; an adult female on September 22, 1959; an adult male on March 2, 1957; and an immature small male on November 24, 1957 (Mitchell and Houck, 1967). Reproduction, growth, and development. The periods of mating and parturition have not been established but in all probability both are protracted. In 1951, 1952, and 1953, embryos 30, 97, and 213 cm long were detected in August; embryos 170 cm long in September; and others 43 cm long in October, 1952 (Omura, Fujino, and Kimura, 1955). An embryo 267 cm long, found in a dead female on Bering Island (Comman- der Islands), was fully developed (Tomilin, 1957). Probably the size of newborn Cuvier’s beaked whales is proximate to the above range. Males attain sexual maturity at a body length of 5.5 to 5.9 m; their testes weigh 883 3.5 to 4.2 kg; females become mature at a body length of about 5.5 т. А female 658 cm long, cast on Amchitka Island (Aleutians), contained at least two traces of corpora lutea of pregnancy. This female had 24-28 layers in her tooth dentine and thus her age was estimated at 12-14 years. A male with a body length of 544 cm, found on the same island, had 13 layers in the tooth dentine or was six-seven years of age (Kenyon, 1961). Enemies, diseases, parasites, mortality, and competitors. Two species of nematodes and one of cestodes are known in Cuvier’s beaked whale. The nematode Crassicauda crassicauda Creplin parasitizes, in addition to Cuvier’s beaked whale, the urogenital system of another species of toothed whales and six species of baleen whales; Crassicauda boopis Baylis, parasitzing the urogenital system of Cuvier’s beaked whale, the humpback whale, and the fin whale, was found in the Atlantic Ocean in the Northern and Southern hemispheres. A cestode, Phyllobothrium sp.., was detected in the blubber of Cuvier’s beaked whale (Delamure, 1955; Tomilin, 1962). Field characteristics. The body length of adult whales averages 5-7 m. The “beak” is broad and short and the forehead low and sloped. The blowhole is crescent-shaped and its horns turned anteriorly. Teeth one pair, located in the anteriormost part of the lower jaw, and not con- cealed by the upper jaw, projecting anterior to its edge when the mouth is closed. These animals live in small groups. While diving, the group disappears under water in unison. (V.A.). Economic Importance Only a few tens of Cuvier’s beaked whales are caught annually and as such this species has no economic importance whatsoever. It is caught along with other species of small whales from the coastal stations along the Pacific Ocean coast of Japan. From 1948 through 1955, 3, 10, 27, 35, 36, and 40 animals were caught annually (Omura, Fujino, and Kimura, 1955). Sexually mature males ranging from 5.5 to 6.5 m in length constituted the major part of the catch. Weight of the Body Parts of Male Cuvier’s Beaked Whale, Body Length 658 cm (kg) (Kenyon, 1961) Flesh, blubber, bones (including head) О) Heart 15.4 Lungs 39.4 Liver 25.9 Kidneys 16.4 884 Reproductive organs 20.7 Stomach: Empty 173) Its contents 18.6 Intestine with contents 41.3 Spleen 0.1 Aorta, bronchi, and connective tissue 31.6 Blood (some inevitably lost) 8.1 Total 2.9837 Hunting is carried out mainly from small boats and using small-bore 654 harpoon guns. A good number of animals fall into nets. The products are used like those of other small whales. On the Commander Islands the flesh and blubber of beached Cuvier’s beaked whales are fed to dogs and foxes. There is no hunting of this whale in our range. (V.A.) Genus of Bottlenose Whales Genus Hyperoodon Lacépéde, 1804 1804. Hyperoodon. Lacépéde. Hist. Nat. des Cétacées, рр. XLIV, 319. Hyperoodon butskops Lacépéde = Balaena ampullata Forster. 1804. Anarnak. Lacépéde. Ibid., pp. XXVIII, 164. Anarnak groenlandicus Lacépéde = Balaena ampullata Forster. (V.H.) Medium-size whales. The “beak” is prominently displayed, more sharply than among other species of the family. The frontal projection of the prominent corpus adiposum [forehead] descends rather steeply from the base of the beak. The corpus adiposum is better developed in males than in females. Two to four longitudinal grooves occur on the throat. The dorsal fin is flexed along the posterior edge. The body is blackish-gray but light gray or yellowish-gray on the abdomen. In the skull the maxillae dorsally form at the base of the rostrum two (right and left) lateral crests which are particularly well developed in adult males. The rostrum is long. The nasal bones are large and concave at the top and the mesethmoid bone is incompletely ossified. Together with the premaxillae, the former bones rise posterior to the bony nares vertically upward and even overhang them. The maxillae have high crests that extend along the edges of the rostrum. One pair, less often two pairs of teeth develop at the tip of the lower jaw. The teeth are larger in males 655 885 than in females. Cervical vertebrae 7, thoracic 9 (8), lumbar 9-11, and caudal 18-20; total 43 - 46. The cervical vertebrae are fused. The sternum consists of three sections. Phalangeal formula: I, _5, Пб-в, Ш, [V4-5, and V,_3. These animals feed mainly on cephalopods. The periods of mating and parturition are prolonged. The duration of gestation is about a year. They are distributed in the North and South Atlantic Ocean, and in the Indian and Pacific oceans. Seasonal migrations take place. The genus comprises two species: northern bottlenose whale, Hyperoodon ampullatus Forster, 1770, and southern flat-faced bottlenose whale, Hyperoodon planifrons Flower, 1882. Mesoplodon planifrons [Mesoplodon pacificus|, described by Long- man in 1926, is sometimes treated as a synonym of Hyperoodon plani- frons (Hershkovitz, 1966), or as a subspecies of Mesoplodon mirus Moore, 1960 (Moore, 1960), or as a lone species of a different genus, Indopace- tus Moore, 1968 (Moore, 1968). Additional data are required before an accurate conclusion can be drawn (descriptions are based on just two skulls). Only the northern bottlenose whale is encountered in the seas of the USSR. There is no special hunting of this species in our waters. (V.S.) NORTHERN BOTTLENOSE WHALE Hyperoodon ampullatus Forster, 1770 1770. Balaena ampullata. Forster. Kalm’s Travel into N. America, I, p. 18. Maldon, Essex, England. 1776. Balaena rostrata. Muller. Zool. Danicae. Prodr., p. 7. Waters of Denmark and Norway. 1789. Delphinus butskopf. Bonnaterre. Tabl. Encycl. Méth. des Trois Reg- nes de la Nature. Cectologie, р. 25. Honfleur, France. 1822. Delphinus hyperoodon. Desmarest. Encycl. Méth. Mamm., 2, p. 520. Thames, England. (V.H.) Diagnosis Only species of the genus found in waters of the USSR. Description Dorsal fin with a concave posterior margin and located above the anal opening (Fig. 386). The flippers are larger in males than in females. The bulge of the crescent-shaped blowhole faces the tail. Two (sometimes 655 656 886 <. Fig. 386. Northern bottlenose whale, Hyperoodon ampullatus (figure by М.М. Kondakov). four) longitudinal grooves extend along the throat. Four hairs are seen on each side of the snout in the embryo. The body is blackish-gray dorsally and light gray ventrally. The fins are grayish-black. With advancing age, the color turns light, almost yel- low. The skin is covered with numerous white spots, apparently caused by a fungal disease. Two pairs of teeth are seen only in some adult animals. In females and many males the posterior pair of teeth does not emerge or, if it does, these teeth are notably smaller than the anterior pair, which may reach a height of 6.5 cm. The anterior teeth are set at the tip of the lower jaw and project Outside when the mouth is closed. The second pair of teeth (when present) is separated from the first pair by a considerable gap. Males are larger than females. The largest of the measured males was 9.4 m in length, of females 8.7 m. Measurements (as percent of animal length) of a young male with a body length of 518 cm and an adult female 716 cm long (Tomilin, 1962) are respectively: from tip of snout to commencement of dorsal fin 52.9 and 57.4; height of dorsal fin 5 and 5.3, length of dorsal fin 7.8 and 8.5; spread of caudal flukes between tips 27.1 and 29.1; length of flippers 10.7 and 8.5, and width of flippers 5.3 (female). The skull dimensions are (Tomilin, 1957) (in cm): condylobasal length of skull 145 (100%), zygomatic width 68 (46.9%), length of 655 887 Fig. 387. Skull of the northern bottlenose whale, Hyperoodon ampullatus (figure by N.N. Kondakov). rostrum 89 (61.4%), width of rostrum at base 35 (21.4%), length of lower jaw 135 (93.1%), and length of mandibular symphysis 25 (17.2%). (V.S.) Geographic Distribution North Atlantic Ocean, usually at depths exceeding 1,000 meters (Jons- gard, 1952). Geographic Range in the USSR Rare in our waters but sometimes encountered in the central part of the Barents Sea, along the Murman coast, and less often along the western coasts of Novaya Zemlya. Transgresses occasionally into the White Sea where instances of some animals being caught are known. Transgres- sions have also been observed in the southern part of the Baltic Sea (in winter). Geographic Range outside the USSR (Fig. 388) In the eastern part of the North Atlantic, these animals inhabit waters from the Cape Verde Islands in the south (15 to 18°N lat.), 656 657 888 NaS, “PAR 30 Fig. 388. Species range of the northern bottlenose whale, Hyperoodon ampullatus (V.A. Arsen’ev). along the entire European coast, and up to the Arctic. They cover the Mediterranean Sea, are encountered along the coasts of Holland, Great Britain, Shetland and Faroe Islands, and coasts of Norway up to Varangerfjord. They inhabit the waters of Iceland and Jan Mayen Island, are common in the Norwegian and North seas, and penetrate the waters of Spitsbergen in the zone of influence of the Gulf Stream (75-78° N lat.) toward the coast of Greenland. They transgress into the southern part of the Baltic Sea (Kiel and Lubeck bays, Rugen Island) but are rare on the Baltic coasts of Sweden and Norway. Known in the west from Rhode Island in the south up along the coast of North America (New York and Massachusetts bays) to Newfoundland, Hudson, Davis, etc. (Tomilin, 1957, 196210; Chapskii, 1963). (V.A.) Geographic Variation Not established. 10 References to the existence of this bottlenose whale in the Pacific Ocean, extensively cited in the literature, including the latest reviews (see, in particular, Hershkovitz, 1966: “Bering Sea and Japan’’), are erroneous and pertain to Вана’; beaked whale (Berardius bairdi, Tomilin, 1952, 1957; see latter). (У. H.) 889 Biology Population. The population of the northern bottlenose whale, compared with that of large whales, is very small. Hunting has never exceeded 500 bottlenose whales per year. Food. Cephalopods (Gonatus fabricu, Onychoteuthis sp., Sepia sp., and Loligo sp.) constitute the main food. Most of the stomachs inves- tigated revealed only the beaks of cephalopods, which numbered up to 10,000 in one stomach. Herring, cod, sea-cucumbers, and starfish were found as random food objects. The characteristic structure of the teeth and the ability of the northern bottlenose whale to dive to great depths confirm the view that these whales specialize in feeding on cephalopods (Tomilin, 1957). Daily activity and behavior. This whale is usually encountered in small groups, sometimes up to 20 animals. Single animals are rare. Groups most often comprise animals of different sexes and ages, including calves. Such groups resemble the harems of sperm whales since usually a single large male is seen in them. However, groups consisting exclusively of adult males have also been observed. At places of food concentration, groups of bottlenose whales sometimes gather into large herds of up to a hundred or more animals. Like the sperm whale, the northern bottlenose whale possesses the ability to descend to great depths and remain submerged for a long while. It has been suggested that the duration of its underwater residence may be as much as one hour. Herein lies some similarity of behavior during diving with the sperm whale. On surfacing after prolonged submergence, the northern bottlenose whale inhales/exhales quite a number of times and remains on the surface for a long time. It then undertakes another long dive. While resting, up to thirty brief intermediate submergences, one every 30 - 40 sec, producing small (50 -60 cm high), barely perceptible blows have been recorded for this whale. A display of the caudal flukes during diving has never been seen. The auditory faculty is thought to be very well developed in the northern bottlenose whale and it is sensitive to extraneous noises. The herd instinct is strong in these animals and they do not usually abandon an injured mate even when confronted with imminent danger themselves. Seasonal migrations and transgressions. The northern bottlenose whale performs regular seasonal migrations but the courses and periods of its spring and autumn migrations have hardly been studied. It is only known. that these whales inhabit the northern part of their range in the summer months and enter the southern part in winter. Spring migrations commence early since they are noticed off the coasts of Norway and 658 890 even Jan Mayen in March and April. The line of sharp increase in depth represents the boundary of distribution of these whales and they usually transgress no farther. Thus they never enter the fjords. Only one case is known; a small herd of bottlenose whales appeared in Oslo Fjord in September, 1939 and one whale beached (Jonsgard, 1952*). In June these whales are sighted along ice edges up to 77° N lat. and have even been encountered (sometimes in large numbers) among drifting ice. Off the coasts of Norway and in other regions of their hunting, the maximum number of bottlenose whales is caught in May and a slightly smaller number in June @stby, 1956, 1957, 1958, 1959). It has been assumed that this species penetrates farthest north in spring and later descends slightly southward to inhabit the waters of the Falkland and Shetland islands and east of Iceland. Most of the northern bottlenose whales are confined to the region where the Gulf Stream waters mix with Arctic currents, 1.е., the most productive regions. They transgress only into those cold waters which fall under the influence of the Gulf Stream. Autumn migrations have not been traced; the whales depart from the northern regions commencing probably September and extending up to late autumn. Wintering sites have not been established; it would appear that they overwinter in the southern part of their range. Many instances of beached bottlenose whales are known on the coasts of France, the FRG, the GDR, Holland, Denmark, Sweden, Nor- way, England, Ireland, Scotland, Faroe and Lofoten islands, and Ice- land. Beaching has also been recorded on the American coast in Rhode Island and Massachusetts states and in New York bay and the Gulf of St. Lawrence. Beached bottlenose whales have even been found on the coasts of Greenland and Spitsbergen. | Finds of bottlenose whales in the White and Baltic seas can be regarded as transgressions (Tomilin, 1957, 1962). Reproduction. The period of mating has not been established; gesta- tion extends for 12 and probably even for 15 months. Births occur in the spring, evidently in March-April or in April-May. The sites of mating and parturition have not been identified. Growth and development. Data on these aspects are extremely frag- mentary and few. The largest of the embryos studied was 305 and even 350 cm long. Apparently the newborn calf ranges from 200 to 250 cm in length, although the smallest of those measured was only 183 cm long. During lactation the calves double in length (duration of lactation is pre- sumably five to seven months). The length of calves fed on milk varied from 244 to 488 cm. By the third year these whales aitain a length of 6 т (Tomilin, 1957). Females become sexually mature at a body length 659 891 of 7 m. The smallest (of measured) pregnant females was 6.4 т long (Jonsgard, 1952*). Enemies, diseases, parasites, mortality, and competitors. The killer whale is regarded as an enemy of bottlenose whales but there are no direct observations of killer whale attacks on them. Diseases have not been studied; some cases of bone tumors have been recorded in museum skeletons. The whale louse Platicyamus thompsoni Gosse, mainly infesting the head, “beak,” and mouth corners, is a known ectoparasite of the northern bottlenose whale. Among the barnacles, Conchoderma cuviert is more commonly encountered and C. auritum rarely, in the teeth and light- colored sections of the skin. Penella crassicornis penetrates the skin. Seven species of helminths have been registered. The cestode Sirobil- icephalus triangularis Diesing, parasitizing the intestines, has been found in the northern bottlenose whale and three species of baleen whales from the waters of Portugal and Brazil; Diphyllobothrium delphini Bosk (larvae) has been detected in the adipose depots of these same whale species. The nematode Anisakis simplex Rudolphi, localizing in the gul- let, stomach, and intestines of the northern bottlenose whale, is very widely distributed among marine mammals. It has been found in many species of baleen and toothed whales and, among pinnipeds, in Steller’s sea lion caught in the North Sea, off the coasts of Kamchatka, and in the waters of Japan and New Zealand. Another nematode, Crassicauda benneti Spaul, has been cited for “bottlenose whale” without mention of the species. Of the two species of acanthocephalans, Bolbosoma tur- binella Diesing parasitizes in addition to the northern bottlenose whale, the intestine of six species of baleen whales from the waters of Iceland and Australia. Finally, Bolbosoma turbinella [sic.| Diesing, found in the intestine of the northern bottlenose whale, has been detected in five species of baleen whales. The mortality of bottlenose whales has not been established (Dela- mure, 1955; Tomilin, 1957). Field characteristics. Long, beak-shaped snout, steeply dropping high forehead, and the white color of old males are prominent features. One pair of teeth on the anterior tip of the lower jaw projects outward when the mouth is closed (sometimes a second pair projects slightly among very old males). The upper side of the body of young animals is black or grayish-black and of old animals light brown. When diving deep, this species flexes the body strongly but never displays the caudal flukes. Between two prolonged submergences, it remains for a long time on the water surface, producing 20-30 blows during this period. The blow 892 is low, bushy, 50-60 cm high, and bearly perceptible. The respiratory sound is short and interrupted with a metallic timbre. (V.A.) Economic Importance Commencing from the 1880s, regular hunting of the northern bottlenose whale was practiced in Norway but the output was low. Hunting inten- sified due to a drop in catch of right whales and several hundreds of bottlenose whales were caught in some years. However, by the begin- ning of the twentieth century, as a result of the rapid development of the hunting of large fin whales and reduction in the population of bot- tlenose whales, the catch of the latter decreased sharply. Hunting was initially carried out predominantly along the coasts of Greenland but later most of the whales were caught in Norwegian waters (Table 67). Commencing in 1929, modern specialized hunting of small whales (including the bottlenose whale) employed small motor boats equipped with small-caliber (50-60 mm) harpoon guns. Among the small whales caught, the Minke whale occupied first place followed by the bottlenose whale. In the 1930s, this hunting, especially of the Minke whale, was of great economic importance to the Norwegian fishermen. The maximum number of bottlenose whales was caught in the water body between Jan Mayen, Iceland, and Faroe Islands, and later west of Medvezhii Island and Spitsbergen and in the deepwater regions off the Norwegian coast. Table 67. Catch of northern bottlenose whales in Norway (international whaling statistics) Year Catch of whales Year Catch of whales 1938 70 1954 70 1939 45 1955 124 1940 8 1956 267 1941 21 1957 163 1942 9 1958 145 1943 34 1959 94 \ 1944 40 1960 193 1945 22 1961 87 1946 22 1962 321 1947 108 1963 267 1948 59 1964 307 1949 220 1965 692 1950 48 1966 340 1951 7 1967 264 1952 17 1968 384 1953 49 660 893 A single northern bottlenose whale yields about a ton of fat, on average (large animals up to two tons), in addition to 200 kg of sper- maceti. The fat is used only for commercial purposes. The spermaceti of the bottlenose whale is similar to that of the sperm whale in chemical composition. The hollow of the lower jaw of the bottlenose whale con- tains a small amount of oil with a typical chemical composition; it does not congeal in the cold and can be used for lubricating fine mechanisms. The raw meat causes disorders of the digestive tract but this adverse phenomenon disappears with cooking and the meat is suitable for feed- ing dogs and fur-bearing animals. It is not used for human consumption. This whale is not hunted in the Soviet Union. (V.A.) W re A) Cato 661 - 684 LITERATURE CITED’ The references listed below include only works cited in the text. Works cited under the synonymy of various species and groups have not been included. Russian Language Sources Akimushkin, 1.1. 1954а. Osnovnoi ob”ekt риашуа butylkonosa (Hyperoodon rostratus Muller) [Main food objects of the bottlenose whale (Hyperoodon rostratus Muller)]. Dokl. AN SSSR, vol. 96, no. 2. Akimushkin, ILI. 19546. Golovonogie mollyuski у pitanii kashalota (Cephalopods in the food of the sperm whale). Dokl. AN SSSR, vol. 96, no. 3. Akimushkin, 1.1. 1955. О kharaktere pitaniya kashalota (Feeding behavior of the sperm whale). Dokl. AN SSSR, vol. 101, no. 6. Akinfov, I. 1848a. Otpiska yakutskogo voevody Ivana Akinfova o tunguzakh, zhivushchikh bliz Okhoty reki, io morzhovom promysle po ostrovam i zalivam Okhotskogo morya (Yakut Commander Ivan Akinfov’s note on Tungus inhabiting near the Okhota River and walrus hunting off the islands and in the bays of the Sea of Okhotsk). Dopolneniya К aktam istoricheskim, sobrannye 1 izdannye Arkheograficheskoyu Komissieyu, vol. 3. Akinfov, I. 1848b. Rospis’ puti ot r. Okhoty do reki Motykhleya (Descrip- tion of the course from Okhota River to Motykleya River). Dopol- петуа К aktam istoricheskim, sobrannye 1 izdannye Arkheografich- eskoyu Komissieyu, vol. 3. Al’banov, У. 1917. Na учр, К Zemle Frantsa-losifa (South of Franz Josef Land). Annexure in Zapisok po gidrografii, vol. 41, no. 2. Aleev, У. 1913. Otchet о poezdke na morskoi zverinyi promysel у gorlo Belogo morya v 1911 g. (An account of the 1911 expedition for hunting marine animals in the neck of the White Sea). Mater. k. poznan. russk. ryb-va, vol. 2, no. 2. * Some references are incomplete in the Russian original. 896 Alekseev, Е.У. 1966. Vozrastnye izmeneniya nekotorykh morfologich- eskikh pokazatelei belomorskogo lysuna (Pagophoca groenlandica Erxleben) [Age-related changes of some morphological indices of harp seal (Pagophoca groenlandica Erxleben)]. In: Ekologo- morfologicheskie issledovaniya nekotorykh zhivotnykh. Kiev. Ambroz, А.1. 1931. Nekotorye nablyudeniya nad morskimi mlekopi- tayushchimi Vostochnogo Sakhalina (Some observations on marine mammals of eastern Sakhalin). Rybn. Khoz. DV, nos. 3 -4. Anbinder, E.M. 1969. Kariotipy baikal’skogo i kaspiiskogo tyulenei v svyazi $ problemoi ikh proiskhozhdeniya (Karyotypes of Baikal and Caspian seals in the context of the problem of their origin). IV Vses. Soveshch. po Izuch. Morsk. Mlekop. (Tez. Dokl.). Moscow. Anbinder, E.M. 1971. Khromosomnye nabory grenlandskogo, kaspi- iskogo i baikal’skogo tyulenei 1 nekotorye voprosy evolyutsii nastoy- ashchikh tyulenei (Chromosome numbers of Greenland, Caspian, and Baikal seals and some aspects of the evolution of true seals). Tsitologtya, vol. 3, no. 3. Andriyashev, A.P. 1954. Ryby severnykh morei SSSR (Fishes of the northern seas of the USSR). Opredelitel’ po Faune SSSR, 53, Zool. In-t AN SSSR. Moscow-Leningrad. Antipin, V.M. 1938. Morskoi zver’ na Novoi Zemle (Marine life on Novaya Zemlya). Sov. Arktika, no. 7. Antipin, V.M. 1938. Fauna pozvonochnykh severo-vostoka Novoi Zemle (Vertebrate fauna of northeastern Novaya Zemlya). Probl. Arktiki, no. 2. Anufriev, [.Р. 1918. О tyulen’em promysle (Seal hunting). Tr. Sev. Коти. dlya Pomoshchi Pomoram Rus. Sev. Arsen’ev, V.A. 1935a. Morskoi promysel mlekopitayushchikh v Chukot- skom i Vostochno-Sibirskom moryakh (Hunting for mammals in the Chukchi and East Siberian seas). Sov. Sever., nos. 3-4. Arsen’ev, У.А. 1935b. 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A, H. 10. 702 INDEX OF LATIN NAMES ОЕ ANIMALS’ Acrodelphidae 436, 441 Agrophiidae 435, 436 Alachtherium 27 Allodesmus 54 —kernensis 19 Anarnak 654 —groenlandicus 654 Archaeoceti 430 Arctocephalinae 54, 75 Arctocephalus 16, 18, 19, 21, 22, 54, 75, 79 —australis 19, 54 —californianus 81 —doriferus 54 —forsteri 54 —gazella 54 —monteriensis 56 —philippii 19, 54 —pusillus 18, 19, 54, 79 Argyrocetus 441 Artiodactyla 6, 8 Aulophyseter 593 Balaena ampullata 653, 654 —glacialis 425 —rostrata 654 Balaenoptera musculus 422 physalus 422, 422,** 424, 427 Basilosauridae 430 Beluga 562 Berardius 420, 627, 629 — агпоих 629 —bairdii 433, 437, 552, 629, 630, 630, 631, 632, 635, 656 —vegae 630 —vegana 630 Caenolestia 6 Caenolestoidea 6 Callocephalus 163 Callophoca 118 Callorhinus 14, 16, 18, 54, 74, 75, 79, 80 —alascanus 81 —curilensis 81 —obscura 166 —ursinus 14, 19, 21, 24, 52, 54, 55, 56, 68, 80, 81, 82, 84, 85, 86 - ~alascanus 87 - -californianus 87 --curilensis 86 - -cynocephalus 87 - -kracheninnikovi 86 --mimica 86 - -niger 86 - -ursinus 86 Callotaria 80 Canoidea 21 Carnivora 6, 8, 11, 13, 15, 21, 52 Caspiopusa Behningi 197 —Dierzawini 197 —kisielewitschi 197 Catodon 594 —australis 594 —macrocephalus 594 Cefus albicans 562 Cephalorhynchus 442 Ceratodon 587, 588 Cetacea 3, 7 “Reproduced from the Russian original. Page numbers of the Russian original appear in the left-hand margin in the text—General Editor. **Раре numbers in italics indicate illustration 990 Chiroptera 7 Clymene 444 —euphrosyne 444 Clymenia 444 Cystophinae 118 Cystophoca 392 Cystophora 14, 18, 118, 119, 392, 393 —borealis 392, 394 —cristata 19, 118, 119, 122, 122, 126, 393 395, 396, 397 Cystophorinae 118, 119, 129, 377, 390, 391, 392 Dasyuria 6 Delphinapterus 482, 562, 568, 587 —beluga 562, 563 —borealis 482 —dorofeevi 563 —freimani 563 —leucas 433, 438, 563, 564, 567, 568 - -leucas 568 - -dorofeevi 568 - -maris-albi 563, 568 Delphinavus 441 Delphinidae 435, 436, 441 Delphinoidea 436, 441 Delphinapteridae 561 Delphinus 442, 446, 452 —acutus 489 —aduncus 472 —albicans 562 —albirostris 487, 492 —algeriensis 455 —alope 451 —bairdii 455 —bredanensis 444 —butskopf 654 —capensis 455 —coeruleo-albus 446 —curvirostris 455 —deductor 522, 523 —delphis 418, 422, 432, 433, 440, 455, 456, 457, 458, 459 - -bairdii 460 - -delphis 460 - -ponticus 455, 460 —desmaresti 647 —dubius 450 —euphrosyne 444, 446 —frontalis 451 —frontatus 444 —gladiator 507 —globiceps 523 —grinda 523 —grisus 518, 519, 520, 521 —hyperoodon 654 —intermedius 535 —lateralis 446 —leucopleurus 487, 489 —malayanus 450 —leucas 562 —melas 523 —nesarnak 472 —orca 506, 507 —pectoralis 501 —peronii 482, 486 —phocaena 536, 558 —phocaenoides 557, 558 —rissoanus 518 —roseiventris 551 —rostratus 442, 444 —sowerbiensis 638 —sowerbensis 643 —styx 446 —truncatus 472 —tursio 472 Desmatophoca 54 Diaphorocetus 593 Didelphida 6 Diplodon 641 Diprotodontoidea 6 Disigatus 54 Dolichodon 641 Doridontidae 430 Electra 486 —electra 487 —obtusa 501 Eocetus 430 Erignathini 131 Erignathus 14, 16, 118, 119, 127, 129, 130, 131 —barbatus 19, 21, 24, 119, 120, 123, 127, 130, 131, 132, 134, 137, 141 - -albigena 142 - -barbatus 139, 141, 142, 160 - -lachtak 142 - -leporinus 142 - -lepechini 142 - -nauticus 139, 141, 142, 144, 156 - - рагзопзи 142 Eumetopias 18, 54, 56, 74, 75 —gillespii 78 —jubata 54 —jubatus 14, 52, 55, 57, 57, 59, 60, 66, 68, 107 Euphrosyne 444 Euphyseter 626 Eurhinodelphidae 436 Eurinodelphinidae 441 Feresa 442, 501, 534 —attenuata 439, 534, 535, 536 —occuleta 535 —uttenu 535 Gladiator 506 Globicephala 442, 522 —macrorhyncha 523 —melaena 438, 522, 523, 523, 524, 526, 527 - -deductor 525 - -globiceps 525 - -melaena 525 - -melas 525 - -scammoni 525 - -sibo 525 - -sieboldii 525 —scammoni 523 —sieboldii 523 Globicephalidae 441 Globicephalus 522 —chinensis 518 —scammoni 523 —sibo 523 Globiceps 522 Grampidae 441 Grampidelphis 518 Grampus 442, 489, 506, 518 —griseus 438, 507, 518 —orca 507 —stearnsii 518 Gryphoca 118, 341 Halichoerus 18, 118, 119, 127, 129, 340 —atlantica 341 —baltica 341 —griseus 340, 341 —grypus 19, 118, 120, 123, 123, 127, 341, 342, 344, 349, 351, 352, 394 991 —gryphus 354 - -atlantica 354 - -baltica 353 - -gris 354 - -grisseus 354 - - вгуриз 353, 354 - -halichoerus 354 - -macrorhynchus 353 - -pachyrhynchus 353 - -thienemanni 354 —macrorhynchus 341 —pachyrhynchus 341 Halicyon 163 —richardii 163, 232 Haliphilus 163 Heliophoca 375 —atlantica 375 Hemisyntrachelidae 436, 441 Hemisyntrachelus 472 Histriophoca 118, 122, 127, 163, 165, 166 167, 286, 329, 330, 331, 332 —fasciata 19 Histriophocina 286 Hoplocetinae 593 Hydrurga 16, 118, 374 —leptonys 115, 119 Hyperoodon 627, 629, 653 —ampullatus 437, 654, 655, 656 —butskops 653 —planifrons 654 Hyperoodon rostratus 422 Hyperoodontidae 436, 627 Indopacetus 654 Inia 422 —goeffrensis 413 Insectivora 6, 8 Kogia 593, 623, 624, 625 —breviceps 437, 627 —simus 437, 623, 627 Kogiidae 436 Kogiinae 593 Lagenodelphis 442 Lagenorhynchus 442, 487, 488, 501 —acutus 440, 489, 490, 491 —albirostris 439, 489, 493 —cruciger 489 —electra 439, 442, 487, 489, 501, 501 992 —obliquidens 439, 489, 495, 496, 498 —ognevi 495 —thicolea 489 Lagomorpha 6, 8 Lemuroidea 6 Leptonychotes 118, 374 —weddelli 19, 115, 119, 129 Leptophoca 19, 118 Leucopleurus 487 Lissodelphis 442, 482 —borealis 438, 482, 483, 484, 485, 486 —peronii 438, 482, 484, 487 Lobodon 16, 118, 374 —carcinophaga 19, 115, 119 Lobodoninae 118, 374 Lobodontini 374 Macrosselidea 6 Marsupialia 6 Mesoplodon 627, 638, 640, 641 —bidens 437, 639, 641, 643, 645, 646 —bowdoini 641 —carlhubbsi 641 —densirostris 639, 641 —europaeus 639 —ginkgodens 639, 641 —grayi 639 —hectori 641 —layardi 639, 641 —mirus 639, 654 —planifrons 654 —stejnegeri 437, 641, 642, 643, 644 Mesotaria 118 —ambigua 393 Miophoca 18, 19 —vetusta 341 Mirounga 14, 15, 18, 118, 119, 374, 392 —angustirostris 18, 119, 392 —leonina 9, 115, 116, 119, 397, 392 Monachinae 16, 118, 119, 129, 372, 373, 374, 377, 393 Monachini 374 Monachus 18, 118, 119, 131, 372, 374, 375, 376, 377 —mediterraneus 377 —monachus 19, 21, 119, 122, 126, 377, 378, 380, 381, 382 —schauinslandi 18, 21, 119, 377, 382 —tropicalis 18, 19, 21, 377, 382 Monodon 562, 587 —monoceros 438, 587, 588, 588, 589, 590 —narhval 588 Monodontidae 436, 441 Monotherium 19, 118 Mustelidae 118 Mutica 3 Mysticeti 6, 419, 430, 431 Narwalus 587 —vulgaris 587, 588 Neomeris 557 Neephoca 18, 19, 54, 75 —cinerea 54 —hookeri 54 Neophocaena 442, 557 —phocaenoides 438, 558, 558, 559, 560 Neotherium 54 Notocetus 627 Odobenidae 4, 12, 13, 20, 21, 22, 23, 24, 25 Odobenus 16, 27 —rosmarus 24, 25, 26, 27, 31 - -arcticus 33 - -cookii 33 - -divergens 33, 41, 44, 45 - -laptevi 28, 33 - -obesus 33 - -orientalis 33 - -rosmarus 33 Odontoceti 3, 6, 430, 431, 435 Ommatophoca 118, 372, 374 —rossi 119 Orca 506 —ater 507 —fusca 507 —gladiator 506 —intermedia 534 —rectipinna 507 Orcaelidae 441 Orcaella 442 Orcinus 442 —orca 317, 438, 507, 508, 509, 510, 511 —rectipinna 512 Otaria 19, 54, 56, 75 —byronia 19, 54 —californiana 74, 75 —gilliespii 74, 75 —japonica 75 —kracheninnikovi 81 —stelleri 58 Otariidae 4, 11, 12, 13, 14, 15, 16, 19—23, 24, 25, 27, 53, 54, 54, 118 Otarlinae 56, 75 Otarioidea 14, 21, 27, 52 Pagomy 163 Pagophila 163 Pagophilus 118, 163, 165, 167, 279, 283, 286, 287, 290, 302 —groenlandica 122 Pagophoca 118, 163 Paleophoca 118 Pappocetus 430 Pelagias 374 Pelagios 374 Pelagius 374 Pelagus 374 Pelagocyon 374 —monachus 374 Peramelia 6 Perissodactyla 8 Phalangeria 6 Peponocephala 442, 487, 489, 501 Роса’ 22, 118, 119 S127 22 23127, 129, 163, 166, 167, 232, 236, 244, 244, 245, 246, 278, 341 —albigena 131 —albiventer 377 —annelata 168 —baicalensis 220 —barbata 130, 131 —bicolor 377 —canina 232 —caspica 14, 19, 119, 121, 121, 124, 124, 129, 166, 167, 171, 197, 199, 200, 201 —chorisii 232 —communis 168 —crinita 377 —cristata 392, 394 —cucullata 394 —curilensis 246 —dorsata 278 —equestris 328 —fasciata 4, 119, 122, 122, 125, 163, 164, 165, 166, 167, 328, 329 — 332 —foetida 163, 167, 168 —groenlandica 19, 21, 118, 121, 125, 127, 129, 163, 164, 165, 166, 167, 278, 279, 283, 286, 287, 290 993 —ochotensis 232 - -groenlandica 291 - -oceanica 291, 302 —grypus 340, 341 —halitschensis 166 —hermanni 377 —hispida 14, 19, 21, 119, 121, 121, 124, 125, 129, 164, 166, 167, 168, 169, 170, 171, 173, 177, 202, 215, 236, 243 - -annelata 178 - -beaufortiana 179 - -birulai 168, 178 - -botnica 178 - -gischigensis 168, 178 - -hispida 178, 179 - -krascheninnikovi 168, 178, 181, 193 - -ladogensis 178, 195 - -nummularis 178 - -ochotensis 176, 178, 193 - -pomororum 168, 178 - -pygmaea 168, 178 - -rochmistrovi 168, 178 - -saimensis 178 - -Schreb 202, 232 - -soperi 179 —insularis 21, 232 —isidorei 394 —Jjubata 56, 58 —lachtak 131 —ladogensis 168 —largha 119, 126, 232, 237, 244 —leonina 58 —lepechini 131 —leporina 131 —leucogaster 377 —leucopla 394 —linnaei 232 —littorea 232 —mimica 81 —mitrata 394 —monachus 374, 377 —nautica 131 —nigra 81 —nummularis 232 —oceanica 278 —ochotensis 168, 232 - -kurilensis 232 —oronensis 220 —parsonsii 131 —petersi 232 994 —richardi pribilofensis 232 —rosmarus 27, 28 —saimensis 168 —semilunaris 278 —schreberi 168 —scopulicola 232 —sibirica 4, 119, 121, 121, 124, 124, 129, 164, 166, 167, 171, 202, 220, 220—222 —steinegeri 232, 237 —thienemannii 232 —variagata 232 —vitulina 19—22, 24, 119—121, 121, 124, ‘125, 126, 126, 163, 164, 166, 167, 171, 197, 220, 231, 232, 237, 238, 242, 244 - -botnica 167 - -canina 245 - -chorisii 246 - -concolor 246 - -insularis 246 - -kurilensis 119, 125, 237, 246, 247, 248, 250, 253 - -largha 125, 126, 232, 236, 244, 244, 245, 246, 250, 251, 252, 282 - -limnaei 245 - -littorea 245 - -macrodens 244, 246 - -nummularis 246 - -mellonae 249 - -ochotensis 244 - -pallasii 232, 244 - -petersi 244 - -richardi 125, 163, 246, 249 - -scopulicola 245 - -steinegeri 246 - -thienemanni 245 - -variegata 245 --vitulina 119, 123, 125, 126, 233, 235, 236, 244, 245 —vitulinoides 166 —ursina 79—81 Phocanella 118 —minor 166 —pumila 166 Phocarctos 18, 19 Phocidae 4, 12, 12, 13, 13, 14, 15, 18—23, 24, 25, 52, 117, 118—120, 127, 129, 300, 372, 390 Phocinae 16, 22, 114, 118, 119, 127—129, 136, 163, 341, 372, 377, 378, 390, 393 Phocoena 442, 470, 536, 537 —communis 538 —crassidens 501, 502 —dalli 549 —dioptrica 538 —phocoena 439, 538, 539 — 541 - -communis 542 - -phocoena 542 - -relicta 542 - -vomerina 542 —relicta 538 —sinus 538 —spinnipinnis 538 —vomerina 538 Phocoenidae 441 Phocoenoides 442, 548, 552 —dalli 439, 549, 549—551, 555 - -dalli 512, 549, 553, 634 - -truei 512, 549, 553 —truei 548, 549 Phocoidea 21, 27, 52 Physeter 539, 593, 594 —australasianus 594 —bidens 638, 643 —breviceps 623 —catodon 436, 594, 595, 596, 598 — 601, 614, 616 - -australasianus 603 - -australis 603 - -catodon 602 - -macrocephalus 602 —macrocephalus 594 —simus 626 —sulcatus 594 Physeteridae 435, 436, 593 Physeteroidea 436 Pinnipedia 3, 7, 17, 23 Pithanotaria 52 Platanista 434 —gangetica 413 Platanistidae 436 Platanistoidea 436 Platyphoca 118, 131 Pliopedia 54 Polyprotodontoidea 6 Pontolis 54 Pontophoca 118, 374 Pontoporia blainvillei 419 Pristiphoca 118, 375 Procidae 16 Prodelphinus 444 Prophoca 118 Prorosmarus 27 —alleni 27 Protocetidae 430 Protocetus 430 Pseudorca 442, 501 —crassidens 439, 502, 503, 504 —mediterranea 502 Pusa 118, 163, 166—168, 171, 178, 179, 193, 202, 232, 236, 286, 287, 291, 341 Rigoon 375 Rodentia 8 Rosmarus 27 Rostrifer nestorésmirnovi 630 Schizodelphis 441 Semantor macrutus 20 Semantoridae 21 Simae 6 Siren cynocephala 80 Sirenia 8 Sotalia 441, 442, 444 Sousa 444 Squalodon 627 Squalodonttdae 435, 436, 441, 627 Stenella 442, 444, 445, 450 —asthenops 446 —clymene 446 —caeruleoalba 440, 446, 447 —449, 512 - -euphrosyne 450 —coeruleoalbus 446, 450 —crotaphiscus 446 —dubia 440, 446, 450 —frontalis 440, 446, 450, 453 —graffmani 446 —longirostris 440, 446, 451, 453—455 —mialayana 446 —pernettyi 446 Stenidae 441, 444 Steno 445 —attenuatus 444, 450 —bredanensis 439, 442, 442, 443 Stenorhynchinae 118, 372 Stenorhynchus 372 995 Sudidae 554 Tarsioidea 6 Tasmacetus 627, 627 Trichechodon 27 Trichechus 27 —arcticus 28 —cookii 28 —divergens 28 —manatus 27 —obesus 28 —orientalis 28 Tridacna 35 True 19 Tupaioidea 6 Tursio 472, 482 Tursiops 442, 446, 470, 472 —abusalam 472 —aducans 472 —gephyneus 472 —gillii 472 —nuuanu 472 —parvimanus 472 —truncatus 422, 439, 472, 473 —476, 480 - - gillii 477 - -ponticus 472, 477 - -truncatus 477 Tylopoda 6 Ursidae 52 Ursus marinus 81 Vegae 630 Vermes 557 Xiphus 647 Zalambdodonta 6 Zalophus 13, 14, 16, 18, 54, 74, 76 —californianus 4, 19, 54, 55, 75, 76, 77 - -californianus 78 - -japonicus 78 - -wollebaecki 78 —cavirostris 473, 647, 647, 648 —650 Ziphiidae 419, 435, 436, 627, 628, 629 Ziphius 627, 647 —grebnitzkii 648 а и № а, м, ny i i om Peale I i 41 oe age es м м о : Te: ey у ие т УИ Ай и И и ий | Hart 1 —#- и в шв и роевомьиы | МИНИН MAMMALS OF THE ‘SOVIET UNION Volume I V.G. HEPTNER, A.A. NASIMOVICH, A.G. BANNIKOV This volume of Mammals of the Soviet Union is devoted to a description of artiodactyls and perissodactyls found in the Soviet Union. These animals are of great scientific and economic interest; information on them, especially in recent Soviet literature, is voluminous. In recent years world literature on this subject has likewise been considerably enriched with new data. The abundance of information on ungulates explains the size of the present volume. All the characteristics of groups have been described according to a common scheme; deviations occur in a few cases, however. These characteristics are stated briefly and pertain to the group as a whole; they are not exclusive to species of the Soviet fauna. All species are described according to a common plan, altered only in the case of some extinct forms. In devising the scheme for descriptions of species not only the convenience of the reader was kept in mind, but the hope that gaps in our knowledge would become self-evident and stimulate further research. MAMMALS OF THE SOVIET UNION Volume II, Part 2 This volume, is part of a three-volume monograph, and is a continuation, of Volume II, Part 1, which was devoted to sea cows and carnivores. It contains species descriptions of terrestrial carnivores and detailed informa- tion on their external morphology, skull, body measurements and other data, affinities with other species, geographic distribution in the historic past and today, geographic variation, practical significance, and biology. Descriptions are presented for orders, families, and genera, and keys given for their identification. The book is richly illustrated with photographs, sketches, and colored illustrations by the famous wild life painter, А.М. Komarov, and the zoologist-artist, N.N. Kondakov. The results of original scientific studies are published here for the . first time, providing readers a vast wealth of material heretofore unknown. This work is intended for teachers and students of faculties of biology and geography in universities, as well as pedagogic, agricultural, and forest institutes, similar organizations, workers engaged in game, fur, forest and fish trades; those interested in the conservation of nature, and all persons interested in zoology and nature study.